Lindsley-Drosophila Variation PDF

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Genetic Variations of
DAN L. LINDSLEY
and
E. H. GRELL
BIOLOGY DIVISION, OAK RIDGE NATIONAL LABORATORY

operated by
UNION CARBIDE CORPORATION
for the
UNITED STATES ATOMIC ENERGY COMMISSION
Carnegie Institution of Washington Publication No. 627

Library of Congress Catalog Card Number 68-15915
ISBN 0-87279-S38-8
First Printing-1968
Second Printing - 1 9 7 2
revised from
THE MUTANTS OF DROSOPHILA MELANOGASTER
Calvin B. Bridges and Katherine F. Brehme
Carnegie Institution of Washington Publication 552, 1944

We dedicate this book to Alexander Hollaender on his retirement as director
of the Biology Division of the Oak Ridge National Laboratory and in recognition
of his continuous encouragement and support-
EDITOR Isabel L. Norton
CONSULTING EDITOR Eileen J. Slaughter
INDEXING Elizabeth S. Von Halle
PROGRAMMING Charles E. Price
TYPING Neva P. Hair
COMPOSITION Ethel Cagle
DESIGN, LAYOUT AND Technical Publication Department
PRODUCTION Technical Information Division
Oak Ridge National Laboratory
Oak Ridge, Tennessee
Atomic Energy Commission
Division of Technical Information Extension
Oak Ridge, Tennessee
PREFACE
The last exhaustive compilation of genetic ance; and (7) wild-type stocks. All except the
variations of Drosophila melanogaster was "The first two groups have relatively few entries.
Mutants of Drosophila melanogaster" prepared by Several new categories of effects unknown or
Calvin B. Bridges and Katherine S. Brehme; it nearly so in 1942 are included here. (1) Pseudo-
was published in 1944 and was complete through alleles: the intensive investigations into pseu-
1942. This volume is a revision of their work; doallelism and complementation dating from the
it contains new information on variants already pioneering work of E. B. Lewis on Star and as-
described and descriptions of variants dis- troid (1945, Genetics 30: 137-66) have produced
covered since 1942; it is reasonably complete information on the genetic fine structure of many
through 1966. The new material was extracted loci. (2) Isozymes: a series of genetically con-
from the literature, from Drosophila Information trolled enzyme polymorphisms and deficiencies
Service, and from voluminous contributions of described mostly in the last decade; their dis-
unpublished material supplied by Drosophila covery was made possible by the development of
geneticists throughout the world. The revision gel electrophoresis. (3) Compound chromosomes:
describes genetic material currently available formed by the attachment of two doses of one
to Drosophila geneticists and extinct material chromosome arm to a single centromere; repre-
that may be encountered in earlier literature on sented by only the attached-^ chromosome in the
the subject. original edition, the various classes of compound
The work of Bridges and Brehme was divided chromosomes now available occupy an entire sub-
into two sections, one describing wild-type section. (4) Marked Y chromosomes: Y chromo-
stocks and the other describing the known depar- somes marked by the genes carried on small at-
tures from the normal genotype. Our revision is tached euchromatic segments derived from the X
divided into seven sections: (1) mutants, with or an autosome. (5) Reciprocal translocations
about 3000 entries; (2) chromosome aberrations, between the X and Y chromosomes. (6) Attached
more than 1500 entries; (3) special chromosomes, XY chromosomes: chromosomes with the portions
i.e., multiply marked chromosomes, balancers, of the X and Y chromosomes necessary for male
compound chromosomes, Y derivatives, and X-Y viability and fertility attached to a single centro-
combinations; (4) cytological markers; (5) depar- mere.
tures from diploidy; (6) nonchromosomal inherit-
Development of the system of nomenclature The conventions adopted fdr naming and sym-
designating genetic variations of Drosophila mel- bolizing different types of genetic changes are
anogaster has been rather haphazard; conse- discussed at the beginning of the different sec-
quently, the system is not a logical structure but tions of the book. Symbols of all genetic vari-
is replete with relics, redundancies, and incon- ants both normal and abnormal are always itali-
sistencies. Revision into a consistent scheme cized but their names in text are printed in roman.
is not practicable, creating as it would a chaotic We are grateful to our colleagues throughout
discontinuity in the literature. Even were such the world for their cooperation in making avail-
a revamping considered desirable, design of such able to us their unpublished observations and
a system is not obvious, since a change pro- in responding to our numerous queries. Special
posed to obviate one inconsistency would likely thanks are due Doctors E. B. Lewis, the late
create more conflicts than it alleviated. There- H. J. Muller, J. Schultz, and A. H. Sturtevant,
fore, with few exceptions, we have adhered to who served as an informal board of consultants.
the conventions established by Bridges and They have contributed a measure of success to
Brehme in the original volume. Some changes this effort but bear no responsibility for its
were made to correct glaring inconsistencies shortcomings.
and others to facilitate automatic handling of
Drosophila symbols.
D. L. L.
July 1967 E. H. G.
guide to
Genetic Variations of MUTATIONS
CHROMOSOME ABERRATIONS
Deficiencies
Duplications
Inversions
Rings
Translocations
Transpositions
SPECIAL CHROMOSOMES
Balancers
Compound Chromosomes
Multiply Marked Chromosomes
X—Y Combinations
Y Derivatives
CYTOLOGICAL MARKERS
DEPARTURES FROM DIPLOIDY
NONCHROMOSOMAL INHERITANCE
WILD TYPE STOCKS
Vn/T(Y;3)1 GENETIC AND CYTOLOGICAL MAPS

Mohr *nd Mossige, 1943, Norske Vldenskaps-Akad. 7: 1—51.
MUTATIONS
A variant exists when there are two or more sive and its name begins with a lower-case
alternative phenotypes. Usually, one is des- letter.
ignated as normal or wild-type because it is the For convenience, a symbol is assigned to each
phenotype characteristic of wild-type flies; the mutant type. This symbol is an abbreviation of
other or others are considered mutant alternatives the name that uniquely designates the mutant in
because they represent departures from normal. question; it combines brevity with information.
Distinction between normal and mutant may be- It usually begins with the same letter as the
come blurred or disappear where both alternatives name, is always italicized, and never contains
are characteristic of wild-type strains, as in iso- Greek letters, subscripts, or spaces; e.g., r for
alleles. The pair or group of alternatives de- rudimentary, R for Roughened, ro for rough, rs
fines a locus, which is given a name that sug- for rose, and ry for rosy. In designations of
gests the main diagnostic features of the mutant genotypes with several mutant genes, symbols
form of the locus without regard to secondary of genes on the same chromosome are separated
characters. The name is concise and is pref- by spaces (e.g., y w i B); symbols of genes on
erably a simple adjective such as black or a homologous chromosomes are separated by a
noun such as Bar. When the main character is slash bar (e.g., y H- / B); symbols of genes on
recognized in the heterozygote, the mutant is noohomologous chromosomes are separated by
considered dominant and its name begins with semicolons and spaces (e.g., bw; v; ey). Names
an upper-case letter; when it is recognized in are not italicized in text.
the homozygote, the mutant is considered reces-
GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER
MULTIPLE ALLELES. The alternatives or al- B+. The plus symbol alone implies the normal
leles at a particular genetic locus are designated (wild-type) allele in any context, such as y/+ or
by the same name and symbol and are differenti- y m //+. Absence of a particular locus may be
ated by distinguishing superscripts. At publication noted by use of a superscript minus sign with
of the original volume, heterozygotes for allelic the symbol; e.g., bb~.
mutants were thought always to show a mutant Loci controlling electrophoretic mobility of
phenotype and segregation of the mutants at mei- enzymes and other proteins require special con-
osis. Since that time, however, discoveries of ventions. Since electrophoretic variants can be
complementation and intra-allelic recombination scored equally well in heterozygotes and homo-
have revealed the widespread existence of com- zygotes, the genes controlling them are consid-
plex pseudoallelic series, with consequent com- ered dominant; e.g., Adh. Alleles specifying
plications in the definition of allelism. By the the variants are differentiated by arbitrary super-
criteria that they occupy virtually identical po- scripts; e.g., AdhF, AdhD, and those specifying
sitions on the genetic map and have similar phe- the absence of a particular enzyme or other pro-
notypic effects, mutants formerly thought to be tein by an appropriate superscript, such as n
at different loci may now be considered changes (negative), a zero, or a minus sign, rather than
in the same pseudoallelic complex; e.g., (a and by a lower-case symbol; e.g., Adhnl. The sole
spl, Iz and amx, and m and dy. The locus will exceptions to the rule that the genetic determi-
probably be defined ultimately as the unit of nants of electrophoretic protein variants be sym-
transcription or, more likely, of translation; but bolized as dominant genes are loci originally
in the current state of knowledge, we have been recognized by recessive phenotypes and so
content to adhere to the historic terminology. named; e.g., v and ry. For proteins with unde-
One school of thought names every recombina- termined activity, we use the symbol Pt- fol-
tionally separable element; another gives all lowed by an arbitrary designation specifying the
members of the same complex a single name with particular protein; e.g., Pt-1. Abbreviations for
arbitrary superscripts. The latter is our pref- the protein and the gene are frequently identical,
erence. and both are used in most discussions. The
The superscript notation designating alleles gene symbol may be differentiated from the pro-
has a number of different forms. A common de- tein symbol by having only its initial letter cap-
vice is an abbreviation that further characterizes italized and by being italicized, whereas the
the particular allele or that was used as the protein symbol is in roman capitals; e.g., ADH.
locus symbol before allelism was established. In several instances where two members of the
This practice is avoided because it has the dis- same allelic series were formerly given different
advantage of preempting useful symbols and locus names, both are here included under one
names from use as locus designations. Another name; e.g., Pm = bwvl. In other cases, we as-
unacceptable device is the use, as superscripts, sume allelism of mutants with similar phenotypes
of elements of the genotype in which the allele and genetic positions even though they have not
arose, since such a designation implies some- been tested for phenotypic interaction. In such
thing more than a trivial connection between instances, the basis for the assumption is usu-
allele and element. More-acceptable super- ally noted. Since the practice has not been con-
scripts for allelic designations are arbitrary sistent, some alleles may be described as dif-
numbers, experiment numbers, capitalized initials ferent genes. We make special effort to infer
of the finder or laboratory, or the date of dis- allelism for Minute loci and for factors causing
covery. The numeral 1 is the implied superscript production of melanotic pseudotumors. Bridges
of nonsuperscripted symbols. Whereas genes in and Brehme made few such inferences. Except
the same allelic series are designated by the in special cases, investigation of allelic inter-
same symbol but with different superscripts, mu- action of sex-linked recessive lethals is not
tants with similar phenotypes at different loci possible; consequently, they are often given dis-
are not given the same symbol and differentiated tinctive symbols where allelism may actually
only by a superscript; this was done extensively exist.
in the past; for example, for genes causing for- MIMICS. Mutants at different loci sometimes
mation of melanotic pseudotumors. have similar phenotypic effects. Such loci may
For a recessive allele of a preponderantly dom- be handled in several ways. The simplest is to
inant series or a dominant allele of a predomi- give each a distinctive name (e.g., vermilion,
nantly recessive series, the superscripts used cinnabar, scarlet, karmoisin, cardinal); this
are r and D, respectively; e.g., Hnr, Hnr2, and method has the effect of scattering such mimics
hwD. Finally, for the normal allele in a series, throughout the alphabetical listing. Or a com-
a superscript plus sign may be used, e.g., 6+ or mon symbol separated by a hyphen from a dis-
MUTATIONS
tinguishing symbol may be used (e.g., tu-la, e.g., 3.0 represents a more accurate location
tu-lb, tu-2 for genes controlling production of than 3. Map units are not computed to the
melanotic pseudotumors). Distinctive suffixes second decimal place. Accuracy of a map
are also useful (e.g., rough, roughoid, roughish, position determination is of course dependent
roughex; plexus, Plexate; dachs, dachsous; on the accuracy of the positions assigned to
maroon, maroonlike). The latter two schemes the reference markers; i.e., on the accuracy
frequently have the virtue of placing like pheno- of the map. We treat the map as a rough
types in sequence in an alphabetical listing. guide to the relative positions of loci but,
Some phenotypes result from mutation at many considered on a refined level, it may be inac-
loci in all chromosomes; these are given a com- curate with respect to both position and order
mon symbol followed by a parenthetical desig- of genes. (We have abandoned the ± used by
nation of the chromosome and then by a distin- Bridges and Brehme to indicate a particu-
guishing designation. Examples of this type of larly low level of reliability.)
mutant are the female steriles, the lethals, the origin: For induced mutants, the agent is
Minutes, and the male steriles [e.g., fs(2)B, given; mutants recovered from untreated par-
1(1)Jl, M(l)n, ms(2)E4, respectively]. Conven- ents or a wild population are listed as spon-
tions for formulating distinguishing symbols are taneous. Isoallelic variants found as major
similar to those for superscripts; use of infor- components of stocks or populations are
mation about the cytological or genetic location listed as naturally occurring alleles. The
is avoided to allow updating such information stock or chromosome in which each mutant
without changing the symbol. arose was listed by Bridges and Brehme;
MODIFIERS. The primary effect of some mutants unless the new mutant is virtually insepa-
is to cause another mutant to exhibit a more- rable from some element of the stock of ori-
extreme departure from normal (enhancer) or a gin (e.g., y31d in In(l)sc8), we omit mention
more nearly normal phenotype (suppressor). Such of the original stock.
mutants are symbolized e or E and sti or Su, fol- discoverer: Name, date of discovery.
lowed in parentheses by the gene modified. Des- synonym: Alternative symbol or name or both,
ignation of the particular allele modified appears mostly obsolete terminology.
as a superscript within the parentheses and references: Sources of the major descriptive
alleles of the modifier gene as superscripts out- material are listed, but bibliographic material
side the parentheses; e.g., su(lz34) and su(Hw)2. may also appear in some of the other cate-
Terms such as dilutor, exaggerator, inhibitor, gories.
intensifies and modifier were also formerly used, phenotype: The most important departures from
but we have usually attempted to classify such normal, which are usually those suggested by
genes as enhancers or suppressors. the name, are described first. Other infor-
FORMAT. Mutants with their descriptions mation about the phenotype follows, and
are now listed alphabetically according to sym- finally there may be data on viability and fer-
bol and cross-indexed according to name. Cur- tility. The last item in the phenotypic de-
rent terminology is listed in bold face. All scription is the rank, abbreviated RK. Mu-
cases of synonymy are also listed in body type tants are classified into three different ranks
with cross-references to current usage. Mutants according to their utility in experiments in
no longer existing in published stock lists or in which counts are made: RK1 mutants are
private stock lists that we have examined are easily scored; RK2 mutants are usable but
considered lost and are preceded in the list by less convenient; RK3 mutants have limited
a star. Each mutant is described according to usefulness. An RK3 mutant may be one with
the following format: good expression and viability but simply not
symbol: name convenient to use in counting experiments;
location: The location is indicated by the e.g., enzyme polymorphisms. The letter A
chromosome number, separated by a hyphen follows the rank of mutants associated with
from the genetic position on the chromosome. chromosome aberrations.
Two levels of accuracy of the genetic loca- other information: This category contains
tion are indicated, those carried to tenths of miscellaneous information that does not fit
a unit being the more accurately determined; into one of the other categories.
a: arc phenotype: Nearly normal; distinguished by its

location: 2-99.2. interaction with certain svr alleles (see Gold-
discoverer: Bridges, 12e24. schmidt, 1945, table 74). RK3.
references: Bridges and Morgan, 1919, Carnegie cytology: Salivary chromosomes normal (Kodani).
Inst. Wash. Publ. No. 278: 202 (fig.). £>a2
Morgan, Bridges, and Sturtevant, 1925, Bibliog. origin: Spontaneous.
Genet. 2: 212 (fig.). discoverer: Goldschmidt.
Bridges, 1937, Cytologia (Tokyo), Fujii Jub. Vol. synonym: bran2.
2: 745-55. references: 1945, Univ. Calif. (Berkeley) Publ.
phenotype: Wings broader, bent downward in slight, Zool. 49: 363-73, 388-89.
even arc, and edges drawn down to diamond shape. phenotype: Wings somewhat more angular than aba.
Sometimes in stock, wings are bent upward instead Interactions with other genes shown in table 74 of
of downward. Crossveins closer together. RK2. Goldschmidt (1945). RK2A.
cytology: Placed between 57F11 and 58E4 on the cytology: Associated with Df(2R)aba2 = Df(2R)58D5-
basis of its inclusion within Df(2R)M-l = 6;58D7-8 (Goldschmidt, 1945).
Df(2R)57Fll-58Al;58F8-59Al but not Dp(2;3)P =
Dp(2;3)58E3-4;60D14-E2;96B5-Cl (Bridges, 1937). origin: Spontaneous.
Likely in band 58D6 or 7 based on Df(2R)ab"2 - discoverer: Goldschmidt.
Df(2R)58D5-6;58D7-8. synonym: bran^.
references: 1945, Univ. Calif. (Berkeley) Publ.
Zool. 49: 386-89.
phenotype: Normal; distinguished by its interaction
with certain svr alleles (see Goldschmidt, 1945,
table 74). RK3.
cytology: Salivary chromosomes normal (Kodani).
origin: Spontaneous. Probably a derivative of aba3.

discoverer: Goldschmidt.
synonym: bran4.
Zool. 49: 389-90, 490.
phenotype: Like aba; distinguished by its interaction
a: arc with certain svr alleles (see Goldschmidt, 1945,
From Bridges and Morgan, 1919, Carnegie Inst. Wash. table 74). RK2.
Publ. No. 278: 148. cytology: Salivary chromosomes normal (Kodani).
aba: arc-broad angular *abadb: arc-broad angular dumpy blistered

origin: Spontaneous. origin: Spontaneous.
discoverer: Goldschmidt, 1934. discoverer: Goldschmidt.
synonym: Always referred to as bran: broad angular synonym: brandb.
by Goldschmidt, but shown by him to be an allele references: 1945, Univ. Calif. (Berkeley) Publ.
of arc. Zool. 49: 370-71, 388-89.
references: 1945, Univ. Calif. (Berkeley) Publ. phenotype: Like aba; distinguished by its interaction
Zool. 49: 351-56, 388-89, 519. with certain svr alleles (see Goldschmidt, 1945,
phenotype: Wings broader and shorter than wild type, table 74). RK2.
blunt at the tip. Frequently shows upturned pos- cytology: Salivary chromosomes normal (Kodani).
terior scutellar bristles. In combination with *abadp; arc-broad angular dumpy
svrPoi, produces soft blistered wing. Other inter- origin: Spontaneous.
actions described by Goldschmidt, 1945, table 74. discoverer: Goldschmidt.
Wing grows in pupal stage to full length and then synonym: brandP.
retracts, possibly with histolysis [Goldschmidt, references: 1945, Univ. Calif. (Berkeley) Publ.
1935, Z. Induktive Abstammungs- Vererbungslehre Zool. 49: 373-86, 388-89.
69: 38-131 (fig.)J. RK2. phenotype: Normal; distinguished by its interaction
cytology: Salivary chromosomes normal (Kodani). with certain svr alleles (see Goldschmidt, 1945,
other information: Claimed to recur repeatedly in table 74). RK3.
certain lines (Goldschmidt, 1945). cytology: Salivary chromosomes normal (Kodani).
other information: Claimed by Goldschmidt to recur
origin: Spontaneous, repeatedly in certain lines.
discoverer: Goldschmidt. *ai>or; arc-broad angular rudimentary
synonym: bran*. origin: Spontaneous derivative of abadP.
references: 1945, Univ. Calif. (Berkeley) Publ. discoverer: Goldschmidt.
Zool. 49: 364-69, 388-89. synonym: branr.
MUTATIONS
references: 1945, Univ. Calif. (Berkeley) Publ. cytology: Salivary chromosomes normal (Hannah-
Zool. 49: 378-79, 388-89. Alava).
phenotype: Wing broad, round and dp-like. Interacts *aM60: arc of Meyer
with certain svr alleles (see Goldschmidt, 1945, origin: X ray induced.
table 74). RK2. discoverer: Meyer, 60f.
references: 1963, DIS 37: 50.
* o Bo ; arc-Broad angular Dominant phenotype: Homozygous lethal. RK3A.
origin: Spontaneous derivative of aba. cytology: Associated with In(2LR)aM6°; breakpoints
discoverer: Goldschmidt. unknown.
synonym: Bran. *A: Abnormal abdomen
references: 1945, Univ. Calif. (Berkeley) Publ. location: 1-4.5.
Zool. 49: 360-63, 388-89. discoverer: Morgan, l l g .
phenotype: aBa/+ resembles aba/aba. aBa/aba synonym: Abnormal.
shows Minute bristles. RK2. references: 1915, Am. Naturalist 49: 384-429 (fig.)-
* o 8 o C : arc-Broad Angular in Canton Morgan and Bridges, 1916, Carnegie Inst. Wash.
origin: Spontaneous. Publ. No. 237: 27 (fig.).
discoverer: Goldschmidt. phenotype: Tergites and sternites raggedly incom-
synonym: Branâ. plete, exposing a thin crinkled cuticle; bristles
references: 1947, J. Exptl. Zool. 104: 197-221. and hairs on abdomen correspondingly eliminated.
phenotype: aBaC/+ is normal; aBaC/aba ig broad Highly variable, wild phenotype in old dry cultures.
angular, but overlaps wild type. aBa^ is dominant A/+ less extreme than A/A and A male; homozy-
in its interaction with certain svr alleles. RK3.. gous female fully viable and fertile. RK2 in well-
cytology: Salivary chromosomes normal (Hannah- fed cultures.
Alava). other information: Lost by reversion to wild type.
*aBapl: arc-Broad angular in silver-pointed A: see bwA
origin: Spontaneous.
discoverer: Goldschmidt, 1947. location: 1- (just to the right of w; judged to be
synonym: BranP0*47"1. allelic to .4).
references: 1947, J. Exptl. Zool. 104: 197-221. origin: Spontaneous.
phenotype: a13^1/+ resembles aba/ab*. RK2. discoverer: Hillman, 53g.
cytology: Salivary chromosomes normal (Hannah- references: 1953, DIS 27: 56.
Alava). Hillman and Barbour, 1963, Proc. Intern. Congr.
*aBop2 Genet., 11th. Vol. 1: 170.
origin: Spontaneous. phenotype: Highly variable; ranging from extreme
discoverer: Goldschmidt, 1947. expression in young cultures to normal in old cul-
synonym: BranPoi47-2. tures. Expression in AS36/AS36 females > A536/Y
references: 1947, J. Exptl. Zool. 104: 197-221. males > AS3&/+ females. Expression varies from
phenotype: Phenotype normal in combination with loss of tergites 2—8 in extreme cases to loss of
aba and +; homozygous lethal. Dominant in inter- lateral part of tergite in one or more segments.
actions with certain svr alleles. RK2 as lethal. RK2A in young cultures.
cytology: Salivary chromosomes normal (Hannah- cytology: Associated with rearrangement of 2-5
Ala va). bands in 3C-D.
* o BoX ; arc-Broad angular from X irradiation a-3: seea(3)26
origin: X ray induced. *A-p; Abnormal abdomen-polygenic
discoverer: Goldschmidt. location: Polygenic.
synonym: Branx. discoverer: Sobels, 49j.
references: 1945, Univ. Calif. (Berkeley) Publ. references: 1950, DIS 24: 62.
Zool. 49: 521-22. 1951, DIS 25: 75-76.
phenotype: Resembles abo2 and abadb, but more or 1952, Genetica 26: 117-279 (fig.).
less dominant. Homozygote never obtained. Inter- 1952, Trans. Intern. Congr. Entomol., 9th. Vol. 1:
actions listed by Goldschmidt (1945, table 153). 225-30.
RK2. synonym: AA; Aay: Asymmetric.
phenotype: Incomplete mediodorsal fusion and one-
origin: Spontaneous. sided reduction of tergites. When more than one
discoverer: Goldschmidt. tergite is abnormal, spiral segmentation types are
synonym: Brany P* bl. most frequent. Expression strongly dependent on
references: 1947, J. Exptl. Zool. 104: 197-221. environment. Penetrance and expressivity corre-
phenotype: Homozygotes usually lethal; rare survi- lated (Bezem and Sobels, 1953, Konixikl. Ned.
vors have short, folded wings and are sterile. Akad. Wetenschap., Proc. Ser. C 56: 48-61). In
aBay/+ is broad-angular, with occasional truncate- strains selected for penetrance of A-p, mediodorsal
like wings. In combination with svrPoi resembles fusion or asymmetrical reduction of head and
rudimentary and blistered. RK2 as lethal. thorax also occur. RK3.
8 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER
*a(1)48: abnormal abdomen in chromosome 1 *A(2)51

location: 1- (not located). location: 2- (not located).
discoverer: Zimmerman, 1948. discoverer: Zimmerman, 1951.
references: 1952, DIS 26: 69. references: 1952, DIS 26: 69.
1954, Z. Induktive Abstammungs- Vererbungslehre 1954, Z. Induktive Abstrammungs- Vererbungslehre
86: 327-72 (fig.). 86: 327-72 (fig.).
phenotype: Has no phenotype of its own but in- phenotype: None. Enhances a(2)48 and a(3)48. RK3.
creases the incidence of abdominal malformations o(3)26
in a(2)48 and a(3)48 and in progeny of such flies. location: 3-27 (to the right of se).
Viability and fertility good. RK3. origin: Spontaneous.
*a(1)50 discoverer: H. A. and N. W. Timofeeff-Ressovsky.
location: 1- (not located). synonym: a-3.
origin: Spontaneous. references: 1927, Arch. Entwicklungsmech. Organ.
discoverer: Zimmerman, 1950. 109: 70-109.
references: 1952, DIS 26: 69. Schaffer, 1935, Z. Induktive Abstammungs-
1954, Z. Induktive Abstammungs- Vererbungslehre Vererbungslehre 68: 336—60 (fig.).
86: 327-72 (fig.). phenotype: Irregular reduction of abdominal tergites,
phenotype: Irregularities in abdomen most frequently sternites, pigmentation, and bristles; more marked
involving the anterior segments. Penetrance 1 per- in females and increased by crowding and dry food
cent. Enhances maternal effects of a(2)48 and (Braun, 1938, Am. Naturalist 72: 189-92).
a(3)48. Viability and fertility good. RK3. Schaffer's data (1935) suggest irregular dominance
*o(l)Sl in heterozygote, overlapping of wild type in homo-
location: 1- (not located). zygote, and genetic modifiers. RK3.
origin: Spontaneous. *a(3)48
discoverer: Zimmerman, 1951. location: 3- (not located).
references: 1952, DIS 26: 69. origin: Spontaneous.
1954, Z. Induktive Abstammungs-Vererbungslehre discoverer: Zimmerman, 1948.
86: 327-72 (fig.). references: 1952, DIS 26: 69.
phenotype: Shows maternal effect only, with 2 per- 1954, Z. Induktive Abstammungs- Vererbungslehre
cent of progeny affected. Abnormalities more 86: 327-72 (fig.).
anterior than those of a(2)4S and a(l)50. Viability phenotype: Only a maternal effect affecting 2.5 per-
and fertility good. RK3. cent of progeny. Irregularities most frequently
*o(2)48 involve posterior segments of abdomen. Viability
location: 2- (not located), and fertility good. RK3.
origin: Spontaneous. A34: see bwV6
discoverer: Zimmerman, 1948. aa: an arista
references: 1952, DIS 26: 69. location: 3-0.
1954, Z. Induktive Abstammungs- Vererbungslehre discoverer: Bridges, 23dlO.
86: 327-72 (fig.), synonym: al~b: aristaless-b.
phenotype: Abdominal irregularities most frequently references: Morgan, Bridges, and Sturtevant, 1925,
involve anterior segments. Penetrance 7 percent. Bibliog. Genet. 2: 218.
Also shows maternal effect. Viability and fertility phenotype: Aristae bare or tufted. Wings somewhat
good. RK3. broader than wild type. Expression variable, over-
*o(2)50 laps wild type often in female and sometimes in
location: 2- (not located), male. RK3.
origin: Spontaneous. cytology: Placed between 61E2 and 62A6, on basis
discoverer: Zimmerman, 1950. of its inclusion in Df(3L)D = Dt(3L)61E2-
references: 1952, DIS 26: 69. Fl;62A4-6 from T(Y;2;3)D.
1954, Z. Inductive Abstammungs- Vererbungslehre
86: 327-72 (fig.), Aa: Altered abdomen
phenotype: None. Six percent progeny affected, i.e., location: 1- (not located).
only maternal effect. RK3. origin: X ray induced in the In(l)dl-49, y w t compo-
nent of C(1)DX, y i of Muller.
location: 2- (not located), discoverer: Cicak, 56f.
origin: Spontaneous, references: Cicak and Oster, 1957, DIS 31: 80.
discoverer: Zimmerman, 1951. phenotype: Heavy deposition of melanin in tergites
references: 1952, DIS 26: 69. of females and males. Males sterile, therefore
1954, Z. Induktive Abstammungs- Vererbungslehre homozygous females not produced. RK2A.
86: 327-72 (fig.). cytology: Probably associated with a rearrangement
phenotype: Penetrance 50 percent. Also shows in addition to ln(l)dl-49.
maternal effect. RK3. A A: see A-p
MUTATIONS
ab: abrupt cytology: Placed in region between 59E2 and 60B10

location: 2-44.0. by Bridges (1937) on basis of its being to the right
origin: Spontaneous. of In(2R)bv/Vr>»1 = In(2R)41B2-Cl;59E2-4 and to
discoverer: Bridges, 16jl6. the left of Df(2R)Px = Df(2R)60B8-10;60Dl-2.
references: Morgan, Bridges, and Sturtevant, 1925,
Bibliog. Genet. 2: 218 (fig.).
phenotype: Vein L5 usually stops after posterior
crossvein. Scutellar bristles usually fewer. Wing
effect probably âcts during contraction period
(Waddington). Overlaps wild type. RK2.
ab: abrupt
Edith M. Wallace, unpublished.
discoverer: Bridges, 23gl6.
synonym: pt: parted.
references: Morgan, Bridges, and Sturtevant, 1925, abb: abbreviated
Bibliog. Genet. 2: 232. From Bridges and Brehme, 1944, Carnegie Inst. Wash.
phenotype: Vein L5 does not reach margin. Scutellar Publ. No. 552: 11.
bristles always fewer than wild type. Hairs parted
down midline of thorax and abdomen. Supra-alar abd: abdominal
bristles sometimes absent. Coxae tend to be location: 3-27 (close to the right of se).
thickened. Males sterile and have rotated gen- origin: Spontaneous.
italia. ab/ab2 resembles ab/ab but has a stronger discoverer: Gottschewski, 1935.
bristle effect. RK2. phenotype: Abdominal bands broken and etched.
Overlaps wild type in test crosses but not in
origin: Spontaneous in In(2L)Cy + In(2R)Cy. homozygous stock. Slightly semidominant. More
discoverer: Edmonds on, 51g. extreme at 19°C. abd/a(3)26 shows slight abd
references: 1952, DIS 26: 60. effect. RK3.
phenotype: A strong allele like ab2. RK2A. abdomen rotatum: see ar
abdominal: see abd
*abt-60h: abrupt-lethal
origin: Spontaneous. *abe: abnormal eye
discoverer: Hall, 60h. location: 1-1.2.
references: 1960, Meyer, DIS 34: 52. origin: Induced by 2-chloroethyl methanesulfonate
phenotype: Homozygote rarely survives. abl"60ll/ab2 (CB. 1506).
has shortened vein L5, but no scutellar bristles discoverer: Fahmy, 1955.
missing, and there is no part down midline of references: 1959, DIS 33: 82-83.
thorax and abdomen. ab1'60fl/ab2 males are phenotype: Eyes rough; either small or deformed.
fertile. RK2. Wings slightly atypical; inner margin frequently
abb: abbreviated removed by large irregular incisions; L4 frequently
location: 2-105.5. stops well short of the wing edge. Flies slightly
discoverer: Bridges, 28d6. smaller than normal. Males about 50 percent as
references: 1937, Cytologia (Tokyo), Fujii Jub. viable as wild type and fertile. Females highly
Vol. 2: 745-55. infertile. RK2.
phenotype: Bristles smaller; especially posterior abero: see abr
scutellars. Developmental time slightly longer Abnormal: see A
than normal. Viability only slightly reduced. abnormal abdomen: see a( )
Classification difficult, especially in early eclo- Abnormal abdomen: see A
sions; improves with age of culture. Enhanced by abnormal eye: see a be
shrunken (2-2.3), making classification easy. abnormal tergites: see abt
RK3; RK2 with shr. abnormal wings: see abw
abr: abero synonym: Called ac11 by Serebrovsky.

location: 2-83. references: 1918, Genetics 3: 133—72.
origin: Spontaneous. Dubinin, 1930, Zh. Eksperim. Biol. 6: 325—46.
discoverer: Bridges, 33blO. 1933, J. Genet. 27: 443-64.
phenotype: Abdominal banding etched and irregular. phenotype: Posterior dorsocentral bristles missing,
Wing margins irregular. Eyes rough. Bristles and anteriors rarely; hairs usually fewer near posterior
hairs sparse and disarranged. abr/+ sometimes dorsocentrals; intraocellar hairs invariably fewer,
lacks anterior dorsocentrals. Viability usually typically absent. Eyes partly devoid of hairs.
poor. RK3. ac/ac or ac/+ partially suppresses h (Sturtevant).
other information: Not allelic to fr or nw. Hw/ac = Hw/+ (Sturtevant). RK1.
abrupt: see ab cytology: Placed in region 1A5-8 on basis of its in-
Abruptex: see Ax clusion in the XD3P element of T(l;3)sc260-20 _
k
abt: abnormal tergites T(l;3)lA8-B1^61Al-2 and in Dp(l;f)sc2 6 0-2 7 =
location: 1-45.6. Dp(l;t)lA8-Bl;l9F, but not being lost from
origin: Induced by 2-chloroethyl methanesulfonate Df(l)260-5 =Dt(l)lA4-5 (Sutton, 1943, Genetics 28:
(CB. 1506). 210-17).
discoverer: Fahmy, 1955. 2
references: 1959, DIS 33: 83. origin: X ray induced simultaneously with s c 3 .
phenotype: Abdomen affected to various degrees, discoverer: Dubinin, 1928.
from extreme deformation of tergites to slight references: 1929, Biol. Zentr. 49: 328-39.
abnormalities in distribution of pigment and hairs. Serebrovsky and Dubinin, 1930, J. Heredity 21:
Eyes also deformed to various degrees from gross 259-65.
alterations in shape to slight derangement of om- phenotype: Since ac2 and sc3 are for practical
matidia. Wings vary from alterations in size, out- purposes inseparable by crossing over, the effect
line, and venation to small incisions of the inner of ac2 alone cannot be assessed. The double
margin. Most-extreme effects not always posi- mutant removes all bristles except scutellars and
tively correlated, and all flies show several postdorsocentrals. ac2/ac2 or ac2/+ suppresses h
atypical characters. Males viable, fertility severly (Sturtevant). Viability of males low; females
reduced. RK3. nearly inviable. RK2.
abw: abnormal wings cytology: Salivary chromosomes normal (Schultz).
location: 1-60. : see ac3
origin: X ray induced.
discoverer: Halfer, 1963. origin: X ray induced.
phenotype: Wing size reduced; wings upturned; L5 discoverer: Dubinin, 1929.
and crossveins absent. Plexus of veins between synonym: Called ac2 by Dubinin, the earlier ac2
L3 and L4. RK1. with sc3 having been omitted from the series.
sci0. sc-'-' (Sturtevant and Schultz, 1931, Proc.
Natl. Acad. Sci. U.S. 17: 265-70).
references: 1930, Zh. Eksperim. Biol. 6: 300—24.
1932, J. Genet. 25: 163-81.
1933, J. Genet. 27: 443-64.
phenotype: Posterior and usually anterior dorsocen-
trals lacking; other bristles wild type. Hairs re-
moved from areas across rear and front edges of
thorax, through mid-dorsal area, and between
ocelli. RK2A.
cytology: Inseparable from In(l)ac3 =
In(l)lB2-3;lB14-Cl (Muller, Prokofyeva, and
Raffel, 1935, Nature 135: 253-55).
other information: Judged to be an allele of ac but
not sc; it is mutant in combination with ac but not
with sc alleles except for scs which may also
show ac variegation and sc3, now lost, which is
thought to be a sc ac double mutant (Sturtevant).
ac3: see ac*
ac: achaete *QC4
From Bridges and Brehme, 1944, Carnegie Inst. Wash. origin: X ray induced in X chromosome carrying sc.
P u b l . No. 552: 12. discoverer: Dubinin, 1929.
a c : achaete synonym: Called ac3 by Dubinin.
location: 1-0.0. references: 1930, Zh. Eksperim. Biol. 6: 300—24.
origin: Spontaneous in X chromosome carrying y. 1932, J. Genet. 26: 37-58.
discoverer: We ins tern, 16b3. 1933, J. Genet. 27: 443-64.
MUTATIONS 11
phenotype: Anterior and posterior dorsocentrals toward the anode more rapidly than those specified
removed; also thoracic hairs. A change also ap- by AdhF. As with AdhF and Adhs, three isozymes
parently induced in expression of sc; called sc13. are specified by AdhD. RK3.
The sc component also removes scutellars and cytology: Placed in region between 34E5 and 33D1,
often ocellars, postverticals, and first and second on the basis of its inclusion in Df(2L)64j =
orbital bristles. Viability low. RK2. Dt(2L)34E5-Fl;35C3-Dl (E. H. Grell).
*ac260-28
origin: X ray induced simultaneously with y260-28m AdhF; Alcohol dehydrogenase-Fast
discoverer: Sutton, 39126. origin: Naturally occurring allele.
references: 1943, Genetics 28: 210—17. discoverer: Johnson and Denniston, 1964.
cytology: Salivary chromosomes appear normal. references: 1964, Nature 204: 906—7.
Ac: see CuA Grell, Jacobson, and Murphy, 1965, Science 149:
occ: acclinal wing 80-82.
location: 1-54.5. Ursprung and Leone, 1965, J. Exptl. Zool. 160:
origin: Induced by triethylenemelamine (CB. 1246). 147-54.
discoverer: Fahmy, 1952. phenotype: Specifies isozymes of alcohol dehydro-
references: 1958, DIS 32: 67. genase that migrate Lin the system of Grell,
phenotype: Wings upheld but slope backward at 45° Jacobson, and Murphy (1965)] toward the anode
angle from abdomen. Viability and fertility good in more rapidly than the isozyme's specified by Adhs.
both sexes. RK1. Homozygote contains three electrophoretically sep-
other information: One allele each induced by arable isozymes. The one moving most rapidly to-
CB. 3007 and by CB. 3026. ward the anode is often not detected in zymograms
achaete; see ac of single adults but is nearly always detectable in
Acpfi-7A: Acid phosphatase-1-A zymograms of single larvae. The faster isozymes
location: 3-101.4. more reliably detected with use of sec-butanol than
origin: Naturally occurring allele. with ethanol as a substrate. AdhF/Adhs heterozy-
discoverer: Maclntyre, 1964. gote contains the parental isozymes plus three
references: 1966, DIS 41: 61. hybrid isozymes. Hybrid enzymes also formed in
1966, Genetics 53: 461-74. melanogaster X simulans hybrids. RK3.
phenotype: Acph-1A/Acph-1A produces acid phos-
phatase-1 enzyme that migrates slowly in starch
gel electrophoresis. Enzyme found in larva, pupa, 70 -i
and adult. RK3. - 9
Acpfi-JB 60-
origin: Naturally occurring allele.
discoverer: Maclntyre, 1964.
50- -8
1966, Genetics 53: 461-74.
phenotype: Acph-1B/Acph-1B produces more rapidly g 40 H -6
migrating acid phosphatase-1 than Acph-1A/ 5 -7
Acph-1A. Acph-1A/Acph-1B produces the two pa- o
cr ^///i/7 * ^
rental enzymes as well as a hybrid enzyme of in- ^ 30 -

termediate mobility. A hybrid enzyme is also -4
2
formed in simulans X melanogaster hybrids. RK3.
20-
ad: arco id
location: 2-60.7.
origin: Spontaneous. 10 -
discoverer: Curry, 38a2.
ORIGIN
phenotype: Wings arched, broad, and somewhat
shortened; crossveins close; scutellar groove
shallow. Legs may be slightly shorter than wild Alcohol dehydrogenase alleles
type. RK3. Adh*; b = AdhF/AdhF; c = Adh
AdhD: Alcohol dehydrogenase-D From Grell, Jacobsen, and Murphy, 2 July 1965,
location: 2-50.1 (one-tenth the distance from el be- Science 149: 80-82.
tween el and rd).
origin: Ethyl rnethanesulfonate-induced derivative of : Alcohol dehydrogenase-negative
Adh^ of Samarkand. origin: Ethyl methanesulfonate-induced derivative of
discoverer: E. H. Grell, 65k8. Adhs of Canton-S.
phenotype: Specifies isozymes of alcohol dehydro- discoverer: E. H. Grell, 66elO.
genase that migrate [in the system of Grell, phenotype: Homozygote shows no alcohol dehydro-
Jacobson, and Murphy (1965, Science 149: 80-82)] genase activity. Sensitive to alcohol, showing
evidence of intoxication within 1 hr of being adpfs: adipose-female sterile

placed on substrate containing 15 percent ethanol; origin: Spontaneous.
death invariably follows within 24 hr. Heterozygote discoverer: Counce, 1956.
with allele producing active enzyme shows evi- synonym: fs(2)adp: female sterile(2) adipose.
dence of formation of a hybrid enzyme with one ac- references: Doane, 1959, Genetics 44: 506.
tive and one mutant polypeptide subunit. RK3. 1960, J. Exptl. Zool. 145: 1-42 (fig.).
Adh»2 1961, J. Exptl. Zool. 146: 275-98.
origin: Ethyl methanesulfonate-induced derivative of phenotype: Adult fat body phenotype like adp60;
Adhs. corpus allatum hypertrophies in mated females to
discoverer: E. H. Grell, 66elO. same degree as in adp6°. Females completely
phenotype: Like Adhnl except no evidence of hybrid sterile; sterility autonomous. Eggs laid by homo-
enzyme in heterozygote with active allele. RK3. zygotes show meiotic or mitotic abnormalities, or
both, never develop beyond early cleavage stages.
origin: Ethyl methanesulfonate-induced derivative of Males 78 percent fertile. Heterozygotes fertile but
Adhs. females become sterile with age. Viability gener-
discoverer: E. H. Grell, 66f. ally good, but longevity reduced; horaozygotes
phenotype: Like Adhn2. RK3. with selective advantage under starvation; hetero-
Adh"4 zygotes superior under desiccation. Average water
origin: Ethyl methanesulfonate-induced derivative of content of well-fed adults reduced; percentage of
AdhD. lipids, as a function of dry body weight, almost
discoverer: E. H. Grell, 66g. double that of wild type. Iodine numbers show
phenotype: Like Adh^2. RK3. greater degree of saturation of mutant lipid
extracts than of wild type. RK3.
origin: Ethyl methanesulfonate-induced derivative of *ae: aeroplane
AdhD. location: 2-55.8.
discoverer: E. H. Grell, 66g. origin: Spontaneous.
phenotype: Small amount of alcohol dehydrogenase discoverer: Mohr, 26k24.
activity in homozygote but ethanol sensitive. Elec- references: Quelprud, 1931, Hereditas 15: 97-119
trophoretic migration of enzyme like that of AdhD. (fig.),
Heterozygote with fully active allele has hybrid phenotype: Wings spread, balancers drooping.
enzyme, presumably with one active and one Adhn$ Overlaps wild type. RK3.
subunit. RK3. *Ae; Aechna
Adhs: Alcohol dehydrogenase-Slow location: 3- (rearrangement).
origin: Naturally occurring allele. origin: X ray induced.
discoverer: Johnson and Denniston, 1964. discoverer: Belgovsky, 45al4.
references: 1954, Nature 204: 906—7. references: 1946, DIS 20: 63.
Grell, Jacobson, and Murphy, 1965, Science 149: phenotype: Wings spread at right angles to body
80-82. axis. Homozygous lethal. RK1A.
Ursprung and Leone, 1965, J. Exptl. Zool. 160: other information: Reduced crossing over in the th-e
147-54. region suggests presence of pericentric inversion.
phenotype: Specifies isozymes of alcohol dehydro- aeroplane: see ae
genase that [with the methods of Greil, Jacobson, *agl: angle winglike
and Murphy (1965)] migrate more slowly to the location: 1- (not located).
anode than those specified by AdhF. There are origin: Recovered among descendants of flies
also three isozymes in Adhs homozygote. RK3. treated with natural gas.
discoverer: Mickey, 49c7.
adp60: adipose synonym: Originally called angle wing, but this
location: 2-83.4. name preoccupied by ang.
origin: Spontaneous. references: 1950, DIS 24: 60.
discoverer: Doane, 1960. phenotype: Wing bent upward in middle. Overlaps
references: 1961, DIS 35: 78. wild type. RK3.
1963, DIS 38: 32. ah aristaless
phenotype: Adult fat body hypertrophies as cells location: 2-0.01 (to the right of net).
become distorted by enormous oil globules. Ab- origin: Spontaneous.
normal fat bodies visible through body wall of discoverer: Bridges, 17k7.
6-day-old and older adults when submerged in 95 references: Morgan, Bridges, and Sturtevant, 1925,
percent alcohol and then water. Adult corpus Bibliog. Genet. 2: 213 (fig.).
allatum of mated females hypertrophies. Females Stern and Bridges, 1926, Genetics 11: 510 (fig.).
fertile but egg hatchability reduced to 45—90 per- phenotype: Aristae strongly reduced. Postscutellars
cent, depending on residual genome; adult emer- widely separated, erect but strongly divergent.
gence lowered to 33—85 percent. Males viable and Scutellum shortened; sternopleurals irregular in
fertile. RK3. size, position, and number; wings slightly bowed
MUTATIONS 13
downward, narrowed, pointed; first longitudinal cytology: Associated with In(2LR)alM60, inferred
vein raised and thickened. RK1. from suppression of crossing over in most of 2L
cytology: Placed in 21C1-2 doublet on the basis of and some of 2R.
its inclusion in Df(2L)al = Df(2L)21B8-Cl;21C8-Dl *o/v: arista!ess-variegated
but not in Df(2L)S5 = Df(2L)21C2-3;22A3-4 (Lewis, origin: X ray induced.
1945, Genetics 30: 137-166). discoverer: E. B. Lewis, 1940.
0/2 references: 1945, Genetics 30: 137—66.
origin: Spontaneous. phenotype: alv/al variegated for al. Homozygous
discoverer: Stern, 26a. lethal. RK2A.
references: Stern and Bridges, 1926, Genetics cytology: Associated with In(2LR)alv =In(2LR)21B-
11: 511. C1.-41.
phenotype: Slightly less extreme than at, but vi- al-b: see aa
ability poorer. RK2. ala: see dyaJa
ala parvae: see dyata
* 0 /3 alarless: see air
origin: Spontaneous. Alcohol dehydrogenase: see Adh
discoverer: Bridges, 33g2.
phenotype: Aristae absent or much reduced. Thorax A/in: Aliesterase-negative
has wide bare area or groove down midline with location: 3- (not located).
divergent hairs and bristles; sternopleurals ab- origin: Spontaneous.
sent. Wings have weakened L2 vein and delta at discoverer: Ogita.
tip of L3. Female sterile. Viability about 10 per- synonym: ali: aliesteraseless.
cent of wild type. RK3. references: 1961, Botyu-Kagaku 26: 93—97.
1962, DIS 36: 103.
phenotype: Homozygotes practically unable to
hydrolyze methyl butyrate, whereas wild type
shows high activity; Alin/+ exhibits intermediate
activity. Homozygotes shown by Beckman and
Johnson to lack a normally present esterase that
migrates slowly on starch gel (their band F). RK3.
Alkaline phosphatase: see Aph
aliesteraseless: see Alin
almond: see Didr
almondex: see amx
almondex-55: see lzK
*o/o; alopecia
location: 1-38.3.
origin: Induced by 2-chloroethyl methanesulfonate
(CB. 1506).
discoverer: Fahmy, 1956.
al3: arrstaless-3 phenotype: Abdominal hairs much reduced in
Edith M. Wallace, unpublished. number; pigmentation frequently lighter and patchy.
Effect very pronounced in females reared at 25°C,
ah but overlaps wild type in both sexes when reared
origin: Spontaneous in In(2LR)bwvl. at a low temperature. Viability and fertility good
discoverer: Bridges, 33127. in males but reduced in females. RK3.
references: 1935, DIS 3: 5. alpha: see tyr-1
phenotype: Slight allele of al in some or all stocks *alr: alarless
of bwyl. RK2A. location: 3- (not located).
*a/3<5 origin: Spontaneous,
origin: X ray induced, discoverer: Steinberg, 40b.
discoverer: Glass, 36c. references: 1940, DIS 13: 51.
references: 1939, DIS 12: 47. phenotype: Outer postalar bristle always missing;
phenotype: Like al. RK1. posterior supra-alar missing in about 80 percent of
the flies. Anterior scutellars, humerals, and noto-
*a/AU0: art stales s of Meyer pleurals frequently duplicated. Never overlaps.
origin: X ray induced, Viability and fertility excellent. RK3.
discoverer: Meyer, 60f. Altered abdomen: see Aa
references: 1963, DIS 37: 50. Alu: Alula
phenotype: Homozygous lethal. May be variegated location: 2-54.9 (Muller places Alu to the left of
position effect. RK3A. pr and spindle attachment).
origin: Spontaneous. *amy: amethyst

discoverer: Bridges, 38al2. location: 2- (not located).
references: Curry, 1939, DIS 12: 45. discoverer: Bridges.
phenotype: Heterozygote has alula fused to main references: Morgan, Bridges, and Sturtevant, 1925,
wing; wings often bent, broader. May overlap wild Bibliog. Genet. 2: 218.
type, but intensified by cold and by heterozygous phenotype: Transparent light-purplish eye color.
ds, with buckling effect increased. Homozygote at RK3.
19°C shows extreme buckling owing to rotation of Amy1: Amylase-1
wing and alula. Homozygote viable and resembles location: 2-77.3 (Doane, 1963, DIS 38: 32).
heterozygote. RK2. origin: Naturally occurring allele.
*AluS6c discoverer: Kikkawa, 1957.
origin: Spontaneous (arose with ltS6c). synonym: Amy+.
discoverer: Meyer, 56c. references: Kikkawa and Abe, 1960, Annotationes
references: 1956, DIS 30: 77. Zool. Japon. 33: 14—23.
phenotype: Similar to Alu. RK2. Kikkawa, I960, Japan. J. Genet. 35: 382—87.
Kikkawa and Ogita, 1962, Japan.J. Genet. 37:
*o/w: arclike wing 394-95.
location: 2- (near 6). Kikkawa, 1963, DIS 37: 94.
discoverer: Sturtevant, 1948. 1964, Japan J. Genet. 39: 401-11 (fig.).
references: 1948, DIS 22: 55. phenotype: Specifies amylase isozyme system
phenotype: Wings evenly bent downward at tips. having, in agar gel electrophoresis, one major and
Overlaps wild type. RK2. one minor component. The major isozyme moves
am: see Didr rapidly toward the anode and occupies position 1.
The minor component migrates immediately behind
*amb: amber it to position 2. Heterozygotes of Amy alleles con-
location: 1-6.8. tain isozymes of both parents. RK3.
origin: Induced by triethylenemelamine (CB. 1246).
phenotype: Pale yellow body color; bristles very
thin and short; hairs less affected. Eyes slightly
brighter red. Males sterile. Viability 10—50 per-
cent wild type. RK2.
other information: One allele each induced by
CB, 1246, CB. 3007, CB. 1506, CB. 1414. Two
alleles induced by CB. 3034.
amb2
origin: Induced by L~p-NN-di(2-chloroethyl)amino-
phenylalanine (CB. 3025). 6
discoverer: Fahmy, 1954. 7
phenotype: Pale yellow body color; bristles slender Origin!—)
1.2 1.3 1.4 1.6 2.6 3.6 46
and only slightly shortened. Male viability and
fertility good; females viable but sterile. RK2. Amylase Alleles
amethyst: see amy Electrophoretic patterns of homozygotes for the various
alleles of Amy. Amy*'2 is on acrylamide gel and the
amx: almondex rest are on agar gel.
location: 1-27.7 [to the left of lz (Green and Green,
1956, Z. Induktive Abstammungs- Vererbungslehre -2
87: 708-21)]. origin: Naturally occurring allele.
origin: X ray induced. discoverer: Doane, 64e6.
discoverer: Ball, 32k20. phenotype: Major Ot-amylase isozymes occupy posi-
phenotype: Eyes slightly reduced, narrower below. tions 1 and 2 in acrylamide gel disc electropho-
Trident pattern stronger than in lz. Homozygous resis (corresponding to positions on agar gel). A
females highly infertile; all progeny that do occur minor component is present at position 0. (In acryl-
are daughters. Infertility does not resemble that of amide gels, minor components migrate more rapidly
lz females, since amx has no effect on the gen- to the anode than major isozymes. In agar gels
italia [Anderson, 1945, Genetics 30: 280-96 (fig.)]. they migrate less rapidly). The total a-amylase
Iz-'mmx is wild type. RK2. activity is quite high, being intermediate between
cytology: Located in 8D (region 8D4 through 8E2) Amy4-6 and Amy1-3. RK3.
by Green mnd Green (1956).
origin: Naturally occurring allele.
MUTATIONS 15
discoverer: Kikkawa. Amys: see Amy2-6

references: 1964, Japan. J. Genet. 39: 401—11 (fig.). Amywh: see Amy1-4
phenotype: Specifies major amylase isozymes that an: an con
occupy positions 1 and 3 after agar gel electropho- location: 2-44 (34-54).
resis and minor components at positions 2 and 7. discoverer: Bridges, 30e3.
RK3. phenotype: Wings and legs somewhat short. Over-
laps wild type. RK3.
origin: Naturally occurring allele. an2
discoverer: Kikkawa. discoverer: Bridges 30c25.
synonym: Amywh; Amy4. phenotype: Wings broad and short. Legs short and
references: 1963, DIS 37: 94. gnarled. Bristles on abdomen straggly; sclerites
1964, Japan.J. Genet. 39: 401-11 (fig.). etched. Eyes small and roughish. an2'/an is like
Doane, 1966, DIS 41: 93. an2. Overlaps wild type. RK3.
phenotype: Specifies major amylase isozymes that anarista: see era
occupy positions 1 and 4 and a minor component at oncon: see an
position 5 after agar gel electrophoresis. The ang: angle wing
isozyme at position 1 was originally considered to location: 2-10.5.
be minor, but Doane considers it major; most pro- origin: Spontaneous.
nounced in young flies. RK3. discoverer: Mittler and Goldberg, 48il6.
Amyl-6 references: Mittler, 1950, DIS 24: 61.
origin: Naturally occurring allele. phenotype: Wings held up from dorsal surfaces and
discoverer: Kikkawa. extended outward 15—90° from the mid-dorsal line.
references: 1964, Japan. J. Genet. 39: 401-11 (fig.). Longitudinal dorsal median muscles 5 and 6 fused
phenotype: Specifies major amylase isozymes that (Goldberg, 1954, Ph.D. Thesis, 111. Inst.
occupy positions 1 and 6 and minor components at Technol.). No increase in expressivity with
positions 2 and 7 after agar gel electrophoresis. temperature. Does not overlap wild type. RK2.
RK3. ang: see ano
Amy* & angle wing: see ang
origin: Naturally occurring allele. angle wing: see agl
discoverer: Kikkawa. angle winglike: see agl
synonym: AmyB. *ano: anomogenitals
references: Kikkawa and Abe, 1960, Annotationes location: 1-35.7.
Zool. Japon. 33: 14-23. origin: Induced by triethylenemelamine (CB. 1246).
Kikkawa, 1960, Japan.J. Genet. 35: 382-87. discoverer: Fahmy, 1952.
Kikkawa and Ogita, 1962, Japan. J. Genet. 37: synonym: Originally symbolized ang, but this
394-95. symbol was preoccupied.
Kikkawa, 1963, DIS 37: 94. references: 1958, DIS 32: 67.
Kikkawa, 1964, Japan. J. Genet. 39: 401-11 (fig.). phenotype: Many bristles on head and thorax either
phenotype: Specifies major amylase isozymes that reduced in size or absent. Thoracic and abdominal
occupy positions 2 and 6 and minor components at hairs appreciably fewer. External male genitalia
positions 3 and 7 after agar gel electrophoresis. invariably abnormal, sometimes completely absent.
RK3. Melanized exudate frequently present in furrow
Amy* 6 between mesonotum and scutellum near anterior
origin: Naturally occurring allele. scutellar bristles. Males sterile; viability less
discoverer: Kikkawa. than 10 percent wild type. RK3.
references: 1964, Japan. J. Genet. 39: 401-11 (fig.).
phenotype: Specifies major amylase isozymes that ant: antennaless
occupy positions 3 and 6 and minor components location: 2- (not located).
that occupy positions 4 and 7 after agar gel elec- origin: Spontaneous.
trophoresis. RK3. discoverer: Gordon, 1936.
Amy4: see Amy1-'* references: 1941, DIS 14: 39.
1941, Proc. Intern. Congr. Genet., 7th. p. 131.
origin: Naturally occurring allele. Gordon and Sang, 1941, Proc. Roy. Soc. (London),
discoverer: Kikkawa. Ser. B 130: 151-84 (fig.).
synonym: Amyad. Vogt, 1947, Biol. Zentr. 66: 388-95 (fig.).
references: 1963, DIS 37: 94. phenotype: Antennae missing on one or both sides.
1964, Japan.J. Genet. 39: 401-11 (fig.), Expression affected by residual genotype, nutri-
phenotype: Specifies major amylase isozymes that tional environment, and temperature. Time of ac-
occupy positions 4 and 6 and minor components at tion about 70 hours after hatching LBegg and Sang,
positions 5 and 7. RK3. 1945, J. Exptl. Biol. 21: 1-4 (fig.)]. Used in ex-
Amy*: see Amy1 periments to locate chemoreceptors IBegg and
Amyad: see Amy4-6 Hogben, 1946, Proc. Roy. Soc. (London), Ser.
B 133: 1—19j and in studies of mating behavior phenotype: Arista tends to be transformed into
(Begg and Packman, 1951, Nature 168: 953). RK3. tarsus; third antennal segment hypertrophied and
deformed. Ocelli reduced in size and number. Ce-
Antp49: Antennapedia phalic capsule deformed. Head bristles reduced in
location: 3- (to the left of p; probably to the right number. Wings held at 45° angle from midline. Ex-
of st; determined for Antp50 by Hannah). pression variable. Homozygous lethal. RK3A.
origin: X ray induced. cytology: Associated with In(3R)AntpL>c =
discoverer: Piternick, 1949. In(3R)84A5-6; 92A5-6.
synonym: Antp4703.
phenotype: Antennae transformed into second legs
plus some differentiation toward first legs (Hannah-
Alava). Lethal in combination with AntpYu,
AntpB, Antp50, and Sex. Possibly lethal with Pc
but semilethal with P c 2 (Hannah-Alava). Quite
variable. Homozygous lethal. RK3A.
cytology: Probably in region 84A (or 83F), based
on Lewis's analysis of AntpB and AntpYu. Ant49
associated with small cytological abnormality of
undetermined nature in 83EF-84AB (Hannah-Alava).
AntpSO
discoverer: Piternick, 1950.
synonym: Antp4715.
phenotype: Variable transformation of entire an- AntpLC: Antennapedia of Le Calvez
tenna into a leg may occur, but effect is often From Le Calvez, 1948, Bull. Biol. France Belg. 82:
limited to slight elongation of third antennal seg- 97-113.
ment. In compounds with Pc and Pc2, the rather
well developed antennal second legs show some AntpR: Antennapedia of Rappaport
transformation into first legs (Hannah-Alava). origin: X ray induced.
Homozygous lethal and lethal in combination with discoverer: Rappaport, 1963.
AntpB, AntpYa, Antp49, and Sex (Hannah-Alava). synonym: s s ^ : spineless-Aristapedia Dominant.
RK3A. references: Falk, 1964, DIS 39: 60.
Antp4?0*: see Antp49 phenotype: Segments added to antennae, usually
Antp4?15: see Antpso distal to aristae. Claw occasionally at end of
antenna. Asymmetry pronounced. Rarely an an-
AntpB: Antennapedia of Bacon tennalike organ on sternopleura. Variable expres-
origin: X ray induced. sion, but expressivity 100 percent in combination
discoverer: Bacon, 50g. with D. Homozygous lethal; lethal in combination
references: Lewis, 1956, DIS 30: 76. withAntpB (Von Halle). RK3A.
phenotype: Antenna partially leg-like, but less ex- cytology: Associated with In(3R)AntpR =
treme than AntpYu and may overlap wild type. En- ln(3R)83F;86C (Ben-Zeev).
hanced by Pc (and Pc2"). AntpB ssa/+ ssa has AntpYu: Antennapedia of Yu
virtually complete antennal leg including coxa, origin: X ray induced.
tibia, femur, and tarsus, but no sex comb in the discoverer: Yu, 1948.
male (B. Holloway). The antennal second leg with reference: 1949, Ph.D. Thesis Calif. Inst. Technol.
some transformation into a first leg completely Lewis, 1956, DIS 30: 76.
developed only in Pc (or Pc2) ssa/AntpB s s a phenotype: Antenna transformed into second leg
compounds. Lethal with AntpYu, Antp49, Antp50, plus some differentiation toward a first leg but
and Sex (Hannah and Stromnaes, 1955, DIS 29: with recognizable arista usually present; not like
121-23 and Hannah-Alava). RK3A. ssa, in which main effect is that the arista be-
cytology: Associated with In(3R)AntpB = comes tarsus-like. Strongly enhanced by Pc and
ln(3R)84A;85E, but apparently mutant and inversion Pc2. Pc +/+ AntpYtl has a pair of excellent an-
are separable (Hannah-Alava). tennal legs complete with tarsae and with sex
combs in the male. Lethal with AntpB, Antp49,
*AntpL-C; Antennapedia of Le Calvez Antp50, and Sex (Hannah and Stromnaes, 1955,
origin: Neutron induced. DIS 29: 121—23; Hannah-Alava). RK3A.
discoverer: Le Calvez. cytology: Associated with T(2;3)AntpYu =
synonym: Ar: Aristapedia; SS^T. T(2;3)22B;83E-F + T(2;3)38E;98A.
references: 1948, Compt. Rend. 226: 123-24. *ap: apterous
1948, Bull. Biol. France Belg. 82: 97-113 (fig.). locotion: 2-55.2.
1948, Arch. Anat. Microscop. Morphol. Exptl. 37: origin: Spontaneous.
50-72. discoverer: E. M. Wallace, 13h.
MUTATIONS 17
references: Bridges and Morgan, 1919, Carnegie references: Meyer, Edmondson, Byers, and
Inst. Wash. Publ. No. 278: 236 (fig.). Erickson, 1950, DIS 24: 59.
Metz, 1914, Am. Naturalist 48: 675-92. phenotype: Compared with ap4 and very similar.
phenotype: Wings and halteres reduced to traces. Almost lethal. RK2.
Bristles eliminated from area around wing base ap
(including posterior notopleurals, anterior, and origin: Spontaneous.
posterior supra-alars, anterior postalars); posterior discoverer: Faulhaber.
scutellars erect when present, but missing in first references: 1963, DIS 37: 48.
counts; dorsocentrals smaller and fewer; hairs on phenotype: Wings vary from clublike to straplike;
thorax sparse and irregular. Sutural furrow re- seldom exceed 30 percent of normal length; lack
duced; thorax disproportionately small. Flies veins and specific bristles. Halteres 25—50 percent
small, pale, weak, and very short-lived. Viability normal length. Postalar, scutellar, and dorsocentral
about 70 percent that of wild type, but erratic. bristles missing. Hind legs sometimes deformed.
Both sexes sterile. RK2. Few ap6/ap6 or ap4/ap6 females survive past 5
cytology: Placed in salivary region 41B-C (Schultz). days; those that do are slightly fertile, the re-
mainder are not (Butterworth and King), ap6/
origin: Spontaneous. M(2)S24 more extreme than ap6/ap6 but some fe-
discoverer: Bridges, 16j20. males fertile and thoracic chaetotaxy more nearly
synonym: ap-c. normal. RK2.
references: 1919, J. Exptl. Zool. 28: 370.
Bridges and Morgan, 1919, Carnegie Inst. Wash. origin: Spontaneous.
Publ. No. 278: 237. discoverer: Ritterhoff, 49j.
Medvedev and Bridges, 1935, Tr. Inst. Genet. references: Glass, 1951, DIS 25: 76-77.
Akad. NaukSSSR 10: 199-209. phenotype: Appears to be somewhat less extreme
phenotype: Like ap but less viable. RK2. than ap4, with which it was compared. Wings and
halteres reduced to vestiges. Bristles and hairs on
origin: Spontaneous. sides absent, including posterior notopleurals,
discoverer: Morgan, 23a. anterior and posterior supra-alars, and anterior
synonym: no-wings; later, ap~c. postalars, but dorsocentrals not reduced in size
references: 1929, Carnegie Inst. Wash. Publ. No. and number, and one or a pair of pre-anterior dor-
399: 183. socentrals may be present. Posterior scutellars
phenotype: Like ap. RK2. not erect when present. Sutural furrow normal;
thorax of normal size. No adults live longer than 3
discoverer: Medvedev, 32al5. days; larval adipose cells persist in adult, and
references: Medvedev and Bridges, 1935, Tr. Inst. adult adipose tissue fails to develop. ap49i/
Genet. Akad. Nauk SSSR 10: 199-209. M(2)S24 adult sterile, short lived, and has abnormal
Beatty, 1949, Proc. Roy. Soc. Edinburgh, adipose tissue and short wing rudiments (Butter-
B 63: 249-70 (fig.). worth and King). Both sexes sterile. RK2.
King and Sang, 1958, DIS 32: 133. other information: Interacts with apXa but not
synonym: ap-d. tested in combination with ap4 or apblt.
phenotype: Wings mostly less than 10 percent normal apSSf
length and lacking veins and specific hairs. Hal- origin: Spontaneous.
teres less than 25 percent normal length, and fre- discoverer: Thompson, 56f.
quently absent. Scutellar and dorsocentral bristles references: Burdick, 1956, DIS 30: 69.
sometimes missing (Butterworth and King). Adults phenotype: Wings club-shaped, 10—30 percent normal
become paralyzed with age and die within 4 days. length, and lack veins and certain types of hairs.
Larval adipose cells persist in imago, and adult Scutellar and dorsocentral bristles missing (Butter-
adipose tissue fails to develop. Female sterile worth and King). Rear and middle legs occasionally
with underdeveloped ovaries; nurse cell nuclei be- twisted, more frequently in female than in male.
come pycnotic after stage 7, and yolk formation is Both sexes fertile when homozygous and in combi-
never initiated (King and Burnett, 1957, Growth 21: nation with other ap alleles. ap56l/M(2)S24 have
263—80). ap4 ovaries develop normally when trans- normal complement of dorsocentral and scutellar
planted into a normal host (King and Bodenstein, bristles (Butterworth and King). RK2.
1965, Z. Naturforsch. 20b: 292-97). Male sterile,
but testes appear normal with motile sperm (King apbh; apterous-blot
and Sang, 1958). ap4/M(2)S24 adult has nearly origin: Spontaneous.
normal complement of bristles but otherwise re- discoverer: Groscurth, 31bl.
sembles ap4 homozygote (Butterworth and King). synonym: bit.
RK2. phenotype: Wings blistered, inflated, often dark be-
ap- cause of dried blood. In extreme cases, a small
origin: Ultraviolet induced. mirror image wing forms by partial twinning of
discoverer: Byers, 49f. wing in third posterior wing cell. According to
18 GENETIC VARIATIONS OF DROSOPHILA MELANOCASTER
Waddington (1939, Proc. Natl. Acad. Sci. U.S. 25: cytology: Shown by Sturtevant (1934, DIS 2: 19) to
299—307), the fundamental effect is partial be associated with T(2;3)apXa = T(2;3)41F;
twinning of wing blade, which leads to difficulties 89E8-F1 which is superimposed on In(2R)Cy and
in clearance of heraolymph after inflated stage. In(3R)P (Morgan, Bridges, and Schultz, 1936,
Much overlapping with wild type. RK3. Carnegie Inst. Wash. Year Book 35: 294; Lewis,
1951, DIS 25: 109).
*apbtt2
discoverer: Whittinghill, 44h.
synonyms: bit2.
phenotype: More extreme than apblt. Wings always
shorter than normal and inflated. Strong tendency
for unequal bifurcation of wing. Wings often break
off and remain attached to pupa cases, hence a
wingless phenotype. Viability about 20 percent of o p * 0 ; apterous-Xasfa
wild type in both sexes. Males fertile but females From Bridges and Brehme, 1944, Carnegie Inst. Wash.
sterile. RK2. Publ. No. 552: 228.
ap blt3
origin: Spontaneous, ap-c: see ap2
discoverer: Semenza, 49k. ap-c: see ap3
synonym: bltS49k. ap-d: see ap4
references: Barigozzi, 1950, DIS 24: 54. Apart: see Apt
phenotype: Wings uniformly inflated, more extreme
than apbtt. Does not overlap wild type. RK2. *apb: apterblister
location: 2-44.7.
apT60: apterous of Thomas origin: Ultraviolet induced.
origin: X ray induced. discoverer: Edmondson, 49K.
discoverer: Thomas, 60g. references: Meyer, Edmondson, Byers, and
references: Meyer, 1963, DIS 37: 50. Erickson, 1950, DIS 24: 59-60.
phenotype: Wings straplike; about 30 percent normal phenotype: Wings always notched, nearly always
length. Adult survives past fifth day. apT6G/ spread and usually blistered, but expression some-
M(2)S24 female lays eggs (Butterworth and King). what variable. Homozygous imagos live less than
RK2. 24 hours, owing to intestinal constrictions that
apXo; apterous-Xasta prevent defecation. Abdomens characteristically
origin: X ray induced in In(2R)Cy; In(3R)P. (The turn dark grey before death, because of accumula-
first X-ray-induced mutation recovered in the tion of digested food products. Although not at
USSR.) same locus as ap, apb +/+ ap4 flies show slight
discoverer: Serebrovsky, 28a. notching of wings and many die within a day;
synonym: Xa. those that survive are fertile, ap5 gives a similar
references: Serebrovsky and Dubinin, 1930, J. heterozygous effect. RK2.
Heredity 21: 259-65. apexless: see apx
phenotype: Wings reduced in length to about 70 per-
cent normal; irregular in outline with a V-shaped Aph°: Alkaline phosphatase deficient
incision with apex at L2, uniformly present, giving location: 3-46.3 (Maclntyre).
wing a mittenlike shape with the thumb between origin: Spontaneous.
marginal vein and L2. Excellent dominant with no discoverer: Johnson.
overlap. Fertile and fully viable in heterozygote. references: 1966, DIS 41: 157-58.
Usually lethal in homozygous conditions, but 1966, Science 152: 361-62.
occasionally ecloses very late as pale dwarf with phenotype: Homozygous larva has no detectable
wings and balancers like vg. Waddington reports alkaline phosphatase activity. Aph°/Aphp larva
deep notch visible in tip of wing fold in prepupa has alkaline phosphatase, which migrates in starch
(1939, Proc. Natl. Acad. Sci. U.S. 25: 299-307; gel electrophoresis to same position as the band in
1940, J. Genet. 41: 75-139 (fig.)]. In homozygotes AphF homozygote. Aph°/Aphs larva has bands of
and in combination with ap4, ®p6, or M(2)S24 wings activity at the Aphs position and at a position
are straplike and 30—70 percent normal length, and slightly faster than the AphF/Aphs hybrid band.
hattere length is 25—50 percent normal; longevity RK3.
fertility like ap4/ap4 except for an occasional long- AphF: Alkaline phosphatase-Fast
lived apx*/M(2)S24 female that may be fertile origin: Naturally occurring allele.
(Schultz; Butterworth and King). apXa'apbtt has discoverer: Beckman and Johnson.
combined pbenoiypes of ap^*,-+ and apblt/apbit references: 1964, Nature 201: 321 (fig.).
(Schultr). RK1A. 1964, Genetics 49: 829-35 (fig.).
MUTATIONS 19
phenotype: AphF/AphF larvae produce an a l k a l i n e Rarely L4 and 5 are interrupted. Males viable and
phosphatase that migrates rapidly in s t a r c h gel fertile; female fertility reduced. RK3.
electrophoresis under conditions d e s c r i b e d by err; abdomen rotatum
Beckman and Johnson (1964). Alkaline phospha- location: 4- (proximal to bt; Fung and Stern, 1951,
tase produced by pupae migrates faster than larval Proc. Natl. Acad. Sci. U.S. 37: 403-4).
enzyme. No enzyme demonstrable in a d u l t s . RK3. origin: Spontaneous.
Aphs: Alkaline phosphatase-Slow discoverer: Beliajeff, 1926.
origin: Naturally occurring allele. references: 1931, Biol. Zentr. 51: 701-8 (fig.).
discoverer: Beckman and Johnson, Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20.
references: 1964, Nature 201: 321 (fig.). Marengo and Howland, 1942, Genetics 27: 604—11
1964, Genetics 49: 829-35 (fig-)- (fig-)-
phenotype: Alkaline phosphatase of Aph^/Aph^ phenotype: Abdomen twisted clockwise through 45°
migrates more slowly in starch gel electrophoresis to 60°. No overlapping with wild type. Male ex-
than that of AphF/AphF. AphF/Aphs larvae pro- ternal genitalia often missing. Males usually
duce a hybrid enzyme of intermediate mobility as sterile; females partially fertile. Puparia not so
well as the fast and slow forms. RK3. smooth as normal; larval segmentation remains.
opp: approximated Puparia have deep constriction near posterior end
location: 3-37.5. just anterior to spiracles. Existing chromosomes
discoverer: Curry, 34a25. marked ar also carry 1(4) and in combination with
references: 1935, DIS 3: 6. Df(4)M show counterclockwise rotation of male ab-
phenotype: Crossveins close together; veins di- domen (Hochman). RK2.
verge at greater angle than wild type; effect vis- cytology: Placed in salivary chromosome region
ible in prepupal wing [Waddington, 1940, J. Genet. 101E through 102B16, on basis of its inclusion in
41: 75—139 (fig.)J. Legs short, with four-jointed Di(4)M =Di(4)101E-F;l02B6-17.
tarsi; the penultimate joint characteristically *ar2
swollen [Waddington, 1939, Growth Suppl. 37—44 origin: Spontaneous.
(fig.)J. Thickset body. Posterior scutellars farther discoverer: Nichols-Skoog, 34el7.
apart than normal. Eyes smaller and flatter than references: Bridges, 1935, Biol. Zh. (Moscow)
normal, also bumpy. Spread wings and thickened 4: 401-20.
veins. RK1. phenotype: Abdomen twisted; male genitalia often
*app61» missing. RK2.
arS7d
discoverer: Puro, 61e. origin: X ray induced,
references: 1964, DIS 39: 64. discoverer: Gloor, 57d.
phenotype: Slightly more extreme than app. RK1. phenotype: Abdomen twisted counterclockwise as
apr: see wa viewed from behind. RK2.
*Apt: Apart
location: 3- (between h and p). origin: X ray induced.
origin: X ray induced. discoverer: Gloor, 57g.
discoverer: Belgovsky, 34e23. phenotype: Abdomen twisted counterclockwise as
references: 1935, DIS 3: 27. viewed from behind. RK2.
phenotype: Wings spread widely. Viability, fertility, At: see Antp1* c
and separability good. Homozygous lethal. RK2A.
cytology: Associated with In(3L)Apt — no salivary arc: see a
analysis, arch: arch
other Information: Apt/D survive; therefore not an location: 2-60.5.
allele of D. origin: Spontaneous.
apterblister: see apb discoverer: Curry, 36g3.
apterous: see ap references: 1937, DIS 7: 5.
*apx: op ex/ess phenotype: Wings curved evenly downward both
location: 1-11.3. longitudinally and transversely, sometimes shorter
origin: Induced by DL-p-NN-di(2-chloroethyl)amino- and blunter, rarely divergent. RK2.
phenylalanine (CB. 3007). arclike wing: see cr/w
discoverer: Fahmy, 1954. arcoid: see ad
references: 1959, DIS 33: 83. arctops: see at
phenotype: Slightly larger fly with large eyes con- arctus ocu/us: see at
taining various numbers of deranged ommatidia. Argentine Curly: see CuA
Wings broad and blunt and in many flies margin Arista: see Ata
removed to various degrees, from a small incision aristaless: see al
of inner margin to removal of entire inner margin, aristaless-h: see aa
costal vein, and parts of the membrane, as far as Aristapedia: see AntpLC
L3. Region from L3 to costal cell unaffected. arp-1: see ssa$P
*as: as cute
location: 3-46. origin: Spontaneous in [n(2L)Cy.
origin: Spontaneous. discoverer: E. B. Lewis.
discoverer: Bridges, 16J21. references: 1945, Genetics 30: 137-66.
references: Bridges and Morgan, 1923, Carnegie phenotype: Similar to ast, but wing veins normal.
Inst. Wash. Publ. No. 327: 170. S/ast3 hatches late; has normal wing veins and
phenotype: Front of scutellum elevated, with partial small eyes similar to but slightly larger than
obliteration of transverse furrow; deep chested. S/ast. RK1A.
Bubble in scutellum or midline of thorax; dried cytology: Same as ast?,
black exudate often at each side of scutellum, may ast*
appear at any of the sutures of head and thorax; origin: Spontaneous recombinational derivative of
black deformed lump behind cheek. Wings droop at ast/ast.
sides. Overlaps wild type. RK3. discoverer: E. B. Lewis.
*as* references: 1945, Genetics 30: 137-66.
origin: Spontaneous. phenotype: ast4/ast4 is usually wild type. S/ast4
discoverer: Bridges, 18116. has smaller eye than S/+; resembles ast/ast in
references: Bridges and Morgan, 1923, Carnegie wing phenotype. RK3.
Inst. Wash. Publ. No. 327: 172. cytology: Salivary chromosomes normal.
phenotype: Same as as. RK3. other information: Recovered as an ast4 ho single
as^9: ascute-hUngende recombinant from an al ast ho/ast female.
origin: Spontaneous,
discoverer: Franke. origin: Spontaneous nonrecombinational derivative
references: 1934, DIS 2: 9. of ast/ast.
Geottschewski, 1935, DIS 4: 15. discoverer: E. B. Lewis,
phenotype: Wings held laterally downward, ends references: 1945, Genetics 30: 137—66.
occasionally resting on legs; eyes small and phenotype: Resembles ast4. RK3.
knobby. RK2. *astr v 1: astero id- re verted
origin: X ray induced in a/ ast ho.
ascutex: see asx discoverer: E. B. Lewis, 1942.
ast: asteroid references: 1945, Genetics 30: 158.
location: 2-1.3 (0.02 unit to right of S). phenotype: Wild type in most combinations, except
origin: Spontaneous. that Df(2L)S4/astrvl slightly more extreme than
discoverer: E. B. Lewis, 38b. Df(2L)S4/+; S** and SM slightly less extreme when
synonym: Sr: Star-recessive. heterozygous with astrvl than with wild type. Ho-
references: 1938, DIS 10: 55. mozygous lethal. RK3A.
1942, Genetics 27: 153-54. cytology: Associated with T(2;3)astrvl =
1945, Genetics 30: 137-66. T(2;3)21E2-3;68C2-3;88D8-9.
1951, Cold Spring Harbor Symp. Quant. Biol. 16: *astry2
159-74 (fig.). origin: X ray induced in al ast ho.
phenotype: Eyes small and rough. Veins L2, L3, discoverer: E. B. Lewis, 1942.
L4, and L5 do not always extend to margin. Over- references: 1945, Genetics 30: 158.
laps wild type rarely. S +/+ ast has very small phenotype: Like astrvl; astrv2/SM overlaps wild
eyes with fused facets; veins L2 to L5 incomplete type. RK3A.
at tip. S ast/+ ast has slightly larger eye than cytology: Associated with ln(2L)astrv3 =
S +/+ ast. S ast/+ + resembles S +/+ +. S +/+ ast In(2L)21E2-3;31.
and ast/ast partially suppress px and net. Eyes of *asfv3
ast/E(S) rough. RK2. origin: X ray induced in net ast dp c/.
cytology: Placed in the 21E1-2 doublet on the basis discoverer: E. B. Lewis, 1942.
of its being included in the synthetic deficiency references: 1945, Genetics 30: 158.
derived by combining the Y-centric portion of phenotype: Wild type in all combinations, except
T(Y;2)21E = T(Y;2)21D4-E1 and the 2-centric por- that S/astrv3 is slightly more extreme than S/+,
tion of T(2;4)astv = T(2;4)2lE2-3;101 (E. B. Lewis, Lethal homozygous and in combination with
1945). Dt(2L)S4 = Dt(2L)21C3-4;22B2-3. RK3A.
cytology: Associated with T(2;3)astrv3 =
origin: Spontaneous in ln(2L)Cy. T(2;3)21E2-3;61C2-3.
discoverer: E. B. Lewis.
references: 1945, Genetics 30: 137-66. asty: asteroid-variegated
phenotype: Similar to ast, but wing veins normal. origin: X ray induced.
S/ast2 lethal. Heterozygote strongly enhanced by discoverer: E. B. Lewis, 1940.
E(S). RK2A. references: 1945, Genetics 30: 137—66.
cytology: Normal except for presence of In(2L)Cy = phenotype: astv/ast and astv/S more variable than
In(2L)22Dl-2;33F5-34Al. but similar to ast/ast and ast/S, respectively;
MUTATIONS 21
suppressed in X/X/Y female. Homozygous lethal. origin: X ray induced.

astv/Df(2L)S2 lethal, RK1A. discoverer: Krivshenko, 1949.
cytology: Associated with T(2;4)ast^ = T(2;4)21E2- synonym: At (symbol preoccupied).
3; 101. references: 1954, DIS 28: 74-75.
astx: asteroid from X irradiation 1955, DIS 29: 73.
origin: X ray induced simultaneously with Sx. phenotype: Lateral branches of aristae reduced,
discoverer: E. B. Lewis. especially branches extending upward from central
references: 1945, Genetics 30: 137—66. axis and situated at base of arista. Axis of arista
phenotype: astx/astx is wild type, but behaves as often abnormal. Wings have small transparent spots
a very slight ast allele in compounds with S and distally. Homozygous lethal. Heterozygous via-
Sx astx. RK3. bility and fertility comparatively high. RK2A.
other information: Separated from Sx by crossing cytology: Associated with T(2;3)Ata =
over in a Sx astx/ast4 ho female. T(2;3)40;66F-67A + T(2;3)47;81.
*osx: ascutex Attenuated: see At
location: 1-26. augenwulst: see awu
origin: Spontaneous. *aw: awry
discoverer: Bridges, 24bl4. location: 1-32 (not allelic to wy).
references: Morgan, Bridges, and Sturtevant, 1925, origin: Induced by ingested radiophosphorus.
Bibliog. Genet. 2: 218. discoverer: Bateman, 1949.
phenotype: Furrow between scutellum and thorax references: 1950, DIS 24: 54.
much shallower, scutellum inflated. Body color 1951, DIS 25: 77.
pale. Legs have blackened leaky joints. Char- phenotype: Wings upcurled, slightly wavy, convex,
acter less extreme in old dry cultures. Viability opaque, or vestigial-like. Variable; overlaps wild
60 percent wild type. RK3. type. Viability about 50 percent wild type. Not
Asy: see A-p enhanced in presence of y as is dvr (1-28.1). RK3.
Asymmettid: see A-p *aw-b: awry-b
*at: arctus oculus location: 1-38 to 39.
location: 2-60.1. origin: Induced by ingested radiophosphorus.
origin: Spontaneous. discoverer: Bateman, 1950.
discoverer: Fernandez Gianotti, 42g28. synonym: aw2.
synonym: bar eye; arctops. references: 1950, DIS 24: 54.
references: 1943, DIS 17: 48. 1951, DIS 25: 77.
1944, DIS 18: 45. phenotype: Like aw. Good expression at 25°C.
1945, Rev. Inst. Genet. Fac. Agron. Vet. Univ. Viability 10 percent that of wild type. Most males
Buenos Aires 2(14): 171-77. fail to eclose. RK3.
1948, DIS 22: 53. awry: see aw
phenotype: Eyes similar to B but with more facets. *awu: augenwulst
Classification, fertility, and viability excellent. location: 2-57.
RK1. origin: Spontaneous.
At: Attenuated discoverer: Rosin, 1951.
location: 1- (in the B region). references: Volkart, 1959, DIS 33: 100.
origin: Induced with soft X rays in phenotype: Eyes deformed; in most extreme expres-
In(l)scSiL-sc8R+dl-49, scsi scsB; associated with sion, deeply indented at middle of anterior margin,
loss of B phenotype. where invaginating integument forms a padlike
discoverer: Valencia and Valencia, 1949. swelling with bristles. Expression variable, often
references: 1949, DIS 23: 64. asymmetrical. Overlaps wild type. Heterozygote
phenotype: In A t/+ females, wings incised medially occasionally has minor effects. Good viability.
and laterally; usually have one large central RK3.
blister. At/At females have badly crumpled, Ax; Abruptex
blistered, and sometimes poorly developed wings. location: 1-3.0.
Wings of At males tend to be more like those of origin: Spontaneous.
At/+ females, although many fall somewhere be- discoverer: Nazarenko, 28a.
tween Af/-s- and At/At in phenotype. Thus there is references: 1930, Biol. Zentr. 50: 385-92 (fig.)-
evidence for only slight dosage compensation for Mohr, 1932, Proc. Intern. Congr. Genet., 6th. Vol.
At. This mutant is similar to some Beadex alleles, 1: 190-212 (fig.).
but allelism with Bx difficult to determine and has phenotype: Homozygous female and male show
not been tested for. Both males and homozygous shortened L5 vein, usually also L4, L2, and some-
females viable and fertile. RK1A. times L3. Wings shortened, arched, thin. Costal
cytology: Associated with ln(l)At ~In(l)16A4- bristles clumped and frayed; costal veins thick-
5;18C4-6;20A2-3. ened. Thorax shows midfurrow with rearranged
*Ata: Arista hair directions; hairs on thorax and head fewer,
location: Not located. with clear patches and streaks. Male genitalia
22 GENETIC VARIATIONS OF DROSOPHiLA MELANOGASTER
often rotated. Ax/+ female shows short L5 in half

of the flies and sparse hair pattern on thorax. discoverer: Nichols-Skoog, 36fl.
Lower temperature (19°C) markedly decreases references: 1937, DIS 7: 5.
expression, and higher temperature enhances it. phenotype: Like b. RK1A in aged flies.
Ax/N8 approaches wild type in all characteristics. cytology: Inseparable from T(2;3)'dp, possibly posi-
Enhanced by H so that Ax/Y; H/+ and Ax/Ax; tion effect or deficiency caused by break distal to
H/+ are nearly lethal at 26° (House, 1959, Anat. 34D. Leads to some ambiguity regarding cytolog-
Record 134: 581-82). RK2 in males. ical location of b.
cytology: A single-band duplication, presumably for
3C7 (Schultz in Morgan, Schultz, and Curry, 1941, origin: Ultraviolet induced.
Carnegie Inst. Wash. Year Book 40: 283). discoverer: Meyer, 50d.
other information: Probably a member of the Notch references: Meyer and Edmondson, 1951, DIS 25: 71.
pseudoallelic complex. phenotype: Somewhat lighter than b. RK2.
origin: Ultraviolet induced.

discoverer: Meyer, 51f.
references: Meyer and Edmondson, 1951, DIS 25: 71.
phenotype: Like b50d. RK2.
*b&: black-Dominant
discoverer: Goldschmidt, 1945.
synonym: 6^.
Zool. 49: 504, 520.
phenotype: bP/b darker than tP/+ or e/e. Homozy-
gous lethal. RK2A.
cytology: Associated with Df(2L)bD =
Df(2L)35C;35D.
Figured in Goldschmidt (1945, p. 520).
B: Bar
location: 1-57.0.
origin: Spontaneous in a female.
discoverer: Tice, 13b.
references: 1914, Biol. Bull. 26: 221-30 (fig.).
Ax; Abruptex Morgan and Bridges, 1916, Carnegie Inst. Wash.
From Mohr, 1932, Proc. Intern. Congr. Genet., 6th. Vol. Publ. No. 237: 66 (fig.).
1: 190-212. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
Genet. 2: 29-33.
phenotype: Eye restricted to narrow vertical bar of
origin: X ray induced. about 90 facets in the male and 70 facets in the
discoverer: Green, 42gl. female as contrasted with normal numbers of about
references: Oliver, 1944, DIS 18: 44. 740 for males and 780 for females [Sturtevant,
phenotype: Similar to Ax, except male lethal. RK2. 1925, Genetics 10: 117-47 (fig)]. Homozygous fe-
b: black male fully viable. B/+ female has about 360 facets
location: 2-48.5. and shows indentation terminating in horizontal
origin: Spontaneous. fissure on anterior margin of eye, producing a
discoverer: Morgan, lOj. kidney-shaped eye. B/B and B/+ completely sepa-
references: Bridges and Morgan, 1919, Carnegie rable from wild type, but in some genetic back-
Inst. Wash. Publ. No. 278: 144 (fig.). grounds B/B overlaps J5/+ slightly. Classifiable
phenotype: Black pigment on body and tarsi and in single dose in triploids by slight anterior nick
along wing veins, darkening with age. Heterozy- in eye (Schultz, 1934, DIS 1: 55); is useful in the
gote shows somewhat darker trident, but is never recognition of triploids. Eyes of female heterozy-
confused with homozygote. Puparium usually gous for a deficiency for B and a normal X are
somewhat lighter than wild type and newly emerged normal (Sutton, 1943, Genetics 28: 97-107). Log
flies not clearly distinguishable from wild type of facet number inversely proportional to tempera-
(Waddington, 1941, Proc. Zool. Soc. London, ture of development (Hersh, 1930, J. Exptl. Zool.
Ser. A 111: 173-80). Tyrosinase formed in adult 57: 283-306).
(Horowitz). RK1 in aged flies.
Nonautonomous over short distances (Sturtevant,
cytology: Salivary chromosomes apparently normal.
1932, Proc. Intern. Congr. Genet., 6th, Vol. 1:
Placed in region between 34E5 and 35D1, on basis
304—7). Facet development enhanced in organ
of its inclusion in Df(2L)64j =•- Df(2L)34E5-
culture by addition of wild type cephalic com-
F1.35C3-D1 (E. H. Grell).
plexes IKuroda and Yamaguchi, 1956, Japan J.
MUTATIONS 23
Genet. 31: 97-102 (fig.)]- Facet number can be in- discoverer: Bridges, 31al5.
creased by addition of a number of compounds to references: Dobzhansky, 1932, Genetics 17:
the medium; probably not a specific inhibition of 369-92.
effect of B (see work of Chevais, Khouvine, Kaji, phenotype: Very slight Bar; merely nick in anterior
Abd-El-Wahab, and DeMarinis). margin of eye in males (no overlap) and in homo-
Embryological studies [Chen, 1929, J. Morphol. 47: zygous females. B4/+ shows slight nick in 10 per-
135-99 (fig.); Steinberg, 1941, Genetics 26: 3 2 5 - cent of cases only. RK2(A).
46 (fig.); 1942, Genetics 27: 171-72; Power, 1942, B36b
Genetics 27: 161, DeMarinis, 1952, Genetics 37: origin: Spontaneous as BB36b in BB chromosome of
75—89 (fig.)] indicate that phenotype results from BB/ln(l)AM female.
reduced number of cells in optic disk and reduced discoverer: Bridges, 36b2.
rate of cell division in anterior part of eye. Facet phenotype: Male resembles standard B; BB36b/+
development responds strongly to environmental female has smaller eye than B/+ but larger and of
factors around 60 hr after oviposition (Luce, different shape than BB/+. Poor fertility both
Quastler, and Chase, 1951, Genetics 36: 488—99). sexes. RK2A.
Pigmented but nonfaceted part of eye shows other information: Homozygous females produce
retinulae and dioptic apparatus lacking, but rudi- wild type and extreme Bar unequal recombinants.
mentary ommatidia present, consisting of hyper-
trophied accessory cells (Wolsky and Huxley, origin: Spontaneous derivative of B in ClB.
1936, Proc. Zool. Soc. London 485-89). RK1A. discoverer: Dempster, 36d9.
cytology: Located in 16A1-2. Associated with references: 1937, DIS 8: 8.
Dp(l;l)B = Dp(l;l)15F9-16Al;16A7-Bl. phenotype: Narrow Bar resembling BB. B36d/+
other information: Since B is a tandem duplication, easy to separate with unaided eye. RK1A.
B homozygotes may give rise to a nonduplicated
chromosome (reversal to normal phenotype) and a origin: Spontaneous in B + .
triplicated chromosome (i.e., double Bar = BB) as discoverer: L. V. Morgan, 36j20.
reciprocal products of unequal crossing over references: 1937, DIS 7: 5.
(Sturtevant and Morgan, 1923, Science 57: 746— phenotype: Slight B; usually stronger than B4, but
47). From successive unequal crossovers in at- shows greater fluctuation and may overlap wild
tached X's, Rapoport (1940, Zh. Obshch. Biol. 1: type. RK3.
235-70; 194.1, DIS 15: 36-37) has been able to *B489
accumulate as many as 7 or 9 Bar regions in a origin: X ray induced in In(l)sc4.
single chromosome. Bar is the first recorded in- discoverer: Yu, 48g.
stance of position effect. Presumably results from references: 1949, DIS 23: 65.
the new band association 16A7-16A1 and can be phenotype: Eyes wider and more variable in width
reversed by rearrangements that separate these than B. Male sterile. RK2A.
bands. Also the first case of cis-trans position cytology: Associated with T(1;2)B486 = T(1;2)15F-
effect, two 16A7-16A1 associations in the same 16A1;33B superimposed on In(l)sc4 =In(l)lB3-
chromosome producing greater facet reduction than 4;19F-2OC1.
two associations in homologous chromosomes;
e.g., facet number in B/B is greater than in BB/+ origin: X ray induced.
(Sturtevant, 1925). discoverer: E. B. Lewis, 5814.
references: Ogaki, 1960, DIS 34: 97.
1960, Japan. J. Genet. 35: 282.
phenotype: At 25°C, male eyes have about five
facets fused into a vertical strip; Bsst/+ female
eyes have about 35. Higher temperature decreases
facet number. Addition of 2.5 percent lactamide to
medium increases facet number to almost 540 in
heterozygous female. Male sterile. RK1A,
cytology: Associated with T(1;3)B5S1 =
B: Bar T(1;3)16A;88F.
Left: heterozygous female. Right: hemizygous male.
From Sturtevant and Beadle, 1939. An Introduction to origin: X-ray-induced partial reversion of B'fl' in
Genetics. Sounders, p. 24. male.
B3 discoverer: Demerec, 34b.
origin: Spontaneous partial reversion of B. references: Sutton, 1943, Genetics 28: 97—107.
discoverer: Stern, 1926. phenofype: Resembles Bl. Viable. RK1A.
phenotype: Eye reduced less than in B and eye cytology: Associated with Dp(l;l)B263-28 -
surface rougher. RK1(A). Dp(l;l)15F9-16Al;16A3-4;16A6-7;16A7-Bl, which
*B4 was derived by deletion of 16A4 of leftmost region
origin: Spontaneous partial reversion of B in a male. through 16A6 of middle region of B^1 triplication.
other information: May be considered to be B1 in size and roughening of remainder. Male sterile;
derived by deletion of one of the regions in the heterozygous female fertile. Interpreted as posi-
BlB1 tandem triplication. tion effect (Dobzhansky, 1936, Biol. Rev.
*B263-34 Cambridge Phil. Soc. 11: 364-84). RK3A.
origin: X-ray-induced reversion of B*B* in male. cytology: Associated with T(l;2)Bbd =
discoverer: Demerec, 34c. T(l;2)16Al-2;48C2-3 + In(2R)41A;47A.
references: Demerec, 1934, Cold Spring Harbor
Symp. Quant, Biol. 2: 110—17.
Sutton, 1943, Genetics 28: 97-107.
phenotype: Eyes wild type. Lethal and cell lethal.
RK2A as lethal.
cytology: B263'34/+ resembles B*B'/+ (Sutton,
1943).
*B263-38
origin: X-ray-induced reversion of BtB* in male,
discoverer: Demerec, 34f.
references: Sutton, 1943, Genetics 28: 97-107. Bbd: Baroid
phenotype: Eyes wild type. Lethal. RK2A as Edith M. Wallace, unpublished.
lethal,
cytology: B263'38/+ resembles BiBl/+ (Sutton, *BDG: Bar of Dubinin and Goldat
1943). origin: X ray induced in B* chromosome in male.
*B263-47 discoverer: Dubinin and Goldat, 1936.
origin: X ray induced in B + male. references: 1936, Biol. Zh. (Moscow) 5: 881—84.
discoverer: Demerec, 38d. phenotype: Eye not described. Lethal when hemi-
references: Sutton, 1943, Genetics 28: 97-107. zygous and homozygous (seems likely that latter
phenotype: Eyes Bar-Mke but larger than Bar. Not claim inferred from former). RK2A as lethal.
lethal. RK1A. cytology: Associated with T(1;2)BD<* = T(1;2)4;15F-
cytology: Associated with ln(l)B263'47 = 16A;20;40-41.
ln(l)16A2-4;20A2-3.
B1: Bar-infrabar
+
origin: X ray induced in B male. origin: Spontaneous partial reversion of B that
discoverer: Bishop, 39i26. occurred in a male.
references: 1939, DIS 12: 61. discoverer: Sturtevant, 1923.
1940, DIS 13: 48. references: 1925, Genetics 10: 117-47 (fig.).
phenotype: Eye reduction in male and heterozygous phenotype: Eye reduction about halfway between B
female between B and B1; size constant in males, and +: B1 male has 478 facets, B^B1 has 320, and
variable in females. Homozygous females viable Bty+ has 716. Facet development inversely pro-
and fertile; show a distinct bb effect. Wings portional to temperature; effective period for
usually leathery and warped at 19°C; normal at temperature treatment is 60 percent through larval
25°. RK2A. life or about 60 hr at 25°C (Luce, 1935, J. Exptl.
cytology: Associated with Tp(l)B263'48 = Zool. 71: 125-47). RK1A.
Tp(l)3E2-3;15F9-16Al;20A2-3. cytology: Apparently no change in Dp(l;l)B in
*Q263-49 which B1 arose.
origin: X ray induced in BB male. *Bl40b
discoverer: Sutton, 41b. origin: Spontaneous in BB male.
references: 1943, Genetics 28: 97—107. discoverer: Steinberg, 40b.
phenotype: Eyes vary in male from BB to wild type, references: 1940, DIS 13: 51.
in homozygous female from BB to B/+. RK2A. phenotype: Similar to B1 both in male and B f /+
cytology: No change in the BB triplication de- female. RK1A.
tectable in salivaries. (Sutton, 1943). cytology: No change from original BB triplication
(Sutton, 1943, Genetics 28: 97-107).
origin: X-ray-induced reversion of BB in male.
discoverer: Sutton, 1940. BMl: Bar of Mullet
references: 1943, Genetics 28: 97-107. origin: X ray induced in B* chromosome.
phenotype: Eyes wild type. Viable. discoverer: Muller, 34e.
cytology: B263'51/+ resembles BB/+ (Sutton, 1943). references: 1935, DIS 3: 29.
; Bar-baroid phenotype: Weak allele of B; always has at least a
origin: X ray induced in B + male. derangement of facets on anterior margin of eye.
discoverer: Dobzhansky, 31bS. RK2A.
references: 1932, Genetics 17: 369-92. cytology: Associated with ln(l)BM1 =
phenotype: Recessive. Eye of male has slight in- In(l)16A2-5;20A3-B (Sutton, 1943, Genetics 28:
dentation of anterior margin, with some reduction 97-107).
MUTATIONS 25
Bridges and Morgan, 1919, Carnegie Inst. Wash.

origin: X ray induced in B+ chromosome. Occurred Publ. No. 278: 148 (fig.).
simultaneously with a reverse mutation of v. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
discoverer: Muller, 34e. Genet. 2: 212 (fig.). 218.
references: 1935, DIS 3: 29. Bridges, 1937, Cytologia (Tokyo), Fujii Jub. Vol.
phenotype: Weak allele of B. RK2A. 2: 745-55.
cytology: Associated with In(l)B^2 = In(l)16A2- phenotype: Wings at first inflated with hemolymph
5;20E button, 1943, Genetics 28: 97-107). to produce blisters and vesicles; venation weak,
*BPar: Bar-partial plexus-like; wings smaller, warped, discolored,
origin: X-ray-induced partial reversal of B in male. divergent. Effect caused by inadequate contrac-
discoverer: Bishop, 1940. tion of epithelium after inflated stage of pupal
references: Sutton, 1943, Genetics 28: 97—107. wing [Waddington, 1940, J. Genet. 41: 75—139
phenotype: Eyes intermediate between B and +. (fig.)]. Sensitive to temperature. RK3 above 25°C;
RK2A. RK2 at 19° or below.
cytology: B duplication unchanged (Sutton, 1943). cytology: Located between 60C5 and 60D2 based on
other information: Six independent partial rever- inclusion within Df(2R)Px = Df(2R)60B8-10;6QDl-2
sions fitting this description found by Bishop. and within Df(2R)Px2 = Df(2R)60C5-6;60D9-10
(Bridges, 1937).
* S R : Bar of Rapoport other information: May be part of a pseudoallelic
origin: X ray induced in normal chromosome. complex with 6s and Px.
discoverer: Rapoport, 1935.
synonym: Bz: Bar of Zuitin.
references: Zuitin, 1935, DIS 4: 6, 16.
1936, DIS 5: 6.
phenotype: More extreme than B. BR/+ resembles
BB/+. RK1(A).
other information: Shows normal crossing over and
reverts to wild type.
*Brev-1: Bar-reversed
origin: X ray induced in B male.
discoverer: Bishop, 1940.
references: Sutton, 1943, Genetics 28: 97—107.
phenotype: Eyes and viability normal.
cytology: B***-1/+ resembles B/+ (Sutton, 1943).
B*: Bar of Sfone

origin: X-ray-induced derivative of B.
discoverer: Stone, 1931. ba: balloon
phenotype: Extreme Bar; produces narrower eye From Bridges and Morgan, 1919, Carnegie Inst. Wash.
than B, both in males and heterozygous females. Publ. No. 278: 148.
RK1A.
cytology: Associated with T(1;4)BS = T(1;4)15F9-
16A1;16A7-B1;1O2F (Griffen, 1940, Genetics 26: origin: Spontaneous.
154-55; Lewis, 1956, DIS 30: 130). discoverer: Banerjee, 58i30.
BS31: Bar-Super inserted in chromosome 3 references: Ray-Chaudkuri, 1959, DIS 33: 99.
origin: Neutron induced in X'YS, sc w B chromo- phenotype: Wings blistered at one or two places,
some. affecting quite a broad area either on the inner
discoverer: Norby. margin or centrally; both wings usually affected.
synonym: BS2; Super-Bar. Wings generally glossy and contracted. Classifica-
references: Muller and Norby, 1949, DIS 23: 61. tion good; viability slightly reduced. RK2.
phenotype: Extreme Bar resembling Bs. RK1A.
cytology: Associated with T(l;3)Bs3i = : see bio
T(1;3)1SF9-16A1;16A7-B1;19-2O;Y;66B13-C1 bio
(Muller; Lindsley). Bag: see Bg
B*: see B*
b-l3 3gis; S ee tri *bal: bandy legged
ba: balloon location: 2- (not located),
location: 2-107.4. origin: Spontaneous.
origin: Spontaneous. discoverer: Stroher, 1958.
discoverer: Morgan, 10k. references: Mainx, 1958, DIS 32: 82.
references: Marshall and Muller, 1917, J. Exptl. phenotype: Legs extremely shortened and crippled.
Zool. 22: 457-70 (fig.). All parts of legs from femur to tarsi shortened,
broadened, and irregularly curved. Tarsal number that suggest that there is enough DNA in 66 + to
not reduced. Deformities most extreme in meta- specify approximately 130 molecules each of 28S
thoracic legs. Movement unsteady and tottering. and 18S ribosomal RNA, these authors view the 66
Manifestation increased by selection. Viability locus as highly redundant and perhaps composed of
poor, especially in males; fertility good. RK2. a very large series of tandem duplications. They
bald: see ra2 interpret bb mutations as partial deletions of the
ballet: see bit locus. They postulate that in 66 flies the rate of
balloon: see ba protein synthesis is limited by the amount of ribo-
Balloon: see Bb somal RNA and the 66 phenotype results in part
balloon wing: see bs3 because normal bristle production represents max-
band: see bn imum protein synthesis on the part of the trichogen
bandy legged: see bal cells during a particular interval in development.
Bar: see B RK2.
Bar + Bar: see BB cytology: Judged to be in 20C2 (or 20C1) by Cooper
Bar double: see BB (1959, Chromosoma 10: 535—88) based on exten-
bar eye: see at sive consideration of published cytology of base
of X chromosome. The 66 locus lies in proximal
bar-3: bar on chromosome 3 heterochromatin of X, probably proximal to and
location: 3-79.1 (not an allele of ro). very close to nucieolus organizer in heterochro-
origin: Spontaneous. matic region hB (Cooper, 1959). Ritossa, Atwood,
discoverer: Ives, 49J20. and Spiegelman (1966), on the other hand, postulate
references: 1950, DIS 24: 58. that the nucieolus organizer is the cytological
phenotype: Like B/B, without significant variation counterpart of the 66 locus. Presence of a
under standard conditions. Viability good. RK1. normal allele of 66 on Y chromosome postulated
Bat-infra double: see 2?* 5* by Burlingame and demonstrated by Stern [l927, Z.
Barlike eye: see Ble Induktive Abstammungs- Vererbungslehre 44: 187—
baroid: see Bbd 231 (fig.)]. This 66 + allele almost certainly in Ys
bat: bat (see Cooper, 1959).
location: 2-71.0. other information: 66 stocks show marked tendency
discoverer: Bridges, 22J26. to accumulate modifiers that suppress the pheno-
synonym: ext-b: extended-b, type. Outcrossing generally brings about return of
phenotype: Wings extended and bent backward. 66 phenotype. Ritossa, Atwood, and Spiegelman
RK2. (1966), however, doubt that this is the case and
*baton: baton postulate that the level of tandem redundancy of
location: 2-52. the locus is subject to frequent stepwise increases
phenotype: Abdomen elongated, with defective or decreases by unequal crossover types of events.
plates; eye resembles L4. Extremely inviable; Many laboratory stocks can be shown by crossing
most homozygotes die in larval and pupal stages, to 6b* to carry 66 alleles of unknown origin.
appearing as elongated corpses. Heterozygote
shows some eye effect. RK3.
bb: bobbed
location: 1-66.0 (Bridges).
discoverer: Sturtevant, 20b.
synonym: 66*. What is now referred to as 6b was
derived from fifth finding of bb. First allele found
was lost and is here omitted from consideration.
phenotype: Bristles of homozygous females de-
creased in both length and thickness. Tergites
etched at sides. Considerable variability of
bristle character, with some overlapping; ab-
dominal character extremely erratic. X/0 male has
phenotype similar to but more extreme than homo-
zygous female. X/Y male is wild type, owing to
presence of normal allele of 66 in Ys; X/X/Y fe-
male similarly normal in phenotype. 66/66' is
extreme 66 in phenotype. Viability variable.
Ritossa, Atwood, and Spiegelman (1966, Genetics
54: 819-34) showed that 66 contains about half as
much ribosomal RNA-complementary DNA as 66 + .
They conclude that the 66 locus is the site of ribo- an extreme bobbed
somal RNA synthesis. On the basis of calculations Edith M. Wallace, unpublished.
MUTATIONS 27
bb$: see bb and bbda is a property of the 66 locus and bb }4

bb<t*>. R K 2 .
origin: Spontaneous in attached X's. *bbG 7: bobbed of Goldschmidt
discoverer: Gabritschevsky, 1926. origin: Spontaneous.
phenotype: When first found, this was a very ex- discoverer: Goldschmidt.
treme 66 with small bristles and very scaly ab- synonym: 66 s P* *P.
domen; it gradually became a weak 6b. Enhances references: 1945, Univ. Calif. (Berkeley) Publ.
expression of gt. RK3. Zool. 49: 3 9 1 - 9 3 .
phenotype: Homozygote normal; bbG1/bbG2 shows
origin: Spontaneous, extreme bristle reduction and abdominal etching.
discoverer: Bridges, 30b24. For interaction with other 66 alleles, s e e
phenotype: bb20/bb is strong bb. bb20 is homozy- Goldschmidt, 1945, table 75. RK2 in some combi-
gous lethal. RK2. nations.
other information: Claimed by Goldschmidt to recur
origin: Spontaneous, in both X and Y chromosomes of certain lines.
discoverer: Stern, 28110.
synonym: bbx. origin: Spontaneous.
references: 1935, DIS 3: 29. discoverer: Goldschmidt.
phenotype: Like 6b. RK2. synonym: bb0 Px aP hi
.
bb': see bbds references: 1945, Univ. Calif. (Berkeley) Publ.
Zool. 49: 391-93.
origin: Thought by Stern and Ogura to be an extreme phenotype: Almost completely lethal homozygous.
66 allele normally occurring on YL in addition to Shows extreme bristle shortening and abdominal
the normal allele occurring on Y$. etching in combination with bbG3. RK2.
discoverer: Stern. other information: Claimed by Goldschmidt to recur
references: Stern and Ogura, 1931, Z. Induktive frequently in certain lines.
Abstammungs- Vererbungslehre 58: 81—121.
phenotype: Only observable evidence of existence origin: Spontaneous.
of 6 6 " i s that, when added to other 66 genotypes, discoverer: Goldschmidt.
it apparently causes them to become slightly less synonym: bt>Poi.
extreme. This could be simply a suppressing references: 1945, Univ. Calif. (Berkeley) Publ.
effect of YL rather than a dosage effect attrib- Zool. 49: 390-93.
utable to a mutant allele of 66. RK3. phenotype: Weak bobbed allele. Extreme in combi-
other information: Inviability of bb*/R(Y)L renders nation with bbG2. RK2 in some combinations*
existence of 6 unlikely. other information: Claimed by Goldschmidt to recur
bba Px SP: s e e bbG1 repeatedly in certain X- chromosomes.
bba Px SP h': see bbG2
*bbD: bobbed-Dominant origin: Spontaneous.
origin: X ray induced. discoverer: Goldschmidt.
discoverer: Lefevre, 48g28. synonym: bbP°i *?.
references: 1949, DIS 23: 58. references: 1947, J. Exptl. Zool. 104: 197-221.
phenotype: Pronounced etching of abdominal phenotype: Bristle effect irregular; no abdominal
tergites; bristles only slightly reduced. Male etching. RK3.
genitalia directed posteriorly rather than ventrally. cytology: Salivary chromosomes normal (Hannah-
Viable in both sexes; fertility of females fair, but Ala va).
of males extremely low. Homozygous females not
produced. Viability and classification good in origin: Spontaneous.
combination with 66 and bbY. RK2- discoverer: Goldschmidt.
other information: Allelism with 66 not definitely synonym: bbPoi hi
.
established. references: 1945, Univ. Calif. (Berkeley) Publ.
bbd*: bobbed-deficiency sensitive Zool. 49: 387, 390-93.
origin: The allele present in some stocks marked phenotype: Homozygous lethal. bbG5/bb produces
66. shortening of bristles and abdominal etching.
synonym: 66 . RK2 as lethal.
phenotype: Females homozygous for bb^s or hetero- bb-': bobbed-lethal
zygous for bbda and a mutant allele of 66 are 66 origin: Spontaneous.
in phenotype. bbd*/Df(l)bb females are lethal. discoverer: Bridges, 1926.
66cfs-like alleles have been reported by Stern and references: Morgan, Bridges, and Sturtevant, 1926,
Ogura (1931, Z. Induktive Abstammungs- Carnegie Inst. Wash. Year Book 25: 308-12.
Vererbungslehre 58: 81—121) and by Lindsley, Stern, 1929, Biol. Zentr. 49: 261-90.
Edington, and Von Halle (I960, Genetics 45: phenotype: Homozygous lethal. 66 J /66 is extreme
1649—70). Presumably, this difference between 66 bb, very late hatching, and invariably classifiable.
bbtybb'/Y = bbl/Y = normal. 66*/0 is lethal. males. In(l)481 = In(l)12E-F;14B induced simulta-
Shown by Ritossa, Atwood, and Spiegelman (1966, neously.
Genetics 54: 819-34) to contain approximately one- *bbOf: bobbed of Offermann
fourth as much ribosomal RNA-complementary DNA origin: X ray induced in T(1;4)A1.
as 66+. RK2A. discoverer: Offermann, 1935.
other information: Segregation from Y chromosome references: 1935, DIS 3: 27.
normal. Crossing over reduced in right end of X in phenotype: Like 6b. RK2A.
6b'/+ females.
origin: X ray induced in In(l)sc8.
L L
origin: Spontaneous in X-Y , bb (X-Y , bb-bb" discoverer: Offermann.
produced X-YL, bbl~2'bb"according to Stern and references: 1935, DIS 3: 28.
Ogura). phenotype: Like 6b. RK2A.
discoverer: Stern, 28k. bbPoi: see bb°3
references: Stern and Ogura, 1931, Z. Induktive bbP°i47: see bbG4
Abstammungs-Vererbungslehre 58: 81—121.
phenotype: Homozygous lethal. bb1'2/bb is bobbed. bbx: see
RK3A. bbY: bobbed on the Y chromosome
origin: X ray induced. discoverer: Bridges, 1926.
discoverer: Lindsley, Edington, and Von Halle. synonym: Ybb.
references: 1960, Genetics 45: 1649—70, phenotype: bb/bbY male has slight 6b phenotype;
phenotype: Lethal as X/0 male and in combination usually separable in crosses, but stock 66/66*'
with In(l)sc4Lsc8R and with bbds. bb1-3a/bb is male tends to change to nearly normal phenotype.
bobbed. RK2A. bbl/btiY male is good 66; always separable and
cytology: Association with Df(l)bbt-3a inferred fertile. RK2 as 66*/66^.
from its irregular segregation from y+Y in males. other information: Chromosome with 66^ described
bbl-74 as Y66 in subsection on Y derivatives.
origin: X ray induced. *bbY-20
discoverer: Lindsley, Edington, and Von Halle, origin: Spontaneous. Arose in combination with
references: 1960, Genetics 45: 1649—70. 6620.
phenotype: Like 66'-3a, RK2A. discoverer: Bridges, 30b24.
cytology: Association with Dt(l)bb1~'''4 inferred synonym: ybb-20t
from irregular segregation from y+Y in males. phenotype: Slightly less extreme but otherwise like
. RK2.
discoverer: Lindsley, Edington, and Von Halle. origin: Spontaneous. Arose in combination with
references: 1960, Genetics 45: 1649—70.
phenotype: Like 66*-3a. RK2A. discoverer: Sturtevant, 31c26.
cytology: Association with Df(l)bbt~iss inferred synonym: Ybb"21<.
from irregular segregation from y+Y in males. phenotype: Like 66*\ RK2.
*bb Y -2 2
origin: X ray induced. origin: Spontaneous. Arose in combination with
discoverer: Lindsley, Edington, and Von Halle. 66".
references: 1960, Genetics 45: 1649—70. discoverer: Curry, 37118.
phenotype: Like 66'-3«. RK2A. synonym: Ybb~22.
cytology: Association with Di(l)bbim4S2 inferred phenotype: Like 66^. RK2.
from slightly irregular segregation from y + F in
male. Bb: Bubble
location: 1- (not located) or 3-48.
origin: X ray induced. origin: X ray induced.
discoverer: Lindsley, Edington, and Von Halle. discoverer: R. L. King, 32d.
references: 1960, Genetics 45: 1649—70. synonym: Balloon.
phenotype: Like bb*-3*. RK2A. phenotype: Wings of heterozygous female smaller,
cytology: Association with Dl(l)bbl-4S6 inferred trimmed, and inflated. Bubble in first posterior
from irregular segregation from y*Y in male. cell. In extreme cases and usually in males, the
wing is a small inflated sac. Sexual difference
origin: X ray induced. in expression may indicate that J56 is on the X.
discoverer: Lindsley, Edington, and Von Halle. Female fertile; male entirely sterile; therefore
references: 1960, Genetics 45: 1649-70. homozygous females not obtainable. RK3A.
phenotype: Like bb'S®. RK2A. cytology: Associated with T(l:3)Bb =
cytology: Association with Dffl)hb!"*8* inferred T(1;3)13E;84F (Morgan, Bridges, and Schultz,
from grossly abnormal segregation from y*Y in 1937, Carnegie Inst. Wash. Year Book 36: 301).
MUTATIONS 29
references: 1925, Genetics 10: 117—47 (fig.).

phenotype: Less reduction in eye than BB. Facet
numbers are 38, 46, and 200 in B'B^B^' female,
BlBl male, and BiBi/+ female, respectively.
Median ocellus lacking or strongly reduced
(Lefevre, 1941, DIS 14: 40). RK1A.
cytology: Associated with Dp(l;l)BB - Dp(l;l)15F9-
16A1;16A7-B1; a tandem triplication.
other information: B* can be recovered as recom-
binant from BiBi/+.
origin: Spontaneous, presumably through unequal

crossing over in B'4(>b/B^0b%
discoverer: Steinberg, 40b.
phenotype: More extreme reduction in eye size than
Bb: Bubble Bi40b. RK2A.
From Bridges and Brehme, 1944, Carnegie Inst. Wash. cytology: Associated with Dp(l;l)BB = Dp(l; 1)15F9-
Publ. No. 552: 23. 16A1;16A7-B1; a tandem triplication.
BB: Bar + Bar bd: see ra2
origin: Spontaneous though unequal crossing over
in B/B (see description of B).
discoverer: Zeleny.
synonym: Bar double; Ultra-bar; double Bar.
references: 1920, J. Exptl. Zool. 30: 292-324
(fig-).
Sturtevant, 1925, Genetics 10: 117-47 (fig.)-
phenotype: Eye more reduced than in B. Facet
numbers are 25, 29, and 45 in BB/BB female, BB
male, and BB/+ female, respectively. Median
ocellus lacking or strongly reduced (Lefevre,
1941, DIS 14: 40). Optic glomerulus reduced
(Power, 1942, Genetics 27: 161). RK1A.
cytology: Associated with a tandem triplication of
the region duplicated in Dp(l;l)B = Dp(l;l)15F9-
16A1;16A7-B1 [Bridges, 1936, Science 83: 210-11
(fig.)]-
BB1: Bar + Bar-infrabar
origin: Spontaneous through unequal crossing over
in B/Bl.
discoverer: Sturtevant.
references: 1925, Genetics 10: 117-47.
phenotype: Like BB. RK1A.
cytology: Associated with Dp(l;l)BB = Dp(l;l)15F9-
16A1;16A7-B1; a tandem triplication, 6 of: Beaded
other information: B is to the left of Bl and both From Bridges and Morgan, 1923, Carnegie Inst. Wash.
types can be recovered as recombinants. Publ. No. 327: 152.
B'B: Bar-infrabar + Bar 6of; Beaded
origin: Spontaneous through unequal crossing over location: 3-93.8.
in B/Bl. discoverer: Morgan, lOe.
discoverer: Sturtevant. references: Dexter, 1914, Am. Naturalist 48:
references: 1925, Genetics 10: 117-47. 712-58 (fig.).
phenotype: Like BB1. RK1A. Bridges and Morgan, 1923, Carnegie Inst. Wash.
cytology: Associated with Dp(l;l)BB = Dp(l;l)15F9- Publ. No. 327: 37, 152 (fig.)-
16A1;16A7-B1; a tandem triplication. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
other information: Bl is to the left of B and both Genet. 2: 45.
types can be recovered as recombinants. phenotype: Wings reduced by marginal excision
B'Bi both anteriorly and posteriorly. Extremely variable
origin: Spontaneous through unequal crossing over and overlaps wild type. Expression and interac-
in B'/Bl. tion studied by Goldschmidt and Gardner (1942,
discoverer: Sturtevant, 1923. Univ. Calif. (Berkeley) Publ. Zool. 49: 103-24).
synonym: Bar-infra double. Almost entirely suppressed by fif. In combination
with many different Minutes, causes incomplete Bridges and Morgan, 1923, Carnegie Inst. Wash.
development of anal and genital imaginal discs in Publ. No. 327: 179 (fig.).
males and less frequently in females (Goldschmidt, Stark and Bridges, 1926, Genetics 11: 249-66.
1948, Proc. Natl. Acad. Sci. U.S. 34: 245-52,' Stark, 1935, DIS 4: 62.
Sturtevant, 1949, Proc. Natl. Acad. Sci. U.S. 35: phenotype: Melanotic tumors appear in larvae and
311-13). Homozygous lethal. RK2 as lethal, RK3 persist in adults. Subject to modification by
as dominant. genetic factors. Nonlethal. RK3.
other information: Bd/In(3R)C, l(3)a was the first Beaded: see Bd
described case of a balanced lethal [Muller, 1918, Beadex: see Bx
Genetics 3: 422-99 (fig.)]. Beadexoid: see Bxd
bending wings: see osbdw
origin: X ray induced. benign tumor in chromosome 3: see be-3
discoverer: Ohnishi, 49116. bent: see bt
references: 1950, DIS 24: 61. bent scutellars: see bsc
phenotype: Like extreme Bd. Variable, overlaps *ber: berrytail
wild type. Homozygous lethal. RK2 as lethal; location: 1-52.4.
RK3 as dominant. origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
; Beaded of Goldschmidt phenylalanine (CB. 3007).
origin: Found among progeny of heat-treated flies. discoverer: Fahmy, 1953.
discoverer: Goldschmidt, 1934. references: 1958, DIS 32: 67.
references: Gottschewski, 1935, DIS 4: 14, 16. phenotype: Abdomen narrow, ending in a berry-like
phenotype: Like Bd but more extreme; not over- protrusion carrying defective genitalia. Wings
lapping wild type in stock. Balancers also re- opaque, with areas of deranged hairs (some with
duced with no overlap. Partially suppressed by H. cut inner margins and interrupted or abnormally
Schultz and Curry report recurrent small or welt- positioned longitudinal veins). Anterior scutellars
like eye effect that is not well understood but is often acutely bent; eyes occasionally misshapen.
in chromosome 3 and may be an effect of BdG. Males sterile and viability about 40 percent wild
Lethal homozygous and in combination with Bd. type. RK3.
RK1. bf: brief
location: 3-95.
discoverer: Goldschmidt. discoverer: Curry, 3813.
references: 1945, Univ. Calif. (Berkeley) Publ. references: 1939, DIS 12: 45.
Zool. 49: 520. phenotype: Fly small; bristles Minute-like. Clas-
phenotype: Like Bd but with more extreme sification perfect, viability fair. Male completely
scalloping effect. RK3. sterile, female with low fertility. RK3.
*BdP: Beaded of Piternick fig; Bag
origin: X ray induced. location: 1-51.6 (to the right of sd).
discoverer: Piternick, 1949. origin: Spontaneous.
references: Goldschmidt, 1953, J. Exptl. Zool. discoverer: Bridges, 33d22.
123: 79-114. phenotype: Heterozygous female with wings shorter
phenotype: Like Bo* but more highly penetrant. and blunter, shortened L5, extra veins or gaps
30—40 percent BdP/+ are phenotypically Bd. near anterior crossvein, and inflated bag centering
Penetrance 100 per cent when heterozygous for in first basal cell. Frequently overlaps wild type.
third chromosome inversions, e.g., In(3L)P + Lethal in male. RK2 as a lethal, RK3 as a
In(3R)P. Interactions with other genes discussed dominant.
by Goldschmidt (1953). Homozygous lethal. RK2 cytology: Probably in 13C, based on Bg-like varie-
as lethal; RK3 as dominant. gation of T(l;3)rasv = T(1;3)9E;13C;81F.
*Bd": Beaded of Wallace *Bg2
discoverer: E. M. Wallace, 15110, discoverer: S^mme.
phenotype: Like Bd but more extreme; ends of L3 synonym: Uw: Uneven wing.
and L4 split or disturbed. Overlaps wild type. phenotype: Heterozygous females with inner wing
Least extreme recognizable phenotype is nick margin frequently nicked or uneven; longitudinal
opposite L3. Homozygous lethal. RK2 as lethal. veins sometimes shortened; one wing often shorter
bdw: see osbdw than the other. Some delay in eclosion. Overlaps
*be~3: benign tumor in chromosome 3 wild type. Viability and fertility fair. Reduces
iocotion: 3-25. size of B eye and is itself exaggerated in combi-
origin: Spontaneous. nation with B. Male lethal. RK2 as lethal; RK3
discoverer: Stark, 16k. as dominant.
references: 1919, Proc. Natl. Acad. Sci. U.S., 5:
573-80 (fig.). origin: Induced by ingested radiophosphorus.
MUTATIONS 31
discoverer: R. C. King, 49h. cytology: Associated with In(3L)Bit; breakpoints

references: Poulson and King, 1949, DIS 23: 62. not determined.
phenotype: Heterozygous female has wings with no bithorax: see bx
crossveins, L5 shortened. Wings asymmetric in Bithoraxlike: see Ubx
size and blistered. L3 very thick; gaps in L3 and bithoraxoid: see bxd
L4. Extra veinlets; veins may fork at wing edge. Bitten: see Bit
Wings often excised terminally and along inner *bk: buckled
margin. Phenotype very variable, overlaps wild location: 1-59.8.
type. Viability 65 percent normal. Male lethal. origin: Induced by p-NN-di-(2-chloroethyl)amino-
RK3. phenylethylamine (CB. 3034).
*BgS2c discoverer: Fahmy, 1955.
origin: Thermal neutron induced. references: 1959, DIS 33: 83.
discoverer: R. C. King, 52c. phenotype: Wings slightly altered in shape and
references: 1952, DIS 26: 65. frequently divergent, with membranes warped be-
phenotype: Like Bg49h', overlaps wild type. Male tween longitudinal veins. Veins slightly thickened
lethal. RK3. at wing margins. Eye shape slightly altered.
hi: bifid Scutellar bristles frequently abnormal, either in-
location: 1-6.9. serted in base atypically, bent, or duplicated.
discoverer: Morgan, Ilk. Males viable and fertile. RK3.
references: Morgan and Bridges, 1916, Carnegie *bk2
Inst. Wash. Publ. No. 237: 28 (fig.). origin: Induced by L-l:6-dimethanesulfonyl mannitol
phenotype: Longitudinal veins fused at base of (CB. 2628).
wing into bifid stalk. L3 delta-like at tip; L4 discoverer: Fahmy, 1960.
often incomplete at tip. Wing margins often ex- references: 1964, DIS 39: 58.
cised at tip of L4. Wings spread in proportion to phenotype: Fly small. Legs shortened; posterior
their shortness. High temperature enhances and pair frequently deformed or absent. Wings
low temperature produces overlapping of wild type. shortened, abnormally shaped, with varying
Stronger in male than in female. Enhances Bx amounts of marginal vein incised. Sex combs may
alleles as well as sd, cp, and vgnP (Waletzky). be enlarged. Bristles stiff; occasionally an extra
RK1. scutellar bristle. bk2/bkl normal. RK3.
cytology: Between 4C7-8 and 4D1-2 according to *Bkd: Blackoid
Demerec, Kaufmann, Fano, Sutton, and Sansome location: 2-65 (Braun).
(1942, Carnegie Inst. Wash. Year Book 41: 191). origin: Spontaneous.
discoverer: Goldschmidt, 1938.
origin: Spontaneous. phenotype: Body color black in homozygote, dis-
discoverer: Gottschewski, 1935. tinctly darker than wild type in heterozygote.
phenotype: Like bi, but w h e r e a s males show 100 RK2.
percent e x p r e s s i o n at 25°C, females show 0—3 *bkl: buckledlike
percent. RK1 in male, RK3 in female. location: 1-59.9.
bis: bistre origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
location: 1-20.1. phenylalanine (CB. 3026).
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- discoverer: Fahmy, 1955.
phenylalanine (CB. 3007). references: 1959, DIS 33: 83.
discoverer: Fahmy, 1954. phenotype: Wings slightly divergent with membranes
references: 1958, DIS 32: 67. warped between longitudinal veins, which them-
phenotype: Very dark brown eye color; ocelli also selves are often slightly thickened. Abnormally
dark. Wings frequently unexpanded. Males sterile. shaped eyes, frequently compressed dorsoven-
Viability varies from less than 10 percent to 70 trally. Both sexes viable and fertile. RK3.
percent wild type. RK2A. other information: Probably a complementing allele
cytology: In bands 7B6 and 7; associated with of bk. One X-ray-induced allele.
Dt(l)bis = Df(l)7B5-6;7B7-8. BI: Bristle
*Bit: Bitten location: 2-54.8 (crossing over may be reduced).
location: 3- (not located; crossing over between ru origin: Spontaneous.
and th almost completely suppressed). discoverer: R. L. King, 25dll.
origin: X ray induced. references: 1927, Biol. Bull. 53: 465-68.
discoverer: Lefevre, 48g5. phenotype: Bristles one-half to two-thirds normal
references: 1949, DIS 23: 58. length, blunt, thicker, and beaded in outline.
phenotype: Inner margin of wing indented. Wings, Posterior scutellars often cross and adhere to
normally folded, appear to have had a bite taken body. Eyes somewhat larger and rougher. Prob-
out of the back. Marginal hairs present unlike N ably affects nature of bristle secretion, partic-
and ct. Flight is impeded, although little wing ularly outer layer [Lees and Waddington, 1942,
area lost. Homozygous lethal. RK1A. DIS 16: 70; Lees and Picken, 1945, Proc. Roy.
Soc. (London), Ser. B 132: 396-423 (fig.)]. Via- origin: X ray induced.
bility of heterozygote is good but erratic; homozy- discoverer: Crowell, 57i.
gotes usually lethal; survivors female-sterile with references: Meyer, 1958, DIS 33: 97.
roughish eye character. RK1 as dominant. phenotype: Eye shape indistinguishable from Bar.
*fi 13 0 Expression of Ble/+ varies; best at 26°C. Ex-
origin: Recovered among progeny of heat-treated cellent expression in homozygote at all tempera-
flies. tures. Ble/Ble in combination with JB results in
discoverer: Plough, 1930. an extremely narrow eye. RK1.
synonym: Sy30: Stubby-30. other information: If Ble represents a transposition
references: Plough and Ives, 1935, Genetics 20: of the Bar locus to chromosome 3, the flanking
42-69. loci of f+ and od+ have not been transposed.
phenotype: Like Bl. RK1. Also against transposition is absence of sexual
dimorphism that dosage compensation of B should
origin: Recovered among progeny of heat-treated produce in such a case.
flies. *bleg: black leg
discoverer: Ives 31119. location: 3- (near p).
synonym: Sy31U9. discoverer: Bridges, 16b23.
references: Plough and Ives, 1935, Genetics 20: references: Bridges and Morgan, 1923, Carnegie
42-69. Inst, Wash. Publ. No. 327: 158.
phenotype: Like Bl. Bl^t/Bl like Bl/Bl and phenotype: Legs black, body color pallid. Wings
poorly viable. Bl311 regularly homozygous lethal. flimsy. RK3.
RK1. blistered; see bs
*h!a: bladderwing Blister I ike: see Bsl
location: 1-43.2. blister/: see by
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- bio: bloated
phenylalanine (CB. 3025). location: 2-58.5.
discoverer: Fahmy, 1953. origin: Recovered among descendants of heat-
references: 1958, DIS 32: 67-68. treated flies.
phenotype: Wings grossly deformed, small, and discoverer: Ives, 33f26.
normally full of fluid. Eyes slightly abnormal in synonym: Originally referred to as ba^: balloon and
shape. Males fertile, females sterile; viability
about 50 percent wild type. RK3. references: Plough and Ives, 1934, DIS 1: 33.
black: see b 1934, DIS 2: 10.
black leg: see bleg 1935, DIS 3: 6.
Blackoid: see Bkd Bridges, Skoog, and Li, 1936, Genetics 21:
bladderwing: see bla 788-95.
Bid: Blond phenotype: Wings spread, crumpled, and vesicu-
location: 1- or 2- (associated with rearrangement). lated; wing shows irregular plexus of extra veins.
origin: Spontaneous in chromosome containing In extreme cases wings unexpanded. Occasional
In(2R)Cy. hooked or wavy bristles. Developmental studies
discoverer: Burkart, 1930. by Waddington [l939, Proc. Natl. Acad. Sci. U.S.
references: 1931, Rev. Fac. Agron. Vet. Univ. 25: 299-307 and 1940, J. Genet. 41: 75-139 (fig.)]
Buenos Aires 7: 393-491. show intervein material spongy and veins swollen,
Burkart and Stem, 1933, Z. Induktive with inadequate contraction after inflated stage of
Abstammungs- Vererbungslehre 64: 310—25 (fig.). pupal wing. Droplets of hemolymph often become
phenotype: Bristles of heterozygote are gleaming clothed with cells liberated from epithelium and
yellow at tips and for varying lengths of more remain along basal processes. Does not overlap
basal regions. Hairs not much paler and bristles wild type, but has poor viability and hatches
of abdomen only slightly affected. Larval mouth later. RK2.
parts wild type. No overlap. Viability and fertility cytology: Not included within and does not recom-
of male and heterozygous female excellent. For- bine with (0/1098) D((2R)Np = Df(2R)44Fl-
merly viable as a homozygous female but in lines 2;45El-2 (Bridges, Skoog, and Li, 1936).
now available the homozygote is lethal, presum- Blond: see Bid
ably owing to a lethal mutation closely linked to blot: see apbit
breakpoint of translocation in chromosome two. *btt: ballet
RK1A. location: 1- (not located).
cytology: Associated with T(l;2)Bld~ T(1;2)1C3- origin: X ray induced.
4,60B12-1J. discoverer: Iyengar.
other information: Bid phenotype associated with references: 1962, DIS 36: 38.
the 2RDXP element of the translocation. phenetype: Wings one-third the normal length;
*Bte: Barlike eye stretched outward and slightly upward; wing tip
location: 3-94. broadened; venation markedly altered as in fused.
MUTATIONS 33
Male viability impaired; females almost com- origin: Spontaneous.

pletely lethal. RK2. discoverer: Fabian, 1941.
bit: see apblt references: 1948, Arch. Julius Klaus-Stift.
*blu: blunt Vererbungsforsch. Sozialanthropol. Rassenhyg.
location: 3- (near ru). 23: 512-17.
origin: Spontaneous. phenotype: Eye color dark brownish red; darkens
discoverer: Wallbrunn, 46j23. with age. Malpighian tubules and testis sheaths
references: 1947, DIS 21: 71. colorless. Male fertile; female sterile. RK2.
phenotype: Wings slightly shorter and broader than *tow: bow wings
normal, giving a squared appearance. Sometimes location: 1- (not located).
difficult to classify. RK3. discoverer: Bridges, 12hl5.
*bn: band references: Morgan and Bridges, 1916, Carnegie
location: 3>72. Inst. Wash. Publ. No. 237: 46 (fig.).
origin: Spontaneous. phenotype: Wings curved downward over abdomen
discoverer: Morgan, 12g. and also sideways like bowl of a spoon. Overlaps
references: Bridges and Morgan, 1923, Carnegie wild type. RK3.
Inst. Wash.. Publ. No. 327: 79 (fig.). bow-legged: see bwl
Morgan, Bridges, and Sturtevant, 1925, Bibliog. bowed: see bod
Genet. 2: 215 (fig.), 218. bp: see bulbP
phenotype: Trident pattern and scutellum darker,
with dark transverse band across anterior portion
of mesonotum. Thorax vacuolated; hairs on thorax
sparse and directed medially, in bowed lines.
RK2.
DO: bordeaux
location: 1-12.5.
discoverer: Nazarenko.
phenotype: Eye color dark wine; not completely
separable from wild type. Red pigment 67 percent br: broad
wild-type level; brown pigment normal (Nolte, Edith M. Wallace, unpublished.
1955, J. Genet. 53: 1—10). Transplantation indi-
cates bo may be nonautonomous (Ephrussi and br: broad
Beadle, 1937, Genetics 22: 65-75). Larval Mal- location: 1-0.6.
pighian tubules bright yellow (Beadle, 1937, discoverer: Bridges, 15i26.
Genetics 22: 587-611). RK3. references: Morgan, Bridges, and Sturtevant, 1925,
bobbed: see bb Bibliog. Genet. 2: 145, 220 (fig.).
bobbed on the Y chromosome: see bbY phenotype: Wings somewhat broader than and about
bod: bowed 80 percent of length of normal, with round full tip;
location: 3-48.3. crossveins closer together. Shape difference vis-
origin: Spontaneous. ible in middle prepupal stage immediately after
discoverer: Nichols-Skoog, 35b20. eversion; probably an influence on cell division
references: 1937, DIS 7: 6. [Waddington, 1939, Proc. Natl. Acad. Sci. U.S.
phenotype: Wings bowed downward over abdomen, 25: 299-307; 1940, J. Genet. 41: 75-139 (fig.)].
curvature along both axes; curvature occasionally RK1.
reversed. Wings somewhat smaller than wild type. cytology: Located between 1C5 and 2C10 (Demerec,
Whole fly smaller and humpy; eyes slightly bulged. Kaufmann, Fano, Sutton, and Sansome, 1942,
Overlaps wild type slightly. Viability 75 percent Carnegie Inst. Wash. Year Book 41: 191).
wild type. RK3. br3
*bord: bordered origin: Spontaneous.
location: 1-70. discoverer: Bridges, 31el.
discoverer: Bridges, 1916. phenotype: Wings like br but more often arc-like
references: Morgan, Bridges, and Sturtevant, 1925, and crumpled. RK2.
Bibliog. Genet. 2: 220.
phenotype: Wings smaller and slightly extended; origin: Induced by DNA.
venation ragged and veins bordered by darker discoverer: Fahmy, 1959.
bands. Viability poor; classification unreliable. phenetype: Extreme expression of or"*? phenotype.
RK3. brS9!/br and bcS9i/ brutl have mutant phenotype.
bordeaux: see bo RK1.
bordered: see bord *brD: broad-Dominant
*bos: bordosteril origin: Spontaneous,
location: 3-0.0. discoverer: Muller, 19h.
references: Muller and Altenburg, 1921, Anat. Rec. discoverer: Fahmy, 1956.
20: 213. references: 1959, DIS 33: 83.
Muller, 1935, DIS 3: 29. phenotype: Fly with broad abdomen and slightly
phenotype: brD/+ resembles br/br. or-D/orand shortened thorax and wings. Wings frequently
brD/Y are lethal. RKl. slightly divergent. Eyes small and dull red with
*Jbr/-o; broad-lethal-a reflection spots. Bristles slightly shortened and
origin: Spontaneous. lying flatter on thorax. Males and females viable
discoverer: Muller, 19h. and fertile. RK2.
synonym: ifljfer^. other information: One allele induced by CB. 3025.
references: Muller and Altenburg, 1921, Anat. Rec. brc: brachymacrochaetae
20: 213. location: 1-0.0 (no recombinants with sc among
Muller, 1935, DIS 3: 29. 6746 sons).
phenotype: brlmB/br female is phenotypically br. origin: Induced by triethylenetnelamine (CB. 1246).
br'-* male dies. RK2. discoverer: Fahmy, 1952.
origin: Spontaneous. phenotype: One or more thoracic bristles much
discoverer: Muller, 19h. reduced in size; scutellars and dorsocentrals most
synonym: l(l)brb. frequently affected. Occasional bristles dupli-
references: Muller and Altenburg, 1921, Anat. Rec. cated. Good viability and fertility in both sexes.
20: 213. RK2.
Muller, 1935, DIS 3: 29. other information: One allele each induced by
phenotype: Like 6r;"a. RK2. CB. 3025, by CB. 1246, and by X rays.
*br*h: broad-short *brd: broadened
origin: Spontaneous. location: 1-33.
discoverer: Bridges, 14g20. origin: X ray induced.
references: 1916, Genetics 1: 151. discoverer: Muller, 26127.
phenotype: Wings one-half to two-thirds normal references: 1935, DIS 3: 29.
length, often arc-like and crumpled. Crossveins phenotype: Wings expanded. Viability 20 percent
closer together than normal and slight plexus wild type. RK3.
effect. Legs gnarled. RK2. *bre: bright eye
*brui: broad-unequal wings location: 1-24.6.
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- origin: Induced by L~p-NN-di-(2-chloroethyl)amino-
phenylalanine (CB. 3026). phenylalanine (CB. 3025).
discoverer: Fahmy, 1955. discoverer: Fahmy, 1953.
synonym: uq. references: 1958, DIS 32: 68.
references: 1958, DIS 32: 77. phenotype: Eye color brighter red. Wings shorter,
phenotype: Wings short and very broad, often often crumpled or waved. Abdomen disproportion-
unequal in length; more extreme at 25°C. Occa- ately large. Male viability and fertility good, but
sionally one wing blistered, or grossly deformed. females have reduced fertility. Not easily classi-
Thoracic hairs irregularly distributed and sparse. fied. RK3.
Ocasional abnormality in bristle position or size. other information: One allele induced by CB. 1540.
Eyes smaller. Viability and fertility good in both brevis: see bv
sexes, bruq/br is wild type. RKl. bri: bright
other information: One X-ray-induced allele. location: 2-54.3.
*Br: Bridged origin: Spontaneous.
location: 1- (right half; crossing over suppressed to discoverer: Nichols-Skoog, 34b23.
the right of v). references: Beadle and Ephrussi, 1937, Am.
origin: X ray induced. Naturalist 71: 91-95.
discoverer: Muller, 2713. phenotype: Eye color bright red like en2 or v2;
references: 1935, DIS 3: 29. difficult to separate from wild type. Malpighian
phenotype: Plexus-like wings, with extra cross- tubules pale yellow (Beadle, 1937, Genetics 22:
veins bridging logitudinals. L4 bent. Wings 587-611). RK3.
arched. Male lethal. RK3A. Bridged; see Br
cytology: Associated with In(l)Br. brief: see bf
Br: see Sp bright: see bri
brachymacrachaetae: see brc bright eye: see bre
bran: see aba Bristle: see Bl
Bran: see aBa Bristled: see Sp
Branchlet: see Bt broad: see br
*brb; broad abdomen broad abdomen: see brb
location: 1-52.9. broadened: see brd
origin: Induced by styrylquinoline (CB. 3086). broader wing: see brw
MUTATIONS 35
2
bronze: see sf effect owing to additive semidominance; was
bronzy: see malbz originally interpreted to indicate allelism. RK2.
brown: s e e few *bs4
brown-like1: see red origin: Spontaneous.
brunette: see Hnr2 discoverer: Goldschmidt.
*6rw: broader wing references: 1947, J. Exptl. Zool. 104: 197-221.
location: 1-39.8. phenotype: More extreme allele of bs. RK2.
origin: X ray induced. cytology: Salivary chromosomes normal (Hannah).
discoverer: Fahmy, 1956. ; s e e bs*>l
phenotype: Wings broad and rounded at the tips. origin: Spontaneous.
Males show reduced viability and are sterile. discoverer: Strangio.
RK3. phenotype: More extreme allele of 6s with variable
expression, but wing generally converted to one
*bs: blistered large blister. Fully penetrant at 20°, 25°, and
location: 2-107.3. 30°C; blistering effect most marked at 20°C. RK2.
origin: Spontaneous. cytology: Salivary chromosomes normal (Strangio).
discoverer: Bridges, I l k l 6 .
references: Bridges and Morgan, 1919, Carnegie origin: Spontaneous.
Inst. Wash. Publ. No. 278: 155 (fig.). discoverer: Mohler, 54j7.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. references: 1956, DIS 30: 78.
Genet. 2: 219 (fig.). phenotype: bsS4i /bsS4i = bssî/bs2 more extreme
phenotype: Wings blistered, small, pointed; vena- and less variable at 25°C than bs2/bs2. Viability
tion thick and plexus-like, with branches from and good in uncrowded cultures. RK2.
parallel to L5 beyond second crossvein, where
there is a semidominant free vein effect. Eye origin: Spontaneous derivative of 6s.
color softened. Temperature sensitive. RK2 at discoverer: Goldschmidt, 38i.
19°C; RK3 at 25°C. synonym: bs3Si.
cytology: Located between 60C5 and 60D2, based references: 1945, Univ. Calif. (Berkeley) Publ.
on its inclusion within Df(2R)Px = Df(2R)60B8- Zool. 49: 408-9, 416-17.
10.60D1-2 and within Df(2R)Px2 = Df(2R)60C5- phenotype: Stronger allele than bs2 but of low
6;60D9-10 (Bridges, 1937, Cytologia (Tokyo), viability and fertility. Wing shows very extended
Fujii Jub. Vol. 2: 745-55). web near the crossvein, which reaches margin of
other information: May be part of a pseudoallelic wing; above web a blister is formed. RK2.
complex with ba and Px.
bscy; blistered-curly
discoverer: King and Poulson, 461.
references: Poulson and King, 1948, DIS 22: 54.
phenotype: Heterozygotes show venation abnormal-
ities, with tiny free veins usually in the third
posterior cell and occasionally in the second in
about 50 percent of males and 90 percent of fe-
males. Less often extra veinlets project from
bs*: blistered-2 posterior crossvein. Homozygotes have bizzare
Edith M. Wallace, unpublished. networks of wing veins. Wings of freshly emerged
adults inflated with hetnolymph, producing large
blister in middle of one or both wings just pos-
discoverer: Bridges, 25k24. terior to the anterior crossvein. Blisters cover
phenotype: More extreme allele of 6s. one-fourth to one-half of total wing area. Lymph
bs2/Df(2R)Px easily separable from +/Df(2R)Px, later dries, leaving wing vesiculated and curled
especially at or below 19°C. Wing effect caused upward. Condition more pronounced in females.
by same mechanism as that of px [Waddington, Flight restricted. bscy/bs2 females are like weak
1940, J. Genet. 41: 75-139 (fig.)]. RK2. bs°y and males like bs2. Viability nearly normal.
S3 RK1.
origin: Spontaneous. bsP
discoverer: Swigert, 31 d. origin: Spontaneous derivative of bs,
synonym: balloon wing; px3id*and px*>«, discoverer: Goldschmidt.
references: Plough and Ives, 1934, DIS 1: 33. references: 1945, Univ. Calif. (Berkeley) Publ.
1935, Genetics 20: 42-69. Zooi. 49: 409-18.
phenotype: Extremely blistered wing. Classifica- phenotype: Stronger allele than 6®. Sensitive to
tion easy in most stocks, + bs3/px + shows wing genetic modification. Females show broad chi-
tinous mass of extra veins; males show extra references: 1914, J. Exptl. Zool. 17: 325-36.
veins branching from L2. RK2. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
*bsPP Genet. 2: 216 (fig.), 219.
origin: Spontaneous derivative of 6s. Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20.
discoverer: Goldschmidt. phenotype: Wings held out at base and bent sharply
references: 1945, Univ. Calif. (Berkeley) Publ. backward. Rear legs often lumpy at first tarsal
Zool. 49: 409-18, 433-39. joint. May have one to four "preleg" or "first
phenotype: Stronger allele than bs. RK2. ventral" bristles on ventral surface of thorax
cytology: Frequently associated with short defi- anterior to first pair of legs, in space otherwise
ciency to the right of 6a (or 6s ?) within devoid of bristles or hairs. Overlaps wild type at
Df(2R)Px2? 25°C, very much at 19°, and little if any at 29°C
other Information: Claimed by Goldschmidt fre- (Metz, 1923, Proc. Soc. Exptl. Biol. Med. 20:
quently to recur by mutation in certain lines. 305-10). RK2 at 28°C.
*bsc: bent scutellars
location: 1-1.1.
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
phenylalanine (CB. 3007).
phenotype: One or more scutellars bent on them-
selves in form of inverted V. Other bristles irreg-
ularly bent. Eyes slightly smaller. Wings slightly
abnormal in shape. Male viability about 50 percent
wild type; fertility much reduced. RK3.
other information: One allele each induced by
CB. 3025 and CB. 3026.
*Bsl: Blister I ike

location: 2-104 (but located 3.2 units from bw
which is at 104.5).
discoverer: Oliver, 29bl. bt: bent
references: 1939, DIS 11: 47. From Morgan, Bridges, and Sturtevant, 1925, Bibliog.
phenotype: Extra veins and blister centering in Genet. 2: 216.
region of posterior crossvein, which is usually
absent. Fluctuation in expression from thickening *btD: bent-Dominant
of veins to blister covering entire wing. Vein L5 origin: X ray induced.
may be short. Homozygous lethal. RK3. discoverer: Schultz, 33all.
cytology: Not done, but Bridges has suggested that references: Bridges, 1935, Biol. Zh. (Moscow) 4:
this is probably a Plexate deficiency. 401-20.
bsp: brown spots phenotype: When found, btD/+ showed regularly
location: 2-40.6. divergent wings with some angular bend near base.
origin: Spontaneous. Legs lumpy at low temperature. Preleg bristles
discoverer: Di Pasquale. present as in bt. Homozygous lethal. RK3 as
references: 1959, DIS 33: 128. lethal.
Di Pasquale and Zambruni, 1963, DIS 37: 73 (fig.). other information: Balanced stocks in existence to-
1966, DIS 41: 119. day show only preleg bristle character and reces-
phenotype: Spots of brown pigment appear in integ- sive lethality (Lewis).
ument of bsp/bsp females only after they have
mated. Di Pasquali and Zambruni (1963) showed *Bt: Branchlet
that copulation with any male, sterile or fertile, location: 1- (rearrangement).
triggers formation of brown spots. Courtship with- origin: Induced by P 3 2 .
out copulation ineffective; virgin females never discoverer: Bateman, 1950.
show brown spots. No phenotype in males. Pene- references: 1950, DIS 24: 54.
trance 60—80 percent; viability excellent. RK3. 1951, DIS 25: 77.
bt: bent phenotype: Heterozygous female has posteriorly di-
location: 4-1.4 [mapped in diplo-4 triploids by rected branchlet on posterior crossvein as well as
Sturtevant (1951, Proc. Natl. Acad. Sci. U.S. 37: other extra venation. Abdominal segments often
405-7)]. poorly chitinized. Male lethal. RK3A.
origin: Spontaneous. First mutant found on chromo- cytology: Associated with Dp(l;l)Bt - Dp(l;l)3B2-
some 4. Cl;6F6-7.
discoverer: MuIIer, 1914. other information: Phenotype may be Co.
MUTATIONS 37
*bu: bulging phanotype: Eye color reddish brown; brilliant

location: 1-58. orange in combination with en. Good viability.
origin: X ray induced. RK1.
discoverer: Muller, 2618. burnt orange: see Jbuo
references: 1935, DIS 3: 29. bv: krmvis
phenotype: Eyes rough and bulging. Semilethal. location: 3-102.7 (recalculated from Sturtevant,
RK3. 1956, Genetics 41: 118-23).
bv: see Hn™ discoverer: Bridges, 33e25.
bu-w61): see vs61i phenotype: Bristles uniformly short and stubby.
Bubble: see Bb Body chunky. Hatches late but viability excellent.
bubble wing: see vsîj RK1.
buckled: see bk bw: brown
buckledlike: see bkl location: 2-104.5.
bul: bulge discoverer: Waaler, 19J15,
location: 3-43.6. references: 1921, Hereditas 2: 391-94.
origin: Spontaneous. Sturtevant and Beadle, 1939, An Introduction to
discoverer: Spencer, 36d28. Genetics, Saunders, p. 64 (fig.).
references: 1937, DIS 7: 6. phenotype: Eye color light brownish wine on emer-
Curry, 1939, DIS 12: 45. gence, darkening to garnet. Red pigments lacking;
phenotype: Eyes very large and bulging; facets ommochromes at 87 percent normal level (Nolte,
rounded, in irregular rows, and some quite large. 1954, J. Genet. 52: 111-26). Adult testes and
Wing margin heavy; end of wing somewhat squared vasa colorless. Larval Malpighian tubules pale
off to L3. RK3. yellow (Beadle, 1937, Genetics 22: 587-611).
bulbP: bulge-bumpy Produces white eyes in combination with v, en, or
origin: Spontaneous, st. Eye color autonomous when transplanted into
discoverer: E. H. Grell, 1955. wild-type host (Beadle and Ephrussi, 1936,
synonym: bp, Genetics 21: 230). RK1.
references: 1955, DIS 29: 72. cytology: Placed between 59D4 and 59E1 by Bridges
phenotype: About one-half the eye surface erupted [1937, Cytologia (Tokyo), Fujii Jub. Vol. 2: 7 4 5 -
into irregular yellowish blisters. Facets larger 55], on the basis of its exclusion from the inner
than normal in nonblistered areas. Homozygotes inversion of In(2LR)bw^l = ln(2LR)21C8-
occur with 1 percent of expected frequency. Sur- Dl;60Dl-2 + In(2LR)40F;59D4-El and its inclusion
viving homozygotes vigorous and male fertility in In(2R)bwvOe2 = ln(2R)41A-B;59D6-El. Based
high; females lay eggs abundantly, but only rarely on the study of bw rearrangements, Slatis (1955,
does an egg hatch. RK3. Genetics 40: 5-23) tentatively places bw in 59D9,
bulging: see bu 10, or 11.
other information: Separable into at least two sub-
*buo: burnt orange units by recombination with bw and bw75 about
location: 2-57.1. 0.001 units to the left of bw** and
origin: Spontaneous. (Divelbiss, 1961, Genetics 46: 861).
discoverer: T. Hinton and Kleiner, 1941.
references: Hinton, 1942, DIS 16: 48. origin: Spontaneous.
phenotype: Eye color bright orange-brown. Mal- discoverer: P. R. Sturtevant, 1921.
pighian tubules colorless in larva (Brehme and references: Morgan, Bridges, and Sturtevant, 1925,
Demerec, 1942, Growth 6: 351-56). RK2. Bibliog. Genet. 2: 220.
other information: Not an allele of en. Allelism phenotype: Darker and redder than bw. RK1.
with ltd (2-56) apparently never tested.
bur: burgundy origin: X ray induced,
location: 2-55.7. discoverer: Demerec, 28dl3.
origin: Ultraviolet induced. phenotype: Like bw2. RKl.
discoverer: Edmondson and Meyer, 49c.
references: Meyer and Edmondson, 1949, DIS 23: origin: Spontaneous,
60. discoverer: Emerson, 32dl9,
phenotype: Eye color dull, darkish brown, like pr; phenotype: Like bw*. RKl.
brilliant orange in combination with en. Classifi- bw*
cation and viability excellent. Fertility of females origin: Spontaneous.
good; of males, variable. RK1. discoverer: Mohr, 31k28.
other information: Not allelic to It, ltd, or pr. phenotype: bw*/bw* is wild type. bw4fbws is
bur2 purpleoid-like. bw4fbw like bw but darker. RK3.
origin: Spontaneous. other information: aw"* originally found in bw4/bw$
discoverer: Hall, 60h. combination and called purpleoid-like. bw4 and
references: Meyer, I960, DIS 34: 52. bw-5 separated by Bridges.
38 GENETIC VARIATIONS OF DROSOPH/LA MELANOGASTER
phenotype: Homozygote not clearly distinguishable

origin: Spontaneous. from wild type. Also wild type in combination
discoverer: Mohr, 31k28. with bw24 and bw7s. Produces a phenotype inter-
phenotype: bws/bw4 is purpleoid-like (see bw4); mediate between bw and wild type in combination
bws/bw is light yellowish brown; bws/+ is wild with bw and bw^l. RK3.
type; bwS/bw* is lethal. RK2A. other information: Shown to lie to right of bw and
cytology: Associated with a deficiency or an inver- bw7S by recombination; nonrecombinant reversions
sion involving 59E1 to 59F1 (Schultz). recovered from bw*9/hw81 (Divelbiss, 1961,
*bw24 Genetics 46: 861).
origin: X ray induced. bw61i: see vs61J
discoverer: Slatis, 1950.
references: 1955, Genetics 40: 5-23. origin: X ray induced.
phenotype: Wild type in combination with bwS9 and discoverer: Slatis.
bw7S, but shows intermediate phenotype in combi- references: 1955, Genetics 40: 5-23.
nation with bw and bwai. Studied only in males phenotype: Like bw. RK1.
(see cytology). RK3A. cytology: Salivary chromosomes normal.
cytology: Associated with but presumably w72
separable from T(Y;2)R24 = T(Y;2)45A;51E. origin: X ray induced.
discoverer: Slatis.
origin: Spontaneous. references: 1955, Genetics 40: 5—23.
discoverer: Clancy, 37g26. phenotype: Like bw. RK1.
references: 1938, DIS 10: 55. cytology: Salivary chromosomes normal.
phenotype: Eye color like bw but darker (like
bw2 ?). Produces yellowish pigment in combina- origin: X ray induced.
tion with v. RK1. discoverer: Slatis, 50f25.
origin: Spontaneous in In(2L)Cy 4- In(2R)Cy, 1955, Genetics 40: 5-23.
Cy en2 sp2. phenotype: Intermediate allele of bw. bw7*/bw7s
discoverer: Ives, 45a. darker than bw7* /bw; about 1 percent of bw7* /bw
references: 1945, DIS 19: 46. flies show twin spots in eye. RK1.
Ives and Scott, 1948, DIS 22: 71. other information: Located to the left of bw81 and
Ives and Evans, 1951, DIS 25: 107. bw59 by recombination (Divelbiss, 1961, Genetics
phenotype: bw4Sa/bw is brown; bW4Sa/bw4 is wild 46: 861).
type; therefore bw45* behaves like bw4. Homozy-
gote has not been tested because bw45* has not origin: X ray induced.
been separated from Cy* RK3A. discoverer: Slatis, 50hl.
other information: Probably occurred simulta- references: 1951, DIS 25: 75.
neously with or45*. 1955, Genetics 40: 5-23.
phenotype: Intermediate allele; more extreme than
bw7*. RK1.
origin: Spontaneous. other information: Located to the right of bw and
discoverer: Ives, 47j. bw7* by recombination; bwsl /bw59 produces non-
references: Ives and Scott, 1948, DIS 22: 71. recombinant reversions (Divelbiss, 1961, Genetics
Ives and Evans, 1951, DIS 25: 107. 46: 861).
phenotype: Brown in combination with In(2L)Cy + *bw+21
In(2R)Cy, Cy en2 bw4** sp2 or4**, but wild type in origin: Isoallele of bw recovered from an X-ray-
combination with bw, bw2b, bw4, or and or4*B. induced brown-Variegated mosaic.
Homozygous normal. RK3. discoverer: Slatis.
other information: Several alleles of this type with references: 1955, Genetics 40: 5-23,
varying degrees of expression found by Ives in a phenotype: Normal, but gives less pigment in
natural population from South Amherst, Mass. combination with variegating alleles than other
bw+ alleles.
origin: Spontaneous, cytology: Salivary chromosomes normal.
discoverer: Clancy, 53i. bw°: brown-amber
phenotype: Intermediate between few and bw37t discoverer: R. C. King, 48fl5.
both alone and in combination with v. RK1. references: Poulson and King, 1948, DIS 22: 54.
phenotype: Eye color light brownish yellow. Adult
origin: X ray induced, testes and vasa colorless. Larval Malpighian
discoverer: Slatis, 50fl6. tubules slightly paler yellow than wild type.
references: 1951, Dig 25: 75. bW/bw gives eye color slightly lighter than bw.
1955, Genetics 4Ch 5-23. RK1.
MUTATIONS 39
: brown-Auburn phenotype: Like bw except testis sheath pale

origin: X ray induced. yellow in older males. Gives some pigmentation
discoverer: Dubinin, of eyes in combination with v. RK1.
synonym: A; Pm^'. by/R3: brown-Rearranged
references: Dubinin and Heptner, 1935, J. Genet. origin: X-ray-induced derivative of bw.
30: 423-46 (fig.). discoverer: Slatis, 48kl6.
Dubinin, 1936, Biol. Zh. (Moscow) 5: 851-74. references: 1955, Genetics 40: 5-23.
phenotype: Nearly uniform brown, but with extra Y phenotype: Heterozygotes with 6w+ strongly varie-
chromosome shows strong variegation. Homozy- gated. Homozygous lethal; lethal or semilethal in
gote usually lethal. RK1A. combination with other bw rearrangements. RK1A.
cytology: Associated with In(2R)bwA = cytology: Associated with In(2LR)bwR3 =
In(2R)41;59D. In(2LR)40F;51F;55E;57E;58D8-9.
*bwR4
*bwAD: brown of A. Das origin: X-ray-induced derivative of bw.
origin: Spontaneous. discoverer: Slatis.
discoverer: Das, 63a7. references: 1955, Genetics 40: 5—23.
synonym: bw3^. phenotype: Expression in bwR4/+ varies from
references: Sarkar, 1963, DIS 38: 28. moderate to strong, depending on origin of bw.
phenotype: Eye color light brown, darkening with Homozygote brown and almost completely lethal.
age. RK1. RK1A.
cytology: Associated with T(2;3)bwR4 =
*bwCB: brown-Chester Beatty T(2;3)S9B2-3;80-81.
origin: Induced by 2-chloroethyl methanesulfonate *bwR 7 2
(CB. 1506). origin: X ray induced.
discoverer: Reddi. discoverer: Slatis.
references: 1960, DIS 34: 53. references: 1955, Genetics 40: 5-23.
phenotype: Lethal allele of bw. RK2. phenotype: Weakly mottled in bwR12/+ heterozy-
cytology: Salivary chromosomes normal (Slizynska). gote; bwR12 homozygote somewhat more extreme.
bwD: brown-Dominant Homozygote semilethal. RK1A.
origin: Spontaneous. cytology: Associated with T(2;3)bwRl2 =
discoverer: T. Hinton, 1940. T(2;3)59D;80C.
1942, DIS 16: 48.
Slatis, 1955, Genetics 40: 246-51. origin: X ray induced.
phenotype: Eye color varies with age from purple to discoverer: Slatis.
brown. Shows slight variegation in combination references: 1955, Genetics 40: 5—23.
with st (Slatis, 1955). Wings pebbled. Variegation phenotype: Variegation intermediate. Homozygous
suppressed by extra Y chromosomes (Brosseau, lethal. RK1A.
1959, DIS 33: 123). Homozygote viable and fertile. cytology: Associated with T(2;3)bwRl* -
Larval Malpighian tubules bright yellow (Brehme T(2;3)59E2-3;80.
and Demerec, 1942, Growth 6: 351-56). RK1A.
cytology: Shultz reports an extra band in 59E that origin: X ray induced.
tends to pair with a band in the homolog, sug- discoverer: Slatis.
gesting a duplication of one band from 59E. Slatis references: 1955, Genetics 40: 5—23.
(1955) reports insertion of three or four bands, phenotype: Variegation intermediate. Homozygote
probably of heterochromatic origin. Reverts to lethal. RK1A.
wild type when extra bands separated from bw cytology: Associated with T(2;3)bwRl$ -
locus (Hinton and GoodSmith, 1950, J. Exptl. T(2;3)59D;80C.
Zool. 114: 103-14).
few**58; brown of Meyer origin: X ray induced.
origin: Spontaneous. discoverer: Slatis.
discoverer: Meyer, 58k. references: 1955, Genetics 40: 5—23.
references: 1959, DIS 33: 97. phenotype: Variegation intermediate. Homozygotes
phenotype: Intermediate allele. Reddish brown lethal. RK1A.
eyes (like g) of somewhat reduced size. May cytology: Associated with ln(2)bwRla =
overlap wild type in old crowded cultures. Testis In(2)40F-41A;59E4-Fl.
sheath light yellow in young but dark in old OWR20
males. Good viability and fertility. RK2. origin: X ray induced.
bwMiS9: brown of Mischaikow discoverer: Slatis.
origin: Spontaneous. references: 1955, Genetics 40: 5—23.
discoverer: Mischaikow, 59e. phenotype: Variegation weak. Homozygote lethal.
reference*: 1959, DIS 33: 97. RK2A.
cytology: Associated with In(2LR)bwR2° = references: 1955, Genetics 40: 5—23.

In(2LR)40D;59D5-6. phenotype: Variegated in heterozygotes with bw + ;
%WR2S strongly variegated in homozygotes, and in
origin: X-ray-induced derivative of bw. heterozygotes with bw and other variegated
discoverer: Slatis. browns. Homozygotes show melanotic clots of
references: 1955, Genetics 40: 5—23. dried hemolymph. RK1A.
phenotype: Variegation not described. Homozygote cytology: Associated with In(2)bwR47 =
like bw. RK2A. In(2)40-41;59DU-El.
cytology: Associated with T(2;4)bwR25 =
T(2;4)59D;101E. origin: X ray induced.
discoverer: Slatis.
origin: X-ray-induced derivative of bw. references: 1955, Genetics 40: 5—23.
discoverer: Slatis. cytology: Analysis incomplete; one break at
references: 1955, Genetics 40: 5—23. 59D2-3.
phenotype: Variegation intermediate; brown in wR 55
combination with bw. RK1A. origin: X ray induced.
cytology: Associated with T(Y;2)bwR27 = discoverer: Slatis, 50d23.
T(Y;2)59D11-E1. references: 1955, Genetics 40: 5—23.
phenotype: bwRS5/+ and bwR55/bw usually show
origin: X ray induced. small amounts of variegated eye tissue; but
discoverer: Slatis. occasionally entire eye nearly lacks red pigment.
references: 1955, Genetics 40: 5-23. Homozygous lethal. RK1A.
phenotype: Variegation intermediate to slight. cytology: Associated with In(2LR)bwRSS =
Homozygote shows extreme variegation and In(2LR)24El-D;42E + In(2R)40F-41A;59D4-5.
reduced viability. RK1A.
cytology: Associated with In(2R)bwR32 =
In(2R)41A;59D. origin: X ray induced.
*by,R33 discoverer: Slatis.
origin: X ray induced. references: 1955, Genetics 40: 5—23.
discoverer: Slatis. phenotype: Variegation intermediate in heterozy-
references: 1955, Genetics 40: 5—23. gote, extreme in homozygote. RK1A.
cytology: Associated with Ia(2R)bwR33 — cytology: Associated with In(2)bwRS6 =
In(2R)41;59D'E. In(2)40F-41A; 59D-E.

discoverer: Slatis. discoverer: Slatis.
references: 1955, Genetics 40: 5—23. references: 1955, Genetics 40: 5—23.
phenotype: Variegation intermediate to slight; phenotype: Variegation weak in heterozygote.
homozygotes show more extreme variegation and RK2A.
reduced viability. RK1A. cytology: Associated with T(Y;2)bwRS7 =
cytology: Associated with In(2)bwR35 = T(Y;2)59D5-6.
In(2)4QF-41A;59Dl 1-E1.
origin: X ray induced. discoverer: Slatis,
discoverer: Slatis. references: 1955, Genetics 40: 5—23.
references: 1955, Genetics 40: 5-23. phenotype: Variegation in heterozygote weak.
phenotype: Shows slight dilution of eye pigment in Homozygote lethal. RK1A.
bwR4°f+ heterozygote. Homozygote presumably cytology: Associated with T(2;3;4)bwRS8 =
lethal. RK2A. T(2;3;4)59D;6 5;101C.
cytology: Associated with Dl(2R)bwR40 -
Df{2R)59C5~6;59E2-3. origin: X ray induced.
discoverer: Slatis.
origin: X-ray-induced derivative of bw. references: 1955, Genetics 40: 5—23.
discoverer: Slatis. phenotype: Variegation in heterozygote moderate to
references: 1955, Genetics 40: 5-23. strong. Homozygote lethal. RK1A.
pfoeno*ype: Variegation of bwR4$/+ slight. cytology: Associated with In(2)bwR67 -
Homoxygote lethal. RK2A. In(2)40F'41A;59E4-Fl.
cytology: Associated with
ln(2)4QF-41A;59E3-4. origin: X ray induced.
discoverer: Slatis.
discoverer: Slatis, 5CM16. phenotype: Like bw. RK1A.
MUTATIONS 41
cytology: Associated with complex rearrangement references: 1930, J. Genet. 22: 299-334 (fig.).
with one break near 58F; mutation and rearrange- Glass, 1934, J. Genet. 28: 69-112 (fig.).
ment presumably independently induced. 1934, Am. Naturalist 68: 107-14.
phenotype: Eye color like bwvi but with browner
origin: X ray induced. ground and numerous discrete very dark granular
discoverer: Slatis. spots; more variable. Larval Malpighian tubules
references: 1955, Genetics 40: 5-23. normal (Glass, Brehme). Homozygote generally
cytology: Associated with In(2)bwR73 = lethal. RK1A.
In(2)40F-41A;59E4-Fl. cytology: Associated with T(2;3)bwV3.
wV*
discoverer: Slatis, 50g26. discoverer: Patterson, 1929.
references: 1955, Genetics 40: 5-23. synonym: Rosy.
phenotype: Strongly variegated in combination with references: Glass, 1934, J. Genet. 28: 69-112 (fig.).
either bw or bw+. Homozygous lethal and lethal 1934, Am. Naturalist 68: 107-14.
or semilethal with most other brown-Variegateds. phenotype: Eye color yellow-brown ground with
RK1A. numerous dark granular spots. Homozygous lethal
cytology: Associated with in 95 percent of cases; survivors have pale rose-
ln(2)40F-41A;59F2-3. brown eye color with few darker spots, which
change to white and yellowish in combination with
: brown-Variegated st. Larval Malpighian tubules normal (Glass,
origin: X ray induced. Brehme). RK1A.
discoverer: Muller, 1929. cytology: Associated with T(2;3)bwV44
synonym: Pm; Plum.
references: 1930, J. Genet. 22: 299-334 (fig.). origin: X ray induced.
Glass, 1934, J. Genet. 28: 69-112 (fig.). discoverer: Patterson, 1929.
1934, Am. Naturalist 68: 107-14. synonym: 143a.
Bridges, 1937, Cytologia (Tokyo), Fujii Jub. references: Glass, 1934, J. Genet. 28: 69-112 (fig.).
Vol. 2: 745-55. 1934, Am. Naturalist 68: 107-14.
phenotype: Eye color like bw or pr, mottled with phenotype: Like bwv3 and bwv* but browner
darker spots that deepen in red color with age. ground color. Rare homozygous survivors. Larval
With sf or v, has pale orange ground with dark Malpighian tubules normal (Glass, Brehme).
orange spots. Extra Y chromosome, as with other RK1A.
variegated browns, suppresses brown color, giving cytology: Associated with T(2;3)bwV5w
red eye sparsely speckled or splotched with *bwV6
darker spots. Larval Malpighian tubules normal origin: X-ray-induced derivative of bw.
(Glass, Brehme). Generally lethal homozygous discoverer: Moore, 1929.
and in combination with other brown-Variegateds. synonym: A34.
Heterozygotes fully viable and fertile. RK1A. references: Glass, 1934, J. Genet. 28: 69-112 (fig.).
cytology: Associated with In(2LR)bwV1 = 1934, Am. Naturalist 68: 107-14.
In(2LR)21C8-Dl ;60Dl-2 + In(2LR)40F;59D4-El phenotype: Like bxvVS. RK1A.
(Schultz and Bridges). cytology: Associated with T(2;3)bwv6.
other information: Lost by reversion.
origin: X ray induced. discoverer: Winchester, 1932.

discoverer: Harris, 1929. phenotype: Eye color like bwvs. Homozygote
synonym: Discolored. moderately viable; eye color of pale rose-brown
references: Muller, 1930, J. Genet. 22: 299-334 ground with few spots. RK1A.
(fig.). cytology: Associated with In(2R)bwv7\
Glass, 1934, J. Genet. 28: 69-112 (fig.).
1934, Am. Naturalist 68: 107-14. origin: X ray induced,
phenotype: Eye color similar to bwvl, but less discoverer: Levy, 1932.
purplish, v, bwv2 has yellower ground color and references: 1935, DIS 3: 7.
browner spots than bwvi; st. Homozygote nearly phenotype: Like bwV^. RK1A.
always lethal. Malpighian tubules of larvae cytology: Associated with T(2;3)bwV8.
normal (Glass, Brehme). RK1A.
cytology: Not studied. Shown genetically to be origin: Radium induced.
associated witn In(2R)bwv'2. discoverer: Van Atta, 291.
synonym: Cream; Dilute-1.
origin: X ray induced, references: Hanson and Winkelman, 1929,
discoverer: Muller, 1929. J. Heredity 20: 277-86.
synonym: Tarnished. Van Atta, 1932, Genetics 17: 637-59.
phenotype: Like bwVl, RK1A. except in the presence of an extra Y.

cytology: Associated with In(2LR)bv/V29l, shows variegation for light (Schultz, 1936). Via-
*bwV30a bility excellent. Homozygous lethal except for
origin: X ray induced. rare survivors with light eye color and somatic
discoverer: Oliver, 30a. abnormalities. Larval Malpighian tubules some-
phenotype: Almost homogeneous brown eye color. what lighter than wild type but not useful in
RK3(A). classification (Brehme and Demerec, 1942, Growth
bwV30k1 6: 351-56). RK1A.
origin: X ray induced. cytology: Associated with In(2LR)bwv32g =
discoverer: Van Atta, 30kl. In(2LR)40F;59E.
synonym: Dilute-2.
references: 1932, Genetics 17: 637-59. origin: X ray induced in In(2L)Cy +In(2R)Cy.
phenotype: Like bwVl. RK1A. discoverer: Oliver, 34k22.
cytology: Associated with In(2LR)bwV30k*-. synonym: Var34k22,
bwV3Okl0 references: 1937, DIS 7: 19.
origin: X ray induced. phenotype: Eye color nearly homogeneous brown,
discoverer: Van Atta, 30kl0. slightly mottled. Larval Malpighian tubules
synonym: Dilute-3. normal (Glass, Brehme). RK1A.
references: 1932, Genetics 17: 637-59. cytology: Associated with In(2R)bwv34k =
phenotype: Eye color blotched heavily with large In(2R)41;59E, which carries as an included
patches of red and brown. Homozygous lethal. inversion In(2R)Cy = In(2R)42A2-3;58A4-Bl.
RK1A. *bwV40b
cytology: Associated with In(2R)bwV3Okio% origin: X ray induced.
*bwV30kl2 discoverer: T. Hint on, 40b.
origin: X ray induced. references: Atwood, 1942, DIS 16: 47.
discoverer: Van Atta, 30kl2. phenotype: Eye light brown with some darker
synonym: Dilute-4. variegation. Homozygous lethal. RK1A.
references: 1932, Genetics 17: 637-59. cytology: Associated with ln(2R)bwv40b =
phenotype: Eye color mostly red with a sprinkling In(2R)41A-B;59D-E.
of spots and facets of brown, appearing as dark
spots. Homozygous lethal. RK2A. origin: Gamma ray induced.
cytology: Associated with T(2;3)bwV30kl2, discoverer: Mickey, 54a6.
origin: X ray induced. phenotype: Variegated for bw. RK1A.
discoverer: Van Atta, 30kl3. cytology: Associated with In(2R)bwv54a =
synonym: Dilute-5. Jn(2R)41A-B;59D4-9.
references: 1932, Genetics 17: 637-59. *bwV54b
phenotype: Eye color mostly red with brown discoverer: Mickey, 54bl2.
spotting. Homozygous lethal. RK2A. references: 1963, DIS 38: 29.
cytology: Associated with T(2;3)bwV30kl3, phenotype: Variegated for bw. RK1A.
cytology: Associated with In(2R)bwv54t> =
origin: X ray induced. In(2R)41A;60D9-ll.
discoverer: Van Atta, 30kl8. *bwV54c
synonym: Dilute-6. origin: Neutron induced.
references: 1932, Genetics 17: 637-59. discoverer: Yanders, 54c5.
phenotype: Eye color mostly red with sprinkling of references: Mickey, 1963, DIS 38: 29.
dark (brown) facets. Homozygous lethal. RK2A. phenotype: Variegated for bw. RK1A.
cytology: Associated with T(2;3;4)bwV30kl8w cytology: Associated with In(2R)bwVS4c =
In(2R)41;59El.
discoverer: Dobzhansky, 32g6. origin: X ray induced in In(2LR)SMl, al2 Cy en2 sp2.
2
synonym: Pm . discoverer: E. H. Grell, 57e.
references: Schultz and Dobzhansky, 1934, references: Hochman, 1961, DIS 35: 85—86.
Genetics 19: 344-64. Welshons, 1962, Genetics 47: 743-59.
Schultz, 1936, Proc. Natl. Acad. Sci. U.S. 22: phenotype: Eyes brown and mottled. Viability of
27-33. bwvS7e/bwvl about 40 percent at 26°C and 20
phenotype: Dominant eye color like brown but percent at 23.5°; body tends to be small, wings
flecked with darker spots. Shows 18 percent divergent and often not expanded; patches of un-
normal red pigment and 88 percent normal brown pigmented microchaetae. RK2A.
pigment (Nolte, 1954, J. Genet. 52: 127-39). cytology: Salivary chromosomes not examined, but
bwv32t/bw is like bw with very few spots. likely that bwVS7e is the result of a rearrangement
t,wV32g/+ jS easily separable from wild type superimposed on SMI.
MUTATIONS 43
: brown-Variegated Dichaete linked phenotype: Like bw in young flies but much darker
origin: X ray induced. than bw in old individuals. RK1.
discoverer: Oliver, 29k24. other information: Apparently never tested for
synonym: Ic D. allelism with ca (3-100.7).
references: 1932, Z. Induktive Abstammungs- bw-l: see red
Vererbungslehre 61: 447-88. *bwl: bow-legged
phenotype: Like bwvl, but redder and more variable. location: 1-21.9.
Homozygous lethal, but not lethal in combination origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
with other brown-Variegateds. RK2A. phenylalanine (CB. 3025).
cytology: Associated with T(2;3)bwVE>% discoverer: Fahmy, 1955.
other information: Irradiated third chromosome references: 1959, DIS 33: 83.
carried D. phenotype: Poorly viable with shorter divergent
bwVDe I; brown-Variegated of Demerec wings. Bristles thinner and shorter. Legs
origin: X ray induced. shortened and either femur or tibia, or both, bow-
discoverer: Demerec, 33i28. shaped. Males sterile. RK3.
references: Bridges, 1937, Cytologia (Tokyo),
Fujii Jub. Vol. 2: 745-55.
phenotype: Eye color variegated brown and red;
extreme dominant brown with pebbled wing.
Lethal when homozygous and in combination
with bw*. RK2A.
cytology: Associated with In(2R)bwVêl -
In(2R)41 B2-Cl;59E2-4.

discoverer: Demerec, 33J14.
references: Bridges, 1937, Cytologia (Tokyo),
Fujii Jub. Vol. 2: 745-55.
phenotype: Eye color mosaic of brown and dark
brown patches. RK2A.
cytology: Associated with In.(2R)bwVDf>2 -
In(2R)41A-B;59D6-El.

discoverer: Demerec, 33jl4.
phenotype: Eye color variegated brown and red. In bx: bithorax
bwVDe3/bw5, wings fail to expand; joints and From Bridges and Morgan, 1923, Carnegie Inst. Wash.
sutures weak with melanotic clots. Homozygous Pubi. No. 327: 152.
lethal. RK2A.
cytology: Associated with T(2;3)bwvE>e3 = bx: bithorax
T(2;3)59D;81F. location: 3-58.8.
discoverer: Bridges, 15i22.
origin: X ray induced. references: Bridges and Morgan, 1923, Carnegie
discoverer: Demerec, 33k22. Inst. Wash. Publ. No. 327: 137, 152 (fig.).
phenotype: Eye color red with brown s p o t s ; Morgan, Bridges, and Sturtevant, 1925, Bibliog.
bwVDe4/bw and X/0; bwVDe4/+ a r e brown. Genet. 2: 79, 214 (fig.).
Homozygous lethal. RK2A. Lewis, 1951, Cold Spring Harbor Symp. Quant.
cytology: Associated with T(2;3)bwyDe4 - Biol. 16: 159-74 (fig.).
T(2;3)59D2~4;80. 1963, Am. Zoologist 3: 33-56 (fig.).
*bwVh brown-Variegated of Ives phenotype: Anterior half of metathorax becomes
origin: Spontaneous. mesothoracic and posterior half remains un-
discoverer: Ives, 38113. changed. This results in the appearance, between
references: 1950, DIS 24: 58. the scutellum and the first abdominal segment, of
phenotype: Like bwD. RKlA. rudimentary anterior mesothoracic elements (i.e.,
cytology: Associated with ln(2R)bwvl - mesonotum and scutellum), with the proper bristles
In(2R)41A;59D (T. Hinton). and hairs. Balancers directed ventrally, enlarged,
*bw-b: brown-b vesiculate or winglike, with typical but rudimen-
location: 3- (between 97.0 and 104.2). tary wing venation and bristles. Metathoracic
origin: X ray induced. tibia has a mesothoracic tibial bristle. Variable,
discoverer: E. L. Smith, 34f. overlapping wild type at 25° and more so at 19°C.
references: Robertson, 1935, DIS 4: 15. Dorsal metathoracic disk of mature larva is 60 per-
Smith and Robertson, 1938, Genetics 23: 167. cent larger than wild type IChen, 1929, J. Morphol.
47: 135—99 (fig.)]. Thoracic musculature studied

by El Shatoury [l956, J. Embryol. Exptl. Morphol. origin: Spontaneous.
4: 228—39 (fig.)] and Pantelouris and Waddington discoverer: Gunson.
[l955, Arch. Entwicklungsmech. Organ. 147: 539— references: 1952, DIS 26: 63.
46 (fig.)]. Bristle patterns studied by Waddington phenotype: Penetrance 0.5 percent at 16°, 1 percent
(1962, New Patterns in Genetics and Development. at 20°, and 29 percent at 25°C. RK3.
Columbia University Press). Slight expression in bxD>: see Ubx
combination with bw34e and bww (Hollander, 1937, *bxw: bithorax-W/scons/n
DIS 8: 77), and no interaction as trans heterozy- origin: Spontaneous; recovered in one third chromo-
gote with bxd or Vbx (E. B. Lewis). RK3. some of the female in whose other third chromo-
cytology: Located close to if not within the 89E1-2 some Ubx was first recovered.
doublet (Lewis, 1951). discoverer: Hollander, 1934.
other information: The leftmost member of a pseudo- references: 1937, DIS 8: 8, 77.
allelic series including from left to right, bx, Cbx, phenotype: Metathorax developed more than any
Ubx, bxd, and pbx. other bx allele and strongly bristled. Halteres
directed ventrally, large, flat, and winglike, with
veins and bristles. Mesothorax shows clear stripe
or bifida condition. Wings usually spread, often
bx Cbx Ubx bxd pbx dragging. Sternopleural and tibial and ventral
bristles of third leg resemble those of normal
0.01 0.008 0.006 0.006 second leg. bxw/bxd = +. Viability and fertility
fair. RK2.
fix; Beadex
Map of the bx region location: 1-59.4.
From Lewis, 1963, Am. Zoologist 3: 33-56. origin: Spontaneous.
discoverer: Bridges, 23a3.
origin: Spontaneous. Bibliog. Genet. 2: 219.
discoverer: Stern, 25b2. Green, 1953, Genetics 38: 91-105 (fig.).
references: 1935, DIS 3: 29. phenotype: Male and homozygous female with
Lewis, 1951, Cold Spring Harbor Symp. Quant. Beaded-like wings; long and narrow and excised
Biol. 16: 159-74 (fig.). along both margins. Male and homozygous female
1955, Am. Naturalist 89: 73-89. fully viable. Heterozygous female less extreme
1963, Am. Zoologist 3: 33-56 (fig.). and overlaps wild type. Some venation abnor-
phenotype: Extreme expression of 6x metathoracic mality. Development studied by Goldschmidt
effect. Little variability. Viability low but can be [1935, Biol. Zentr. 55: 535-54 (fig.)]. According
maintained homozygous. Balancers enlarged in to Waddington (1940), embryological effect is same
heterozygote. Interactions with other bx pseudo- as that of vg. RK2 (RK3 as Sx/+).
alleles described by Lewis (1951, 1955, 1963). Al- cytology: Locus in salivary region 17A, B, or C,
most completely suppressed by su(Hw)2. RK3. based on limits of Dp(l;l)Bxr49k (E. B. Lewis).
Salivary chromosomes of Bx normal. (Lewis).
discoverer: Schultz, 34e20.
Lewis, 1951, Cold Spring Harbor Symp. Quant.
Biol. 16: 159-74.
1955, Am. Naturalist 89: 73-89.
1963, Am. Zoologist 3: 33-56 (fig.).
phenotype: Metathoracic outpushing is a uniform,
narrow hairy band. Balancers depressed, inflated, Bxh Beadex-2
with elongated pointed tip and heavy line of costal Edith M. Wallace, unpublished.
bristles. Base of third leg shows sternopleural
bristles like those of normal second leg. Meta-
thoracic development of bx34e/bx340 > origin: Spontaneous.
bx3«»/bxw> bx34°/bx (Hollander, 1937, DIS 8: discoverer: Mohr, 24129.
77). bx^4e/Ubx has round, flat, winglike references: 1927, Nyt Mag. Natur 65: 265—74.
halteres; bx^4e i/bx/+ + = Ubx/+ (Lewis). Comple- Green, 1953, Z. Induktive Abstammungs-
ments hxd. Reasonable viability and fecundity. Verebungslehre 85: 435—49.
Highly constant expression and easy separability. phenotype: Wings of males and homozygous females
Expression increased slightly at 29°C, decreased narrowed by marginal excision. Wings often bubbly
at 15°C (Villee, 1943, Anat. Record 87: 475). RK2. and ragged. Homozygous female fully viable.
MUTATIONS 45
2
Bx /+ less extreme; overlaps wild type. Classi- other information: Bx/Dp(l;l)Bxr produces recom-
fiable in a single dose in triploids (Schultz, 1934, binants of genotype Bx+Bx and BxBx+, which are
DIS 1: 55). RK1 (RK3 as Bx2/+). more extreme than Bx. Same holds for
Bx2/Dp(l;l)Bx*.
discoverer: Gershenson, 1927.
references: Gaissinovitsch and Gershenson, 1928,
Biol. Zentr. 48: 385-87 (fig.).
phenotype: Extreme allele usually without the
bubbles in the wing. Shortened L5 a constant
character (few Bx2 show this). Wings more pointed
than Bx2 and hairs at tip of wing clumped.
Scalloping visible in prepupal wing bud
[Waddington, 1940, J. Genet. 41: 75-139 (fig)],
fully separable. RK1. Bx^9k: Beadex-recessive 49k
From Green, 1953, Z. Induktive Abstammungs-
origin: Spontaneous. Vererbungslehre 85: 435-49.
discoverer. T. J. Lee, 59h.
phenotype: Like Bx. RK2. origin: Spontaneous.
discoverer: Mossige, 49k22.
*Bxc: Beadex of Catcheside synonym: Bxl2.
discoverer: Catcheside, 39c3. Green, 1953, Z. Induktive Abstammungs-
references: 1939, DIS 12: 49. Vererbungslehre 85: 435-49 (fig.).
phenotype: Posterior wing margin excised as far as phenotype: Slight scalloping of posterior wing
and including most of L5; end of wing notched and margin only; overlaps wild type. RK3A.
anterior margin weakly excised. Wings blistered, cytology: Associated with Dp(l;l)BxrJ*9k -
especially basally. BxC/+ is like Bx^/Y. RK1. Dp(l;l)17A;17C (E. B. Lewis).
other information: This duplication undergoes
Bx-I: Beadex of Jollos unequal crossing over readily and forms triplica-
origin: Induced by heat treatment. tions and quadruplications. Duplication is reces-
discoverer: Jollos, 1930. sive; triplication is dominant. Phenotypic interac-
synonym: Ptd: Pointedoid. tion with Bx same as for BxT.
references: 1933, Naturwissenschaften 21: 831—34.
Gottschewski, 1935, DIS 4: 7, 14, 16.
Jollos and Waletsky, 1937, DIS 8: 9.
phenotype: Wings reduced to slender strip; only
posterior cell present at tip. Femur shortened or
legs otherwise abnormal, especially third pair.
Homozygous female viable. Interacts with bi to
give more nearly normal wings. Embryology like
Bx [Goldschmidt, 1935, Biol. Zentr. 55: 535-54;
Waddington, 1940, J. Genet. 41: 75-139 (fig.)].
RK1.
Bxr: Beadex-recessive
discoverer: Ives, 35k.
Green, 1952, Proc. Natl. Acad. Sci. U.S. 38:
949-53.
1953, Genetics 38: 91-105 (fig.).
phenotype: Bxr/+ is normal. Male and homozygous
female show less extreme narrowing of wings than hxd: hithoraxoid
Bx. Anterior crossvein short and thickened and From Bridges and Morgan, 1923, Carnegie Inst. Wash.
that region blistered. May overlap wild type in Publ. No. 327: 225.
old crowded cultures at 25°, more extreme at 19°C.
RK3A. bxd: bithoraxoid
cytology: Associated with Dp(t;l)Bxr ~ location: 3-58.8.
Dp(l;l)17A;17E-F (Green, 1953, determined by origin: Spontaneous.
E. B. Lewis). discoverer: Bridges, 19127.
references: Bridges and Morgan, 1923, Carnegie references: 1945, Univ. Calif. (Berkeley) Publ.
Inst. Wash. Publ. No. 327: 225 (fig.). Zool. 49: 507, 520.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. phenotype: Like a strong Bx. RK2.
Genet. 2: 214 (fig.), 219.
Lewis, 1951, Cold Spring Harbor Symp. Quant. Bxl: see Ubx
Biol. 16: 159-74 (fig.). by: blistery
1955, Am. Naturalist 89: 73-89. location: 3-48.7.
1963, Am. Zoologist 3: 33-56 (fig.). origin: Spontaneous.
phenotype: Posterior portion of metathorax becomes discoverer: Glass, 33a.
mesothoracic; anterior metathorax unaffected. references: 1934, DIS 2: 8.
Enlarged metathoracic postnotum forms two phenotype: Wings blistered in subterminal region,
wedges of tissue, devoid of hairs and bristles, and wing surface dusky and warped. Thorax
meeting at the mid-dorsal line. Balancers enlarged humpy. RK1.
into circular disks that are cupped and bent down- *by46h
ward and show venation but are bare of bristles origin: Spontaneous.
except for weak ones at rear margin. Stalk of discoverer: Ives, 46hl5.
balancer wide and flat. First abdominal segment references: 1948, DIS 22: 53.
shows thoracic modification (modification of phenotype: Like by but without thoracic effect.
anterior portion is metathoracic-like, posterior RK1.
portion is partially mesothoracic-like). Rudiments hi: see malbz
of a first pair of abdominal legs and, in certain
combinations, of partially wing-like first abdominal
halteres may occur. Interactions with other bi-
thorax pseudoalleles described by Lewis (1951,
1955, 1963). RIG.
cytology: Locus probably 89E3-4 (E. B. Lewis).

references: 1951, Cold Spring Harbor Symp. Quant.
Biol. 16: 159-74.
phenotype: More extreme bithoraxoid phenotype
than bxd. RK2A.
cytology: Associated with Tp(3)bxdi00 -
Tp(3)66C;89B5-6;89E2'3. bx+ but not bxd included
in transposed section.
origin: X ray induced in ss bx Su(ss)2.

discoverer: E. B. Lewis,
phenotype: Like hxd*O°. RK2A.
cytology: Associated with T(3;4)89E = T(3;4)89E2-
3;101F. c; curved
From Bridges and Morgan, 1919, Carnegie Inst. Wash.
origin: X ray induced in pP bx sr ea. Pubi. No. 278: 165.
phenotype: Like bxd10®. RK2A. c: curved
cytology: Associated with Tp(3)bxd11(> = Tp(3)89E2- location: 2-75.5.
3;91C7-Dl;92A2-3. origin: Spontaneous.
discoverer: Bridges, 11124.
discoverer: E. B. Lewis. Inst. Wash. Publ. No. 278: 164 (fig.)-
references: 1951, Cold Spring Harbor Symp. Quant. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
Biol. 16: 159-74. Genet. 2: 211 (fig.).
1963, Am. Zoologist 3: 33-56 (fig.). phenotype: Wings thin textured, divergent, uplifted
phenotype: hxdm/bxd shows infrequent develop- at base, and curved downward throughout their
ment of abdominal wing-like halteres as well as length. RK1.
abdominal legs. RK2. *C"K: Curved of Krivshenko
hxd*>: see Vbx location: 2- or 3- (rearrangement).
*Bxd: Beadnxoid origin: X ray induced,
location: 1-45. discoverer: Krivshenko, 5513.
origins Spontaneous. references: 1956, DIS 3th 74.
dlscov»r«r: Goldschmidt. synonym: CK.
MUTATIONS 47
phenotype: Wings are thin textured, slightly diver- CHL: see In(2L)NS
gent, uplifted basally, and then curved downward. CIIIRE: see In(3R)C
Homozygous lethal. RK2A. co: claret
cytology: Associated with T(2;3)C-K = location: 3-100.7.
T(2;3)52;76;81;86. origin: Spontaneous.
C( ): Crossover suppressor discoverer: Bridges, 19112.
The terminology originally used for dominant sup- references: Bridges and Morgan, 1923, Carnegie
pressors of crossing over. These effects were Inst. Wash. Publ. No. 327: 219 (fig.).
found to be rearrangements and are so treated here. phenotype: Eye color ruby. Red and brown pig-
The symbol C in this context has been dropped ments 27 and 29 percent of normal levels, respec-
except where included under synonymy. tively (Nolte, 1955, J. Genet. 53: 1-10). With en,
*c(l)a: recessive crossover suppressor for eye color is deep reddish yellow; with bw, trans-
chromosome 1 lucent brownish yellow (Mainx, 1938, Z. Induktive
location: One factor in X and probably several Abstammungs- Vererbungslehre 75: 256—76).
autosomal modifiers, Larval Malpighian tubes colorless (Beadle, 1937,
origin: Spontaneous. Genetics 22: 587—611). Eye color autonomous
discoverer: Bridges, 1916. when larval optic disk from ca is transplanted into
references: Bonnier, 1923, Hereditas 4: 81—110. wild type or v. Wild-type disk in ca not entirely
Morgan, Bridges, and Sturtevant, 1925, Bibliog. autonomous (Beadle and Ephrussi, 1936, Genetics
Genet. 2: 220. 21: 230); ca flies produce less v + substance than
phenotype: Reduces recombination between v and f wild type (Clancy, 1942, Genetics 27: 417-40).
from 23 to 15 percent and between w° and v from Slightly narrow body and pointed wing. RK1.
31 to 10 percent. c(l)a was probably the cause of cytology: Judged to be in 99C-E, based on
a secondary nondisjunction frequency of 15—30 In(3R)cav = In(3R)81F;99C-B (E. B. Lewis).
percent. RK3. co2
other information: Validity of phenotypic descrip- origin: Spontaneous in In(3R)P = In(3R)89C2-
tion seems dubious. 3; 96A18-19.
C(2)R: see In(2R)NS discoverer: Bridges, 32f22.
C(2;3): see In(2L)t phenotype: Eye color like ca. Same body and wing
C(2L)HR: see In(2L)t effect as ca. Larval Malpighian tubes colorless
C(2L)T: see ln(2L)t (Brehme and Demerec, 1942, Growth 6: 351-56).
c(3)G: recessive crossover suppressor in RK1A.
chromosome 3 of Go wen ; clarei-nondisjunctional
location: 3-57.4 (1.0 to the left of sbd2, 4.0 to the origin: X ray induced.
right of cv-c). discoverer: E. B. Lewis and Gencarella.
origin: Spontaneous, references: 1952, Genetics 37: 600—1.
discoverer: Gowen and Gowen, 1917. phenotype: Eye color like ca, but homozygous
synonym: ex (Gowen, 1928, Proc. Natl. Acad. Sci. females lay mostly inviable eggs; surviving
U.S. 14: 475-77). progeny often show results of chromosome nondis-
references: 1922, Am. Naturalist 56: 286-88. junction and loss; meiosis normal in cand/ca
1932, Proc. Intern. Congr. Genet., 6th. Vol. 2: females. Similar in action to ca of Dzosophila
69-70. simulans (Sturtevant, 1929, Z. Wiss. Zool. Abt. A
1933, J. Exptl. Zool. 65: 83-106. 135: 323—56). One experiment yielded 1373
phenotype: In homozygous females, crossing over in progeny of females homozygous for cand, and gave
entire chromosome complement reduced to a small the following results: 42 percent regular offspring,
fraction of normal. Production of triploids and in- 15.0 percent exceptional (X/X/Y and X/0), 32.0
tersexes 300—500 times normal. Nondisjunction in- percent haplo-4, 3.9 percent haplo-4 mosaics, and
creased; egg hatching very low, probably owing to 5.4 percent gynandromorphs. The remaining 0.8
aneuploid zygotes. Core structures characteristic percent included a triploid female, triploid inter-
of electron microscope preparations of normal sexes, probable superaales, and a few other kinds
oocyte nuclei absent in c(3)G/c(3)G oocytes of mosaics. Causes nondisjunction and loss of
(Meyer, 1964, Proc. Eur. Reg. Conf. Electron. major autosomes (Davis, D. G., 1963, Ph.D.
Microscop. 3rd, pp. 461—62). Somatic crossing over Thesis, Univ. Georgia). Hinton and McEarchen
normal (Le Clerc, 1946, Science 103: 553-54). In- (1963, DIS 37: 90) reported haploid-diploid
creased recombination observed in c(3)G/+ females mosaic. Crossing over between X chromosomes
(Hinton, 1962, Genetics 47: 959; 1966, Genetics normal (Davis). cand ovaries transplanted into
53: 157—64). Meiosis in males not affected. RK3. normal host behave autonomously (Roberts, 1962,
cytology: Included within Df(3R)sbd*05 - DIS 36: 112). Chromosome segregation normal in
D((3R)88F9-89Al;89B4-5 (Lewis, 1948, DIS 22: cand males. RK3.
72-73). *: claret-variegated
C2L: see ln(2L)NS origin: X ray induced.
C3: see In(3R)C discoverer: E. B. Lewis.
phenotype: cav/ca slightly variegated. Can be *Cb: Curled blistered

confused with wild type. cav/cand females pro- location: 1-13.
duce normal progeny- Homozygous lethal. RK3A. origin: Spontaneous.
cytology: Associated with In(3R)cav = discoverer: Villee, 40b.
In(3R)81F;99C-E. references: 1945, DIS 19: 47.
*cal: coal phenotype: Heterozygous or homozygous Cb give
location: 3-59.5. curled and blistered wings only in presence of
origin: Spontaneous. homozygous pxCb. RK3.
discoverer: Grout, 47120. *cbd: cluboid
references: Ives, 1948, DIS 22: 53. location: 3- (about 30 units to right or left of D).
phenotype: Black body color similar to e4. Via- origin: Spontaneous.
bility reduced slightly. RK2. discoverer: Bridges, 16il5.
canopy wing: see cpw references: Bridges and Morgan, 1923, Carnegie
Inst. Wash. Publ. No. 327: 169.
car: carnation phenotype: Wings not expanded. Dwarfish. Low
location: 1-62.5. fertility. RK3.
origin: X ray induced. *cbf: clubfoot
discoverer: Patterson, 28c20. location: 1-45.
references: 1934, DIS 1: 31. origin: X ray induced.
phenotype: Eye color dark ruby. Body shape and discoverer: Cantor, 46d20.
proportions seem rounded. With st, eye color is references: 1946, DIS 20: 64.
yellow-brown, with bw, brownish yellow to brown phenotype: Leg segments greatly shortened; ab-
(Mainx, 1938, Z. Induktive Abstammungs- normally shaped tarsi and metathoracic legs.
Vererbungslehre 75: 256—76). Maipighian tubes Wings slightly warped, wide in center, and
pale yellow in mature larva (Beadle, 1937, tapering at ends. All flies emerging show both
Genetics 22: 587—611) but hard to distinguish wing and leg effects but expression variable.
from wild type before third instar. Eye color Only about 3 percent of cbt flies eclose. RK3.
autonomous in transplant into wild-type host other information: Not tested for allelism to pi
(Beadle and Ephrussi, 1936, Genetics 21: 230). (1-47.9).
Contains 33 percent wild-type red pigment and 47 Cbx: Contrabithorax
percent brown pigment (Nolte, 1959, Heredity 13: location: 3-58.8 (to the right of bx; to the left of
233-41). RK1. Ubx).
cytology: Shown to lie in doublet 18D1-2 by defi- origin: X ray induced. Arose simultaneously with
ciency analysis (J. I. Valencia). pbx.
discoverer: Bacon, 49h.
discoverer: Nolte, 1952. references: Lewis, 1954, DIS 28: 76.
references: 1954, DIS 28: 77. 1954, Proc. Intern. Congr. Genet., 9th. 1: 100-5.
phenotype: Visually resembles car, but contains 1955, Am. Naturalist 89: 73-89.
only one-half the amount of red pigment of car, 16 1963, Am. Zoologist 3: 35—56 (fig.).
percent of wild-type red pigment, and 48 percent phenotype: Cbx/+ and Cbx/Cbx virtually indistin-
of wild-type brown pigment (1959, Heredity 13: guishable and have small wings. The posterior
233-41). RK1. portion of the mesothorax transformed into a struc-
CQr26-48 ture that resembles posterior portion of meta-
origin: Induced by mustard gas. thorax. Wings have veins LI, L2, and L3 only. In
discoverer: Sobels and Jansen, 571. extreme cases, entire mesothorax resembles meta-
references: Sobels, 1958, DIS 32: 84. thorax so that wings resemble halteres. Meta-
phenotype: Eye color darker than car and tends to thorax is wild type. Cbx acts as dominant sup-
resemble pn more than car. RK1. pressor of homozygous pbx and partial suppressor
cardinal: see ccf of bx and bxd. Interactions with other bx pseudo-
carmine: see cm alleles described by Lewis (1955, 1963). RK2.
carnation: see car cytology: Salivary chromosomes normal. Located
Cat: see spaCat close to if not within the 89E1-2 doublet (Lewis).
*cb: club other information: A member of the bithorax pseudo-
location: 1-16.5. allelic complex (see bx).
origin: Spontaneous. *cc; chlorotic
discoverer: Morgan, 13e. location: 1-0.0 (0.1 to left of sc).
references: Morgan and Bridges, 1916, Carnegie origin: Spontaneous.
Inst. Wash. Publ. No. 237: 69 (fig.). discoverer: Mohr, 19jl8.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. references: 1923, Studia Mendeliana (Brunae):
Genet. 2: 78 (fig.), 266-87.
phenotype: Wings club-like in about half the flies. phenotype: Body color greenish yellow. Flies
Sternopleural bristles absent from all flies. RK3. small. Mortality 90 percent. RK3.
MUTATIONS 49
*CCY: chlorotic-yellowish discoverer: Glass, 39a28.

discoverer: Morgan, 21i. phenotype: Varies from almost complete fusion of
references: 1929, Carnegie Inst. Wash. Publ. No. veins L3 and L4 to wild type. Intermediate types
399: 190. have narrowing and closing of first posterior wing
phenotype: Body color pale yellow. Viability and cell with extra veins in region of anterior cross-
fertility low. Like cc. RK3. vein and deltas at ends of L3 and L4. Expression
ccw: concave wing is better above 25°C. Homozygous lethal. RK3.
locotion: 1-23.4., cytology: Placed in salivary chromosome region
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- 101E through 102B16, based on inclusion of Ce^
phenylalanine (CB. 3025). within Dt(4)M = Df(4)101E-F;102B6-17.
phenotype: Wings shorter and narrower, with L3 and discoverer: Green.
L4 shifted toward each other; occasionally trun- references: 1952, DIS 26: 63.
cated. Wing membrane depressed in center into phenotype: Ocelli reduced or absent; ocellar and
slight concavity, giving slight scooped effect. scutellar bristles absent; wing veins L3 and L4
Not easily classified. RK3. converge, giving wing phenotype much like fa,
other information: One allele induced by CB. 3025. although wing phenotype variable. Homozygous
cd: cardinal lethal; lethality occurs during embryonic period
location: 3-75.7. (Hochman). RK3.
origin: Spontaneous. other information: Allelism based on phenotype and
discoverer: Johnson, 19k24. lethal interaction with Df(4)M (Hochman).
references: Bridges and Morgan, 1923, Carnegie *Ce3
Inst. Wash. Publ. No. 327: 217 (fig.). origin: X ray induced.
phenotype: Eye color yellowish vermilion, discoverer: Green, 59cll.
changing toward wild type with age. Brown pig- references: 1959, DIS 33: 94.
ment 15 percent of normal (Nolte, 1954, J. Genet. phenotype: Identical to Ce2. RK3.
52: 111—26). Ocelli white, showing no effect of *cf; cleft
age. Eye color autonomous in transplant of larval location: 1-65.6.
optic disk into wild type, ca, en, st, or v larval origin: Spontaneous.
host (Beadle and Ephrussi, 1936, Genetics 21: discoverer: Bridges, 14J28.
230). Larval Malpighian tubes bright yellow; not references: Morgan, Bridges and Sturtevant, 1925,
distinguishable from wild type. RK2. Bibliog. Genet. 2: 55 (fig.).
*cc/3 phenotype: Wings smaller and somewhat spread.
origin: Spontaneous. L3 split just beyond first crossvein; extra cross-
discoverer: Ives, 32c30. veins and branches. Gap in L4 beyond second
references: Plough and Ives, 1934, DIS 1: 34. crossvein. Males sterile.. Viability good. RK2.
1935, Genetics 20: 42-69. *Cf: Confluent
phenotype: Like cd. RK2. location: 2- (not located).
cd63 origin: Spontaneous.
origin: Spontaneous. discoverer: Bridges, 14i23.
discoverer: Clancy, 63a. references: Bridges, 1916, Genetics 1: 151.
references: 1964, DIS 39: 65. Bridges and Morgan, 1919, Carnegie Inst. Wash.
phenotype: Like cd. RK2. Publ. No. 278: 255.
*Cd: Coildex phenotype: Veins thickened and knotted, especially
location: 2-54.6 (0.1 unit to the right of pr). L2 opposite anterior crossvein and at costa. Wing
origin: X ray induced. smaller. Low fertility in heterozygote; good
discoverer: Bateman, 1954. viability. Homozygous lethal. RK2.
synonym: Coiled. other information: May have been a Plexate.
phenotype: Similar to Cy, but wing curvature more Cf-3: see Die(-3
extreme; wings opaque and greyish. Anterior eg: comb gap
margin of wing invaginated at point where LI location: 2-71.1.
meets wing margin. When expression is weakest it origin: Spontaneous.
appears only as a slight wave in the wing margin. discoverer: Bridges, 25kl6.
In 10—15 percent of the flies, wings also curve phenotype: Sex combs of male extremely large.
downward over flanks before curling upward. In y; Some distortion and shortening of legs. Wings
Cd flies, curvature reduced to a shallow spoon. show gap in vein L4 between posterior crossvein
Cd epistatic to Cy. Homozygous lethal. RK2. and margin. Wings slightly curved. Effects result
*Ce: Cell from a combination of overgrowth and irregular
location: 4- (not located). folding of imaginal rudiments during the pupal
period. Strong exaggeration in compound hotnozy- phenotype: Wings short, papery, and downcurved
gotes with genes such as d, fj, ds, and ss fl . with short, broad alulae. Males sterile. Homozy-
Double heterozygote for eg and ci often shows gap gotes short lived and balanced stock
in L4 (Waddington, 1952, J. Genet. 51: 243-58). en che bw sp/In(2L)Cy + In(2R)Cy,
Double heterozygote en cg/++has slight degree of al% Cy en2 L4 sp? has a generation time 30 per-
L4 interruption and thinning at low temperature. cent longer than normal. RK3.
Triple heterozygote en cg/++; ci/+ has L4 inter- cfte*
ruption in half the flies (House, 1961, Genetics 46: origin: Ultraviolet induced.
871). ciw interacts strongly with eg. cg/+; ciw/+ discoverer: Meyer and Edmondson, 481.
resembles ciw/ciw (House, 1953, Genetics, 38: references: 1951, DIS 25: 71.
669—70). Females sterile. Oogenesis highly irreg- phenotype: Similar to cfte in wing characteristics
ular (Beatty, 1949, Proc. Roy. Soc. Edingurgh and male sterility. RK3.
B 63: 249-70). RK2.
eft: chubby
Left: wild-type larva. Right: chubby larva.
From Dobzhansky and Duncan, 1933, Arch. Entwicks-
lungmech. Organ. 130: 109-30.
*cfte3
eg; comb gap discoverer: Meyer, 60g.
From Bridges and Brehme, 1944, Carnegie Inst. Wash. references: 1963, DIS 37: 50.
Publ. No. 552: 40. phenotype: Wings curved, spread, and short. Less
extreme than che. che3/che sterile in both sexes.
eft: chubby che3/che3 not tested for sterility because fs(2)B
location: 2-72.5. and ms(2)2 on same chromosome. RK2.
origin: Spontaneous. chilblained-b: see cft-£>
discoverer: Bridges, 17J26. eft/: chaetelle
references: Morgan, Bridges, and Sturtevant, 1925, location: 2-60.8.
Bibliog. Genet. 2: 222. discoverer: Bridges, 33a4.
phenotype: Adults, pupae, and larvae thickset and references: Beatty, 1949, Proc. Roy. Soc.
short. Difficult to distinguish from wild type. Edinburgh B 63: 249-70.
Chubby larvae shorter than wild type at hatching phenotype: Bristles very small. Wing venation
[Dobzhansky and Duncan, 1933, Arch. Entwick- slightly plexus-like; exaggerates px when com-
lungsmech. Organ. 130: 109-30 (fig.)]. RK3. bined with it. Body size small. Rotated genitalia
*ch-b: chilblained-b in many males. Blunt-tipped abdomen. Females
location: 1-23.8. infertile, but ovary and oocytes appear normal.
discoverer: Moriwaki, 39e22. RK2.
references: 1939, DIS 12: 50. cft/orof/c: see cc
phenotype: Tarsi conglutinated. RK3. cfto: chocolate
cftofe//e: see eft/ location: 1-5.4 (left of ec).
*che: cherub origin: X ray induced,
location: 2-62.0. discoverer: Weigle, 1955.
origin: Ultraviolet induced. references: Sturtevant, 1955, DIS 29: 75.
discoverer: Meyer, 48g. phenotype: Eye color brown with whitish highlights.
references: Meyer and Edmondson, 1951, DIS 25: 71. Paler than se, less purplish than pa. Malpighian
MUTATIONS 51
tubes of larvae and adults contain brown pigment or absent. Other gaps and scattered branch veins
like red. Larvae easily distinguished from wild in region of crossveins. At 19°C nearly all flies
type. Brown pigment of Malpighian tubes absent have a mutant phenotype; at 25°C there is slight
when cho is combined with v, en, or sr mutations, overlap with wild type; at 30°C virtually all flies
which prevent formation of brown eye pigment. are wild type. Dosage effect such that ci/0
Eye color of cho v is yellowish, but cho g cannot haplo-4's are more extreme than ci/ci diplo-4's,
be distinguished from g. Separability, viability, which are more extreme than ci/ci/ci triplo-4's.
and fertility excellent. RK1. For interactions of ci with en, H, ve and eg, see
other information: Not included in Df(l)N8 = House, 1953, Genetics 38: 199-215, 309-27;
Df(l)3B4-Cl;3D6-El. 1955, Anat. Record 122: 471; 1959, Genetics, 44:
516; 1961, Genetics, 46: 871. Expression of ci
origin: Spontaneous. sensitive to genetic background; selection pos-
discoverer: Green, 1955. sible for more and less extreme phenotypes (House
references: Sturtevant, 1955, DIS 29: 75. and Yeatts, 1962, Genetics 47: 960). Phenotypic
phenotype: Dark brownish eye; indistinguishable effect visible in prepupa by absence of the lower
from cho. RK1. longitudinal vein. RK1 at 19°C and higher rank
chocolate: see cho with higher temperatures.
*chr: chrome cytology: Placed in salivary chromosome region
location: 1- (not located). 101F2-102A5, on the basis of its inclusion in
origin: Spontaneous. Df(4)M63a = Df(4)101F2-102Al;102A2-5.
discoverer: Bridges, 13115. other information: The expression of ci + can be
references: Morgan and Bridges, 1916, Carnegie altered in direction of ci by certain chromosome
Inst. Wash. Publ. No. 237: 74. rearrangements that have one break in vicinity of
phenotype: Body color brownish yellow or tan. ci locus. Rearranged fourth chromosomes carrying
Abdominal bands clear yellow. RK3. a mutant allele of ci, R(ci), may also show altered
other information: Probably a tan allele. expression of gene (Stern and Kodani, 1955). R(ci)
chubby: see c/i and R(ci+) terminology not retained here; interac-
chy: chunky tion with ci included in descriptions of aberrations
location: 2- (between 8 and 28). involving chromosome 4.
origin: Spontaneous. ci3 6l
discoverer: Bridges, 38blO. origin: Spontaneous.
phenotype: Body short and heavy set. Wings discoverer: Curry, 361.
shorter than wild type. Difficult to classify. RK3. phenotype: Less extreme than ci. Ranges from
appearance of a plexus in L4 between crossveins
to gaps in L4 and L5 posterior to crossveins.
RK3.
c/ + 2; cubitus interruptus-wild-type isoallele

origin: On fourth chromosome carrying ey2.
discoverer: Stern and Schaeffer, 1943.
references: 1943, Proc. Natl. Acad. Sci. U.S. 29:
361-67.
ch cubitus interruptus phenotype: Homozygote wild type at 14° and 26°C.
Wings showing from no interruption (extreme left) to ci+2/Df(4JM wild type at 26°C; shows some
complete absence (extreme right) of the cubital vein. thinning and interruption of L4 at 14°C. ci+2/ci
From Stern and Kodani, 1955, Genetics 40: 343-73. wild type at 26°C; at 14°C fewer flies show
thinning or interruption of L4 than ci + C/ci.
ci: cubitus interruptus ci*2/ciW shows significantly greater amount of
location: 4-0 (most proximal mutant in 4). thinning and interruption of L4 than c / + c / c i w \
origin: Spontaneous. RK3.
discoverer: Tiniakov and Terentieva, 30b. c/ + 3
references: Terentieva, 1931, Zh. Eksperim. Biol. discoverer: Stern and Schaeffer, 1943.
7: 187-90 (fig.). referenees: 1943, Proc. Natl. Acad. Sci. U.S. 29:
Tiniakov and Terentieva, 1933, Genetics 18: 361-67.
117-20 (fig.). phenotype: Homozygote wild type at 26° and shows
Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20. some thinning of L4 at 14°C. About half of
Sturtevant, 1951, Proc. Natl. Acad. Sci. U.S. 37: ci+3/ci heterozygotes are not wild type at 25° and
405-7. about three-fourths are not at 14°C. Only a few
Stern and Kodani, 1955, Genetics 40: 343-73 (fig.). ci*3/ciw individuals overlap wild type. RK3.
phenotype: Vein L4 shows one or more gaps both C/+5
distal and proximal to posterior crossvein, gen- origin: A male of the Cockaponsett wild stock.
erally nonterminal. Anterior crossvein shortened discoverer: Hochman, 551.
references: 1961, Evolution 15: 239-46. synonym: It: Interntptus.

phenotype: Wild type at 25°C; at 17°C a small frac- phenotype: Homozygote is extreme ci type. Wings
tion of flies display wing vein abnormalities; how- sometimes almost twice normal width, arclike, and
ever, not involving L4. Over 80 percent of ci+5/ci virtually lack veins. Often present is a well-
flies show gaps in L4 distal to posterior cross- organized pattern of venation in which the pos-
vein. RK3. terior crossvein flows smoothly into L5. Legs
C/+C lumpy; sex combs larger than normal; antennae
origin: Canton-S wild type. enlarged; eyes smaller; and extra bristles present.
discoverer: Stern and Schaffer, 1943. Heterozygote shows gap in L4 in 80 percent of
references: 1943, Proc. Natl. Acad. Sci. U.S. 29: flies. ciw enhanced by H, en, and Cy (House,
361-67. 1953, Genetics 38: 669-70; 1959, Genetics 44:
phenotype: When homozygous, wild type at 14°, 18°, 516), Temperature effect described by House
and 25°C. ci+c/Df(4)M wild type at 26°C; very (1955, Genetics 40: 576). RK2.
few flies show thinning or interruption of L4 at cinnabar: see en
14°C. ci^C/ci wild type at 26°; some flies show ck: crinkled
thinning or interruption at 14°C. ci+c/ciw causes location: 2-53.
significant thinning or interruption of L4 at 26 C. origin: Spontaneous.
RK3. discoverer: Bridges, 30c30.
ciD: cubitus interruptus-Dorninant phenotype: Wings flimsy, crinkled, or wavy.
origin: X ray induced. Irregular stubby or wavy bristle effect. Viability
discoverer: Ruch, 32al8. poor. RK3.
references: Bridges, 1935, Biol. Zh. (Moscow), cl: clot
4: 401-20. location: 2-16.5.
phenotype: Wings show interruptions of L4 in two origin: Spontaneous.
places; proximal to and distal to, anterior cross- discoverer: Bridges, 27a3.
vein. L5 also shows distal interruption. L3 and phenotype: Eye color dark maroon to sepia-like
L5 thick. Considerable plexus effect and knotting with age; less extreme than sepia. Sixty percent
of veins. Wings broader and warped or concave more red pigment than wild type (Nolte, 1954,
upward, regularly extended and bent backward. J. Genet. 52: 127—39). Eye color autonomous
Alula fused with and in same plane as blade of when larval optic disk is transplanted into wild-
wing. Black dried blood from axillary spiracle. type host (Beadle and Ephrussi, 1936, Genetics
Slight scalloping of inner wing margin, with hairs 21: 230). Larval Malpighian tubes pale yellow,
and tufts. Direction and extent of temperature distinguishable from wild type (Brehme and
effects depends on genetic background (Scharloo). Demerec, 1942, Growth 6: 351-56). RK1.
In general, no overlapping wild type. H/+ inhibits cytology: Placed in salivary chromosome region
scalloping of ciD but greatly enhances L4 inter- 25E1 to 26C1 (E. H. Grell).
ruption (House, 1959, Genetics 44: 516). Fully c/2
dominant in triplo-4's (Sturtevant, 1936, Genetics discoverer: Terry, 1928.
21: 448). Homozygous lethal. Lethal acts in phenotype: Eye color like cl but darker. Larval
embryonic stage (Hochman). RK1. Malpighian tubes pale yellow (Brehme and
cytology: Salivary study by Bridges revealed no Demerec, 1942, Growth 6: 351-56). RK1.
chromosomal aberration. claref: see ca
other information: Not allelic, at least with respect *c/c/: cloudy
to its lethality, since ciD/Df(4)M63a survives, location: 2-96 to -101.
whereas ci/Dt(4)M63a is mutant (Hochman, 1965, origin: Gamma ray induced.
DIS 40: 60). discoverer: Wallbrunn, 61 j6.
c / 0 - 6 ; cubitus interruptus-Dominant of G/oor references: 1964, DIS 39: 59.
origin: Obtained by recombination between chromo- phenotype: Wings opaque from fluid between upper
somes with ci® and spaP°K and lower membranes; occasionally fluid forms
discoverer: Gloor. small blisters. Males sterile, females highly
references: Scharloo, 1963, DIS 38: 32. infertile. RK2.
phenotype: Less extreme than ciD. Interruption of cleft: see cf
L5 is infrequent. Usually a terminal interruption elf; see wtwdf
of L4 distal to second crossvein and a gap prox- *Cli: Clipped wings
imal to first crossvein. Wings neither spread nor location: 1- (to the left of f).
warped. No black dots present in axillary discoverer: Agol.
spiracles, but overlaps wild type at lower temper- references: 1936, DIS 5: 7.
atures. Good expressivity at 25°C (Scharloo). phenotype: Dominant wing mutant (no description
RK1. given). Viable in male and homozygous female.
ci*; cuhitus interrvptvs of Wallace RK3.
origin: Spontaneous. clip wing: see
discoverer: E. M. Wallace, 36d20. clipped: see cp
MUTATIONS 53
Clipped wings: see Cli More precisely located by Hannah-Alava in 6E,

Clipt: see Cpt probably in or near 6E6.
elm: clumpy marginals cm: see emp
location: 1-32.6.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- origin: Induced by mustard gas.
phenylalanine (CB. 3025). discoverer: Sobels, 571.
discoverer: Fahmy, 1953. references: 1958, DIS 32: 84.
references: 1958, DIS 32: 68. phenotype: Eye color more translucent and ruby-like
phenotype: Irregularly bent marginal hairs, than cm. RK1.
especially on posterior border of wings. Bristles cm ^
stiff and frequently bent or split. Viability and origin: X ray induced in R(l)2.
fertility of males good. Homozygous females discoverer: Muller, Valencia, and Valencia, 1946-53.
reduced in viability and fertility. RK2. references: Valencia, 1966, DIS 41: 58.
other information: One allele each induced by cytology: Associated with Dt(l)cmR8BH4 =Df(l)6E
CB. 1506 and CB. 3007. (J. I. Valencia).
clot: see c/
cloudy: see eld
cloven thorax: see c/v
club: see cb
clubfoot: see cbf
cluboid: see cbd
clumpy marginals: see elm
*c/v-7; cloven thorax no. 7

location: 1-0.0. Cm: Crimp
origin: X ray induced. Edith M. Wallace, unpublished.
discoverer: Muller, 19h.
references: 1935, DIS 3: 29. *Cm: Crimp
phenotype: Thorax often has long cleft; partially location: 3-43.5.
dominant. Semilethal at low temperature, viable origin: Spontaneous.
at high one. RK3. discoverer: Bridges, 28a28.
*clv-2 phenotype: Heterozygote has crimped wings ruffled
location: 1-42.0. on rear edge. Classification good in first 4 days'
origin: X ray induced. hatch, then Cm overlaps wild type progressively.
discoverer: Muller, 26111. Better at 25° than at 19°C. Homozygous lethal.
references: 1935, DIS 3: 29. RK2 as lethal; RK3 as dominant.
phenotype: Thorax has logitudinal cleft, sometimes
half thorax. One wing often reduced or like vg.
Partially dominant. Semilethal. RK3.
*cly.2S2b
discoverer: Bateman, 52b.
phenotype: Like clv-2. Some flies have no dorsal
thorax at all. Viability 30 percent. Penetrance 50
percent. The apparently wild-type males are
fertile; abnormal ones sterile. Completely
recessive. RK3.
other information: Allelism with clv-2 not tested.
cm: carmine
location: 1-18.9.
discoverer: Mohr, 27d27. crop: crumpled
references: 1927, Z. Induktive Abstammungs- Edith M. Wallace, unpublished.
Vererbungslehre 45: 403—5.
phenotype: Eye color translucent dark ruby. With emp: crumpled
st, eye color deep orange; with brown, slightly location: 3-93.
lighter than bw alone. Larval Malpighian tubes origin: Spontaneous.
very pale yellow. RK1. discoverer: Bridges, 22d2.
cytology: Locus lies between 6A3-4 and 6F10-11 synonym: cm.
(Demerec, Kaufmann, Fano, Sutton, and Sansome, references: Bridges and Morgan, 1923, Carnegie
1942, Carnegie Inst. Wash. Year Book 41: 191). Inst. Wash, Publ. No. 327: 247.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. cytology: Associated with a tandem duplication,
Genet. 2: 223. Dp(l;l)Co = Dp(l;l)3C4-5;3D6-El (Schultz, 1941,
phenotype: Wings about two-thirds normal size and DIS 14: 54-55). Result of duplication of 3C7,
greatly crumpled or blistered. Marginal hairs deficiency for which gives Notch (Morgan, Schultz,
irregularly clumped. Legs irregularly shortened and Curry, 1941, Carnegie Inst. Wash. Year Book
and gnarled. Bristles somewhat short and thick. 40: 283).
Posterior scutellars slightly divergent. Branches other information: Reversion to wild type occurs in
of aristae bent anteriorly near middle, with apices Co/Co by unequal crossing over. A member of the
parallel to main axes of aristae. Viability and Notch pseudoallelic complex.
fertility may be low. RK3.
en: cinnabar
location: 2-57.5.
discoverer: Clausen, 2018.
references: 1924, J. Exptl. Zool. 38: 423-36.
phenotype: Eye color bright red, like v or st.
Ocelli colorless. Eye color darkens with age, but
ocelli remain colorless. Larval Malpighian tubes Co: Confluens
pale yellow (Beadle, 1937, Genetics 22: 587— Edith M. Wallace, unpublished.
611). Nonautonomous in development of pigment
of transplanted eye disks (Beadle and Ephrussi, Co-3A: see l(2)S3a
1936, Genetics 21: 230), en blocks conversion of Co-7: see 1(2)S7
kynurenine to 3-hydroxykynurenine, which has coal: see cal
been identified as the en "*" hormone (Butenandt, *coc: collapsed ocelli
Weidel, and Schlossberger, 1949, Z. Naturforsch. location: 1-61.5.
4b: 242-44). RK1. origin: Induced by D-l:6-dimethanesulfonyl mannitol
cytology: Proximal to 44C, based on its inclusion in (CB. 2511).
Dp(2;3)P32 = Dp(2;3)41A;42D-E;44C-D;89D7-El discoverer: Fahmy, 1960.
(E. B. Lewis). references: 1964, DIS 39: 58.
cn2 phenotype: Ocelli small and flat; deflated owing to
origin: Spontaneous in In(2R)Cy. lack of eye fluid. Anterior ocellar hairs frequently
discoverer: L. Ward, 1921. missing. Other slight alterations in body size and
references: 1923, Genetics 8: 276—300. wing shape. RK3.
phenotype: Eye color slightly brighter than normal cytology: Placed in salivary region 18A4 through
and ocelli pale. cn2/cn bright scarlet like cn/cn 18B8 on the basis of its inclusion within the defi-
and has colorless ocelli but darkens more rapidly ciency carrying the left end of In(l)y4 = In(l)lA8-
with age. Malpighian tubes of en2/en pale yellow Bl;18A3-4 and the right end of In(l)sc^ =In(l)lB2-
(Brehme and Demerec, 1942, Growth 6: 351—56). 3;18B8-9 (Norton and Valencia, 1965, DIS 40: 40).
RK2.
en*: cinnabar-sterile Co/: Coiled
origin: Spontaneous. location: 2-48.7.
discoverer: Ives, 40el8. origin: X ray induced.
phenotype: Eye color like en, but females sterile. discoverer: Carlson, 57g.
RK2, references: 1960, DIS 34: 48.
other information: Possibility that sterility factor phenotype: Coi/+ has curled wings like Cy/+ or j.
separable from en not completely eliminated. Coi/Coi viable, with strongly curled wings similar
Found in chromosome carrying In(2L)t = to Cy/+; JD/+. Coi/j shows no interaction. Excel-
In(2L)22D3-El;34A8-9. lent viability and fertility. RK1.
Co: Confluens cytology: Not included in Df(2L)64j = Df(2L)34E5-
location: 1-3.0. F1;35C3-D1 (E. H. Grell).
origin: Recovered among progeny of cold-treated Coiled: see Cd
fly. Coiledex: see Co'
discoverer: Gottschewski 34c. collapsed ocelli: see coc
references: 1935, DIS 4: 7, 14, 16. *co/n: compressed
1937, Z. Induktive Abstammungs- Vererbungslehre location: 3-48.5.
73: 131-42. origin: Spontaneous.
1937, DIS 8: 12. discoverer: Bridges, 18k27.
phenotype: Veins irregularly thickened, especially references: Bridges and Morgan, 1923, Carnegie
toward tips, which are usually deltas and fused Inst. Wash. Publ. No. 327: 193.
broadly to marginal vein. Stronger expression in phenotype: Head flattened ventraliy. Eyes small,
males than in females. Co/N8 wild type except for displaced. Vibrissae tufted. Aristae crumpled.
slightly thicker L3 vein. Co/Ax like Ax/+. RK1A. Humeral patches elevated. Wings droopy. Poor
viability and fertility. RK3.
MUTATIONS 55
*com~d: compres sed-dilapida tor corr; corrugated wing

location: 3-68.5. location: 2-36.
discoverer: Bridges, 19c8. discoverer: Mayeda, 61g.
phenotype: Flies small, pale, weak, with defective references: 1963, DIS 38: 31.
legs and wings. RK3. phenotype: Wings wrinkled and wavy, reduced to
comb gap: see eg three-fourths normal size. Whole wing corrugated
compressed: see com at 20°C, only posterior third at 25°. Good classi-
*con: condensed fication. RK2.
location: 1-27.1.
origin: Spontaneous. Corroded eye: see Cor
discoverer: Bridges, 36dll. corrugated wing: see corr
references: 1937, DIS 7: 6. costakink: see csk
phenotype: Thorax and abdomen shortened; abdomen cp: clipped
dilated, exposing ventral skin to side view. Eyes location: 3-45.3.
slightly roughened, occasionally kidney shaped discoverer: Mainx, 34g.
and somewhat dark. Wings short, bluntly rounded, references: 1936, Z. Induktive Abstammungs-
with crossveins closer together than normal. Vererbungslehre 71: 303-4 (fig.).
Bristles shortened and somewhat fine at 19°C, Pollitzer, 1937, DIS 8: 91.
stubby at 25°C. Postscutellars semierect and phenotype: Wing margins snipped, most often along
crossed; posterior verticals shortened or missing. marginal vein. At 19°C character slighter but
Male entirely sterile. Viability 50 percent wild completely penetrant. RK1.
type. RK2.
cytology: Salivary chromosome studies (Demerec,
Kaufmann, Fano, Sutton, and Sansome, 1942,
Carnegie Inst. Wash. Year Book 41: 191) show
locus to lie between 7C4-5 and 8C1-2. Further
restricted to 7E1 through 8C2, on the basis of its
genetic location to the right of oc which is ex-
cluded from Di(l)sn = Df(l)7B2-3;7D22-El (Hinton
and Welshons, 1955, DIS 29: 125-26).
concave wing: see ccw cp; clipped
condensed: see con Edith M. Wallace, unpublished.
Confluens: see Co
Confluent: see Cf *cpl: cupola
Confluent-3: see Dlcf-3 location: 1-0.0 (no crossing over with sc in 584
contorted: see ctt males).
Contrabithorax: see Cbx origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
convex wing: see cvw phenylalanine (CB. 3025).
cop; copper discoverer: Fahmy, 1953.
location: 1-43.3. references: 1959, DIS 33: 83-84.
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- phenotype: Small, inviable fly. Wings shorter and
phenylalanine (CB. 3026). curved to form canopy over abdomen with tips
discoverer: Fahmy, 1953. converging toward mid-dorsal line. Head and eyes
references: 1958, DIS 32: 68. slightly deformed. Abdominal tergites abnormal;
phenotype: Brownish red eye color. Best classifi- from irregular pigmentation to absence or gross
cation in newly emerged flies. Occasionally wings deformation of the sixth and seventh tergites.
show cutaway inner margins. Excellent viability Males sterile. RK3.
and fertility in both sexes. RK2. *Cpt: Clipt
other information: Two alleles induced by CB. 3025. location: 2-43.7.
Cor: Corroded eye origin: Spontaneous.
location: 3- (not located), discoverer: Sturtevant, 26bl8,
origin: X ray induced, phenotype: Bristles short, like those of Sb. Homo-
discoverer: Muller. zygous lethal. Male sterile. RK1.
references: 1946, DIS 20: 66. *cpw: canopy wing
phenotype: Cor/+ shows slight irregular flecking of location: 1-2.5.
eye. In combination with v, expression enhanced, origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
producing patchy diminution in color, especially phenylalanine (CB. 3025).
near posterior margin of eye, giving impression discoverer: Fahmy, 1953.
that color was washed or eaten away, especially references: 1958, DIS 32: 69.
from deeper layers; regions of surface often phenotype: Wings short and very broad; longitudinal
blackened. Homozygote not described. RK2. veins frequently do not reach wing margin and
often diverge. Eyes large and slightly rough- Cream: see Cr

Head bristles reduced in number (ocellars most Cream: see
frequently affected). Thorax broad, one or more cream underscored: see cru
bristles occasionally absent; hairs more widely creased: see cs
separated, with noticeable hairless areas. Males creeper: see rk4
sterile. Viability 40 percent wild type. RK3. Crimp: see Cm
eg; see rk4 crinkled: see ck
*cr: crisp *crip: cripple
location: 1- (not located). location: 2- (between pr and en).
discoverer: Agol. discoverer: Komai, 1924.
references: 1936, DIS 5: 7. references: 1926, Genetics 11: 280-93.
phenotype: Bristles like forked. RK2. 1927, Mem. Coll. Sci. Univ. Kyoto, Ser. B 2: 211 —
other information: Not an allele of I or sn. 57.
*Cr-2: Cream in chromosome 2 phenotype: Middle and hind legs twisted and
location: 2- (not located). shortened. Thirty percent penetrance. RK3.
origin; Spontaneous. crisp: see cr
discoverer: Bridges, 13il5. *crk: crooked setae
references: 1919, J. Exptl. Zool. 28: 337-84. location: 1-60.1.
Bridges and Morgan, 1919, Carnegie Inst. Wash. origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
Publ. No. 278: 239 (fig.). phenylalanine (CB. 3026).
phenotype: Specific dilutor of w0, we; Cr-2/Cr-2 discoverer: Fahmy, 1953.
has a pale cream eye color. we; Cr-2/+ has eye references: 1958, DIS 32: 69.
color between eosin and cream. RK3. phenotype: Bristles thin and slightly shortened;
*cr-3: cream in chromosome 3 occasional missing scutellar. Acrostichals de-
location: 3-36.5. ranged. Abdominal hairs of female frequently
origin: Spontaneous, missing; tergites occasionally abnormal. Classifi-
discoverer: E. M. Wallace, 14b27. cation difficult. Viability and fertility good. RK3.
references: Bridges, 1919, J. Exptl. Zool. 28: other information: One allele induced by CB. 3025.
337-84. *crm: cramped
Bridges and Morgan, 1923, Carnegie Inst. Wash. location: 1-1.4 (based on crm^).
Publ. No. 327: 112 (fig.). origin: Induced by P32.
phenotype: Homozygote has slightly diluted eye discoverer: Bateman.
color. Eye color of we; cr-3 cream. Larval Mal- synonym: stap: stubarista from P32.
pighian tubes of w°; cr-3 white, those of cr-3 references: 1951, DIS 25: 78.
bright yellow (Brehme and Demerec, 1942, Growth 1953, DIS 27: 55.
6: 351-56). RK3. phenotype: Antennae stumpy with shrunken, warped
*cr-a: cream-a aristae, usually lying back on head. Initially
location: Autosomal, not located. showed narrowed and scalloped wings and eyes
origin: Spontaneous. shaped like inverted pears, but these effects vari-
discoverer: Bridges, 13gl5. able. Viability low. Sterile. RK3.
references: 1916, Genetics 1: 147.
1919, J. Exptl. Zool. 28: 337-84. origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
phenotype: Strong specific dilutor of we. RK3. phenylalanine (CB. 3025).
*cr-b discoverer: Fahmy, 1953.
location: 2-24. synonym: cramped-like.
discoverer: Bridges, 14clO. phenotype: Antennae stumpy and short, with
references: 1916, Genetics 1: 149. reduced, abnormal aristae. Wings frequently
1919, J. Exptl. Zool. 28: 337-84. abnormally held, with cuts on inner margins. Eyes
Bridges and Morgan, 1919, Carnegie Inst. Wash. pear shaped. Small extra sex combs on second
Publ. No. 278: 245 (fig.). tarsal segment. Not easily classified. Viability
phenotype: Specific dilutor of w e . RK3. and fertility slightly reduced in males, greatly
*cr-c reduced in females. RK3.
location: 2- (near S). other information: Allelism to crm inferred from po-
origin; Spontaneous. sition of ctni at 1:14 and phenotype. Seven other
discoverer: Bridges, 16gl3. alleles: 1 each induced by CB. 1540, CB 1592,
phenotype: Weak specific dilutor of we. RK3. CB. 3007, CB. 3025, CB. 3034, and 2 induced by
cramped: see crm CB. 1506.
cramped-like: see crm2 cro: see ptg3
CRB: see T(1;4)A1 crooked: see fwc
etc: see l('2)crc crooked setae: see crk
cream: see cr crossover suppressor: see c( )
MUTATIONS 57
crossveinless; see cv *csk: costakink

crowri: see ptg3 location: 1-33.0 (no crossovers with v in 526
crs: cru sterile males).
location: 2- (between px and bw). origin: Induced by triethylenemelamine (CB. 1246).
discoverer: Muller. discoverer: Fahmy, 1953.
1955, DIS 29: 146. phenotype: Eyes smaller. Wings slightly reduced in
phenotype: Male sterile. RK2. size and abnormally held; costal vein frequently
cytology: Located between 58E3 and 59A2 on basis kinked near L2. Not fully penetrant. Male
of sterility in combination with Di(2R)P + viability and fertility good, but female viability
Dp(2;Y)bw+ = Di(2R)58E3-Fl;60D14-E2 + and fertility reduced to about 50 percent wild
Dp(2;Y)YL;58Fl-59A2;60D14-E2 (Muller, 1955). type. RK3.
other information: Male sterility formerly associated other information: One X-ray-induced allele.
with but separable from cru. cf: cuf
crt: crumpled tips location: 1-20.0.
location: 1-40.3 (7.3 units from v, based on 3035 origin: Spontaneous.
flies). discoverer: Bridges, 15J12.
origin: Induced by triethylenemelamine (CB. 1246). references: Morgan, Bridges, and Sturtevant, 1925,
discoverer: Fahmy, 1952. Bibliog. Genet. 2: 35, 223 (fig.).
references: 1959, DIS 33: 84. phenotype: Wings cut to points and edges scalloped.
phenotype: Wing tips frequently shriveled, pleated, Eyes smaller and somewhat kidney shaped.
or crumpled, and often turned up or down. Wings Abdominal bands warped. Antennae often
vary from completely unexpanded to wild type. deformed. RK1.
Viability and fertility good in both sexes. RK2. cytology: Placed in salivary gland chromosome
other information: Twelve other alleles: 1 each bands 7B3-4 (Demerec, Kaufmann, Fano, Sutton,
induced by X rays, CB. 1246, CB. 1522, and and Sansome, 1942, Carnegie Inst. Wash. Year
CB. 3025; 2 induced by CB. 3034; 3 each induced Book 41: 191). Hannah-Alava agrees.
by CB. 1592 and CB. 3007.
cru: cream underscored
location: 2-52.5. origin: X ray induced.
origin: Spontaneous. discoverer: Hannah, 1947.
discoverer: Bridges, 20a5. references: 1949, Proc. Intern. Congr. Genet., 8th.
phenotype: Specific dilutor of we and P. Slight pp. 588-89.
dominant but used as a recessive. Originally phenotype: Lethal. Shows cut phenotype in combi-
thought to be male sterile, but this was caused by nation with viable ct alleles; lethal in combination
a factor in 2R, crs. Larval Malpighian tubes of with lethal ct alleles. RK2A.
we; cru colorless; those of +; cru bright yellow cytology: Associated with Dl(l)ctâ^ =
(Brehme and Demerec, 1942, Growth 6: 351—56). Df(l)7B3-6;7B6-7.
RK3. *cf2a3
crumpled: see crop origin: X ray induced.
crumpled tips: see erf discoverer: Hannah, 1947.
cru sterile: see crs references: 1949, Proc. Intern. Congr. Genet., 8th.
*cs; creased pp. 588-89.
location: 1-56. phenotype: Like ct**?. RK2A.
origin: X ray induced. cytology: Associated with D((l)ct2a3 =
discoverer: K. C. Atwood, 41i. Dt(l)7B2-3;7Cl-2.
references: 1942, DIS 16: 47. *ct2cl
phenotype: Wings logitudinally creased in first origin: X ray induced.
posterior cell from distal end of L3 virtually to discoverer: Hannah, 1947.
anterior crossvein. Fertility and viability good. references: 1949, Proc. Intern. Congr. Genet., 8th.
RK1. pp. 588-89.
CS53 phenotype: Like ct2a^. RK2.
origin: X ray induced. cytology: Salivary chromosomes normal.
discoverer: Krivshenko, 53k5. *ct*
references: 1956, DIS 30: 74. discoverer: Morgan, 17a22.
phenotype: Wing longitudinally creased. Lateral references: Morgan, Bridges, and Sturtevant, 1925,
edges of wings bent slightly downward distally. Bibliog. Genet. 2: 35 (fip.).
RK1. phenotype: Extreme ct allele. Wings short, dark,
other information: According to crossover data of and crumpled as well as cut and scalloped.
M. Aronson and description, this is an allele of Abdominal bands warped. Antennae flattened and
cs, which was lost before this mutant was embedded. Aristae concave forward. Eyes smaller
discovered. and kidney shaped. Vibrissae gone. More extreme
expression in females than in males; females have phenotype: Lethal. Expression in combination with
much poorer viability. 1 female:3 males in stock. ct6 suppressed by YL. RK2A.
RK3. cytology: Associated with Tp(l)ct6al =
* c f3o2 Tp(l)7B2-Cl;19;20.
discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th.
pp. 588-89.
phenotype: Lethal in male and homozygous female,
as well as in combination with other lethal alleles
of ct. Extreme ct phenotype in combination with
ct6. Like ctn in combination with ctn, Phenotype
reportedly suppressed by addition of Y chromo-
some material. RK2A.
cytology: Associated with In(l)ct3a2 —
In(l)7B2-Cl; 19-20.

pp. 588-89.
phenotype: Lethal, but unlike other lethal ct
alleles tested by Hannah, fails to survive in
combination with Dp(l;3)sn13al. RK2A.
cytology: Associated with In(l)ct3bI =
In(l)3A4-Bl;7B2-5.
ctf>: cut-6
origin: X ray induced. From Morgan, Bridges, and Sturtevant, 1925, Bibliob.
discoverer: Hannah, 1947. Genet. 2: 35.
pp. 588-89. *C/7a7
phenotype: Like cf 2 °2. RK2A. origin: X ray induced.
cytology: Associated with Df(l)ct4bi = discoverer: Hannah, 1947.
Di(l)7B2-4;7C2-4. references: 1949, Proc. Intern. Congr. Genet., 8th.
*ct4d pp. 588-89.
origin: X ray induced. phenotype: Lethal. ct7al /Dp(l;3)sn13al males
discoverer: Hannah, 1947. show reduced viability and are probably sterile.
references: 1949, Proc. Intern. Congr. Genet., 8th. RK2A.
pp. 588-89. cytology: Associated with T(l;2)ct7ai = T(1;2)7B.
phenotype: Like ctl*2. R K 2 . * c f7o2
cytology: Salivary chromosomes normal. origin: X ray induced.
origin: Spontaneous. pp. 588-89.
discoverer: Bridges, 2Oc20. phenotype: Like c*2a2. RK2A.
references: Morgan, Bridges, and Sturtevant, 1925, cytology: Associated with Df(l)ct?a2 =
Bibliog. Genet. 2: 35 (fig.). Di(l)7A5-Bl;7C4-9.
phenotype: Wing character uniform and reliable; *cf7b2
usually lacks the abdomen, antenna, arista, and origin: X ray induced.
eye effects of ct. Vibrissae gone or displaced discoverer: Hannah, 1947.
downward to bottom of eye. Developmental study references: 1949, Proc. Intern. Congr. Genet., 8th.
by Waddington Il939, Proc. Natl. Acad. Sci. U.S. pp. 588-89.
25: 299-308; 1940, J. Genet. 41: 75-139 (fig.)] phenotype: Like cf 2fl 2. RK2.
shows wing bud narrower than wild type as early cytology: Salivary chromosomes normal.
as just after eversion of wing in early pupa. RK1. * c f 7c7
other information: Allele most used as genetic origin: X ray induced.
marker. discoverer: Hannah, 1947.
origin: X ray induced, pp. 588-89.
discoverer: Hannah, 1947. phenotype: Like ct3*2. RK2A.
references: 1949, Proc. Intern. Congr. Genet., 8th. cytology: Associated with T(l;2)ct7cl -
pp. 588-89. T(1;2)7B2-3;8E2-3;25C.
MUTATIONS 59
*ct7c2
discoverer: Hannah, 1947. discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th. references: 1949, Proc. Intern. Congr. Genet., 8th.
pp. 588-89. pp. 588-89.
phenotype: Like ct2a2. RK2A. phenotype: Like ct**2. RK2A.
cytology: Associated with Df(l)ct?c2 = cytology: Associated with T(l;3)ctâ =
Df(l)6Fll-7Al;7B8-Cl. T(1;3)1B; 7B2-3;8E-F;84B.
*cf9bl * c f72a7
discoverer: Hannah, 1947. discoverer: DeFrank, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th. references: Hannah, 1949, Proc. Intern. Congr.
pp. 588—89. Genet., 8th. pp. 588-89.
phenotype: Similar to ct6. RK1. phenotype: Lethal in hemizygote and in combina-
cytology: Salivary chromosomes normal, but tion with other lethal ct alleles. Expression vari-
staining of 7B1-2 and 7B5 lighter and darker than able in combination with viable ct alleles.
normal, respectively. ct12al /Dp(l;3)ani3al males show low viability
other information: Induced simultaneously with an and are sterile. RK3.
independent but closely linked recessive lethal cytology: Salivary chromosomes normal.
mutation. *C/J2o2
*cf9b2 origin: X ray induced.
origin: X ray induced. discoverer: DeFrank, 1947.
discoverer: Hannah, 1947. references: Hannah, 1949, Proc. Intern. Congr.
references: 1949, Proc. Intern. Congr. Genet., 8th. Genet., 8th. pp. 588-89.
pp. 588-89. phenotype: Like ct2a2 except that lethal not
phenotype: Semilethal but semilethality not shown covered by Dp(l;3)sn13al. RK2A.
to be at ct locus; males appear rarely; sterile with cytology: Associated with In(l)ct12a2 =
small yellowish tan bodies. Margin notched from ln(l)4E2-3;7B2-4.
the costal cell around wings to base of inner * c f72c7
margin. RK2A. origin: X ray induced.
cytology: Induced simultaneously with but appar- discoverer: Hannah, 1947.
ently independently of a complex inversion and references: 1949, Proc. Intern. Congr. Genet., 8th.
translocation between X and 3R. pp. 588-89.
phenotype: Lethal, In combination with viable ct
* c f70o7 alleles, some flies show abnormal venation.
origin: X ray induced. ctl2cl/Dp(l;3)snl3al males viable but sterile.
discoverer: DeFrank, 1947. RK2A.
references: Hannah, 1949, Proc. Intern. Congr. cytology: Associated with T(l;3)ctI2cl =
Genet., 8th. pp. 588—89. T(l ;3)7B2-3;7D2-6;85.
phenotype: Like ct2a2. RK2A.
cytology: Possibly associated with Dt(l)ct1Oal = origin: X ray induced.
Df(l)7B3-4;7B6-7. discoverer: Hannah, 1947.
origin: X ray induced. pp. 588-89.
discoverer: Hannah, 1947. phenotype: Like ct2*2. RK2A.
references: 1949, Proc. Intern. Congr. Genet., 8th. cytology: Possibly associated with Dt(l)ct12c2 —
pp. 588-89. Df(l)7B2-3;7B6-7.
phenotype: Like ct2a2'. RK2A.
cytology: Associated with Dl(l)ct1Obl = origin: Spontaneous.
D((1)6D8-E1;7B7-C1. discoverer: Bridges, 21f7.
* c f70c7 references: Morgan, Bridges, and Sturtevant, 1925,
origin: X ray induced. Bibliog. Genet. 25: 35 (fig.).
discoverer: DeFrank, 1947. phenotype: Like ct but females usually sterile.
references: Hannah, 1949, Proc. Intern. Congr. RK2.
Genet., 8th. pp. 588-89. Cf73o7
phenotype: Males have notched but unexpended origin: X ray induced.

wings; may have abnormal antennae and vibrissae. discoverer: Hannah, 1947.
Males show very low viability, usually dying in references: 1949, Proc. Intern. Congr. Genet., 8th.
larval stage. Surviving males sterile. Lethal in pp. 588-89.
combination with other lethal ct alleles; heterozy- phenotype: Like ct2®2. RK2A.
gous females made using Dp{l;3)&n13l*l. RK2. cytology: Associated with In(l)ct*3mi ~
cytology: Salivary chromosomes normal. In(l)7B2-3; 19-20.
cytology: Associated with Df(l)ct14cl =

origin: X ray induced. Df(l)7B3-4;7B6-9.
references: 1949, Proc. Intern. Congr. Genet., 8th. origin: X ray induced.
pp. 588-89. discoverer: Hannah, 1947.
phenotype: Similar to ct6. RK1. references: 1949, Proc. Intern. Congr. Genet., 8th.
cytology: 7B1-2 show abnormal staining and pp. 588-89.
ectopic pairing with heterochromatic regions. phenotype: Like ct2*2. RK2A.
cytology: Associated with Df(l)ct1Sbl =
origin: X ray induced. Df(l)7B2-4;7B6-7.
references: 1949, Proc. Intern. Congr. Genet., 8th. ct15B4
pp. 588-89. origin: X ray induced in In(l)scslLsc8R+dl-49.

phenotype: Like ctê?. RK2A. discoverer: Muller, Valencia, and Valencia, 1946-53
cytology: Associated with T(l;4)ctl3bl - references: Valencia, 1966, DIS41: 58.
T(l;4)lA;7B2-3;101A-D;102. phenotype: Male viable. RK2.
*cf36b
origin: X ray induced. origin: Spontaneous.
discoverer: Hannah, 1947. discoverer: Stalker, 36b28.
references: 1949, Proc. Intern. Congr. Genet., 8th. references: Spencer, 1937, DIS 7: 20.
pp. 588-89. phenotype: Slight nick at tip of one or both wings.
phenotype: Slightly less extreme than ct6. RK1. Less than 50 percent penetrance at 19°C, 85 per-
cytology: Salivary chromosomes normal. cent penetrance at 25°, and 100 percent pene-
*cfUa2 trance at 29°. RK3.
origin: X ray induced,
discoverer: Hannah, 1947. cf43aHl
references: 1949, Proc. Intern. Congr. Genet., 8th. origin: X ray induced.
pp. 588-89. discoverer: Muller, Valencia, and Valencia, 1946-53.
phenotype: Like ct^^. RK2A. references: Valencia, 1966, DIS 41: 58.
cytology: Associated with T(l;2)ctl4a2 = phenotype: Male lethal. RK2A.
T(l;2)7B2-4;19-20;41El-2. cytology: Associated with In(l)ct43aH1 = In(l)4Bl-
*cfl4a3 4;7B4-C1 + In(l)10D5-6;20B-C.
discoverer: Hannah, 1947. origin: X ray induced.
references: 1949, Proc. Intern. Congr. Genet., 8th. discoverer: King and Poulson, 461.
pp. 588-89. references: Poulson and King, 1948, DIS 22: 54.
phenotype: Like ct2a2f but ct^'fa3/Dp(l;3)sn^3ai phenotype: Distal edges of wings scalloped in area
shows reduced viability. RK2A. between L3 and L4, and, occasionally, lateral
cytology: Associated with In(l)ct14a3 = surface of wing toward L5 scalloped. Abdomen,
ln(l)7B2-3;20. antenna, arista, and eye effects of ct absent.
Classification of males and females reliable.
origin: X ray induced. More extreme than ctn and more viable than ct6.
discoverer: Hannah, 1947. ct461/ct6 flies have slightly nicked wing tips
references: 1949, Proc. Intern. Congr. Genet., 8th. resembling ctn. RK1.
pp. 588-89. * c f50e
phenotype: Like cf2«2, RK2A. origin: Spontaneous.
cytology: Associated with Dt(l)ct14bl = discoverer: Bakkum, 5Oe.
Df(l)7B2-3;7C3'4. references: Mickey, 1951, DIS 25: 74.
*CfUb2 phenotype: Wings cut to points. Eyes slightly
origin: X ray induced. ovoid. Viability and fertility lowered, especially
discoverer: Hannah, 1947. in females. RK2.
pp. 588-89. origin: Recovered among progeny of radiofrequency-
phenotype: Like c t ^ . RK2A. treated male,
cytology: Associated with In(l)ct14l>2 - discoverer: Mickey.
In(l)3D2-5;7B2'4. references: 1963, DIS 38: 28.
*cfUc1 phenotype: Like ct. RK1.
origin: X ray induced. *cf62f
discoverer: Hannah, 1947. origin: Spontaneous in Base.
references: 1949, Proc. Intern. Congr. Genet., 8th. discoverer: Mickey, 62f8.
pp. 588-89. references: 1963, DIS 38: 28.
phenotype: Like cf-?«2. RK2A. phenotype: Lethal in male. RK2A.
MUTATIONS 61
*cf268-l *cf268-21
discoverer: Demerec, 33j. discoverer: Hoover, 35i.
phenotype: Lethal and cell lethal. RK2. phenotype: Lethal. RK2A.
cytology: Salivary chromosomes normal (Hoover). cytology: Associated with T(l;3)ct268-21 -
* c f268-2 T(l ;3)7B3-4; 7B4-5; 96 F.
origin: X ray induced. *cf268-23
discoverer: Demerec, 33k. origin: X ray induced.
phenotype: Lethal but not cell lethal. RK2. discoverer: Hoover, 35g.
cytology: Salivary chromosomes normal (Hoover). phenotype: Lethal but not cell lethal. RK2(A).
*cf268-3 cytology: Salivary chromosomes show possible
origin: X ray induced. deficiency for 7B3.
discoverer: Demerec, 33k. *cf268-24
phenotype: Lethal but not cell lethal. RK2. origin: X ray induced.
cytology: Salivary chromosomes normal (Hoover). discoverer: Hoover, 35i.
*cf268-5 phenotype: Lethal. RK2A.
origin: X ray induced. cytology: Associated with T(l;2)ct268-24 -
discoverer: Demerec, 33k. T(1;2)7B2-5;41F6-42A1.
phenotype: Lethal. RK2A. *cf268-26
cytology: Associated with T(l;3)ct268-5 = origin: X ray induced.
T(l;3)7B2-3;90C4-Dl. discoverer: Hoover, 35j.
*cf268-6 phenotype: Lethal. RK2A.
origin: X ray induced. cytology: Associated with T(l;2)ct268-26 -
discoverer: Demerec, 331. T(1;2)7B3-C1;36E.
phenotype: Lethal but not cell lethal. RK2. *cf26B-27
cytology: Salivary chromosomes normal (Hoover). origin: X ray induced.
*cf268-13 discoverer: Hoover, 35j.
origin: X ray induced. references: 1938, Z. Induktive Abstammungs-
discoverer: Demerec, 34f. Vererbungslehre 74: 420-34.
references: Hoover, 1937, Genetics 22: 634-40. phenotype: Lethal but not cell lethal. RK2A.
1938, Z. Induktive Abstammungs- Vererbungslehre cytology: Associated with In(l)ct268-27 =
74: 420-34. In(l)3D6-El;7B3-5.
phenotype: Lethal but not cell lethal. RK2A. *cf268-29
cytology: Associated with In(l)ct26S-l 3 ~ln(l)2E3- origin: X ray induced.
F1;2F2-3;7B2-3;7B4-5;19A4-5;19A6-B1. Salivary discoverer: Demerec, 38d.
bands 2F1-2, 7B3-4, and 19A5-6 missing. phenotype: Lethal but not shown that lethality at
* c f268-75 ct locus. RK2A.
origin: X ray induced. cytology: Induced simultaneously with but presum-
discoverer: Demerec, 34g. ably separable from T(l;3)lz268'29 =
phenotype: Lethal but not cell lethal. RK2. T(1;3)8D8-9;81F.
cytology: Salivary chromosomes normal (Hoover). *cf268-30
*cf268-17 origin: X ray induced.
origin: X ray induced. discoverer: Hoover, 38d.
discoverer: Demerec, 34h. phenotype: Lethal and cell lethal. RK2A.
phenotype: Lethal. RK2A. cytology: Associated with Df(l)ct268-30 =
cytology: Associated with T(l;2)ct268-l7 - Df(l)7B2-3;7C3-4.
T(l;2)7B2-5;41E2-4 (Hoover). *ct268-31
*cf268-18 origin: X ray induced.
discoverer: Demerec, 34i. discoverer: Demerec, 38d.
references: Hoover, 1938, Z. Induktive phenotype: Lethal. RK2A.
Abstammungs- Vererbungslehre 74: 420—34. cytology: Associated with T(l;3)ct2^S-3l -
phenotype: Lethal. RK2A. T(1;3)3D2-3;7B2-5;84D4-5;86B4-C1;88F (Hoover).
cytology: Associated with InCl^t269"1^ = *cf268-32
In(l)7B2-3;7B4-5;lW8-9. origin: X ray induced.
*cf268-20 discoverer: Demerec, 38e.
origin: X ray induced. phenotype: Lethal. RK2A.
discoverer: Demerec, 35g. cytology: Associated with T(l;'2)ct26S-32 =
references: Hoover, 1938, Z. Induktive T(l;2)lE-F;3D-E;7B2-5;46 (Hoover).
Abstammungs- Vererbungslehre 74: 420—34. *ct268-33
phenotype: Lethal and cell lethal. RK2A. origin: X ray induced.
cytology: Associated with In(l)ct26S"20 = discoverer: Demerec, 38e.
In(l)6Fll-7Al;7B5-6;l OBI 1-12. phenotype: Lethal. RK2A.
cytology: Associated with T(l;2)ct268~33 - cfK; cut of Krivshenko

T(1;2)7B2-5;41E (Hoover). origin: Spontaneous.
*cf268-35 discoverer: Krivshenko.
origin: X ray induced. references: 1956, DIS 30: 74.
discoverer: Demerec, 38k. phenotype: Both margins, as well as tips, of wing
phenotype: Lethal. RK2. are scalloped. Bristles of mesonotum, and
cytology: Salivary chromosomes normal (Sutton). especially scutellum, are fine as in Minutes.
*cf268-36 Bristle abnormality cannot be separated from wing
origin: X ray induced. effect by crossing over. Viability and fertility of
discoverer: Demerec, 39j. both sexes high. RK1.
phenotype: Lethal. RK2A. cytology: Salivary gland chromosomes appear
cytology: Associated with T(l;3)ct268-36 - normal.
T(1;3)7B2-C1;66F. ct": cut'notch
*cf268-37 origin: Recovered among progeny of heat-treated
origin: X ray induced. flies.
discoverer: Demerec, 39k. discoverer: Ives, 32c3.
references: 1940, Genetics 25: 618—27 (fig.). references: Plough and Ives, 1935, Genetics
Sutton, 1940, Genetics 25: 534-40. 20: 42-69.
phenotype: Lethal. RK2A. 1934, DIS 1: 31.
cytology: Associated with T(l;3)ct268-37 - phenotype: Wings notched at tips. Classification
T(l;3)5D2-3;7B2-3;80C-F. of males reliable, of females harder, but perfect
*cf268-38 at higher temperatures. Viability excellent. RK1
origin: X ray induced. in male.
discoverer: Demerec, 39k. *Cfn4
phenotype: Lethal. RK2. origin: Spontaneous.
cytology: Salivary chromosomes normal (Sutton). discoverer: Mischaikow, 58g.
*cf268-39 references: 1958, DIS 32: 83.
origin: X ray induced. phenotype: Wings notched at tips and inner margins,
discoverer: Demerec, 40a. similar to ctn. Excellent viability and fertility.
phenotype: Lethal. RK2. RK1.
cytology: Salivary chromosomes normal (Sutton). *Ctn63
*Cf268-40 origin: Spontaneous.
origin: X ray induced. discoverer: Datta, 63bll.
discoverer: Demerec, 39k. references: Sarkar, 1963, DIS 38: 28.
phenotype: Lethal. RK2A. phenotype: Wings cut to point and notched.
cytology: Associated with T(l;2;3)ct268-40 - Abdominal bands somewhat warped. Expression
T(l;2;3)7D2-3;10A5-6;21B-C;28-29;40-41;75B- more extreme in females than in males. Classifi-
C;87D;88C;92. cation, fertility, and viability excellent. RK1.
*cf268-4l *cts°: cut of Sytko
origin: X ray induced. discoverer: Sytko.
discoverer: Demerec, 391. references: Agol, 1936, DIS 5: 7.
phenotype: Lethal. RK2A. phenotype: Deeply notched wing tips. Good
cytology: Associated with T(l;2)ct268~41 = expression and viability. RK1.
T(l;2)7B2-5;37C2-3. *ctt: contorted
cf268-42 location: 1-0.3.
origin: X ray induced. origin: Induced by ethyl methanesulfonate (CB.
discoverer: Demerec, 40a. 1528).
phenotype: Lethal. RK2A. discoverer: Fahmy, 1956.
cytology: Associated with D((l)ct26S'42 = references: 1959, DIS 33: 84.
Dt(l)7A5~6;7B8-Cl. phenotype: Wings shorter than normal and abnor-
*Cfdo'vg: cut-dominigene for vestigial mally shaped; frequently curved either convexly or
discoverer: Goldschmidt. concavely. Eyes rough and slightly altered in
references: 1935, Z. Induktive Abstamrnungs- shape. Bristles thinner and straggly; orbitals
Vererbungslehre 69: 36—131 (fig.). frequently reduced or absent. Male genitalia
1935, Biol. Zentr. 55: 535-54. frequently slightly twisted and abnormal. Males
Gardner, 1942, Univ. Calif. (Berkeley) Publ. fertile and females sterile. RK2.
Zool. 49: 85-102. cu; curled
phenotype: ctdo~vif+ interacts with vg/+ to produce location: 3-50,0.
scalloped wings. RK3. origin: Spontaneous.
other information: Presumed by Goldschmidt to discoverer: Morgan, 15115.
enhance dominance of vg and thus termed a references: Morgan and Bridges, 1923, Carnegie
' 'dominigene.'' Inst. Wash. Publ. No. 327: 152 (fig.).
MUTATIONS 63
Morgan, Bridges, and Sturtevant, 1925, Bibliog. phenotype: Curled in combination with cu; homozy-
Genet. 2: 215 (fig.), 223. gous lethal. RK2.
Whittinghill, 1937, DIS 7: 22. cytology: Salivary chromosomes normal.
phenotype: Wings curved upward throughout length *Cu: Curl
and slightly divergent. Body color dark. Post- location: 2-55.2 (located using b and en).
scutellars erect and crossed. Good nutrition of origin: Spontaneous.
larvae enhances curled character as does high discoverer: Ives, 48e3.
temperature in last day of pupal life. (Nozawa, references: 1948, DIS 22: 53.
1956, Japan J. Genet. 31: 321-26). RK1. phenotype: Distal part of wing curled as in Cy;
cytology: Shown to be in region 86D2 through 87B2 proximal part to just beyond alula maintains
by its inclusion within the synthetic deficiency lateral compression and indentation fold of
with 3R proximal derived from T(3;4)86D = unfolded marginal wing. Usually an extra cross-
T(3;4)86D2-3;101F and 3R distal derived from vein beyond posterior crossvein extending across
T(Y;3) P102 = T(Y;3)87B2-3 (Cleland). L2, L3, and L4. Cu/+ and Cu/Cu identical; both
have good viability. RK1.
*CuA: Curl-Argentine
discoverer: Fernandez Gianotti.
synonym: Ac: Argentine Curly.
phenotype: Wings curled more strongly than Cy;
waxy texture. Homozygous viable; viability and
fertility good. RK2.
other information: Allelism inferred from similarity
in phenotype and genetic location (2-56.6) to Cu.
Cu-3: Curl in chromosome 3
location: 3-66.0.
discoverer: Erickson and Meyer, 51c.
synonym: Cur; Curl preoccupied.
references: Meyer, 1952, DIS 26: 66.
phenotype: Heterozygote has curly wings with
parchment-like texture resembling Cy. Homo-
zygous lethal. RK2.
cu-X: curled-X
cu: curled location: 1- (not located but not allelic to ex).
From Bridges and Morgan, 1923, Carnegie Inst. Wash. origin: Spontaneous in In(l)dl-49+BM1, y sc v.
Publ. No. 327: 152. discoverer: Krivshenko, 57j29.
*cu700.69 phenotype: Males have wings that are bent upwards
origin: X ray induced. and diverge slightly. cu-X is never expressed in
discoverer: Alexander. females. It represents a mutation whose pheno-
references: Ward and Alexander, 1957, Genetics 42: typic expression is sex limited. Expressed
42-54. equally well in males with and without a Y
phenotype: Like cu. RK1. chromosome. RK2.
cytology: Associated with T(l;3)cul 00.69 - cubitus interruptus: see ci
T(1;3)6B1-C1;88A4-B1. cuh curvi
location: 2-23.4 (1.4 to the right of Sp and 0.5 to the
*cu100.384 right of lys).
discoverer: Alexander. discoverer: Nicoletti.
references: Ward and Alexander, 1957, Genetics 42: synonym: curved.
42-54. references: 1957, DIS 31: 84.
phenotype: Curled in combination with cu; homozy- phenotype: Distal half of wing curved upward.
gotes semilethal. RK2. Viability and expressivity very good. RK1.
cytology: Salivary chromosomes normal. cupola: see cpl
cur: curvold
* c u 300.2J5 location: 3-30.
discoverer: Alexander. discoverer: Bridges, 33cl4.
references: Ward and Alexander, 1957, Genetics 42: phenotype: Wings divergent and curved down.
42-54. Resembles c. Viability erratic. RK3.
Cur: see Cu-3 cv-c: crossveinless-c

Curl: see Cu location: 3-54.1 (4.7 units to the left of sb<J2 by
curled: see co C. Hinton).
Curled blistered: see Cb origin: Spontaneous.
curl ex: see ex discoverer: Stern, 25gl3.
Curly: see Cy references: 1934, DIS 1: 35, 36.
Curlyoid: see Cyd phenotype: Posterior crossvein usually absent or
curved: see c greatly reduced. Anterior crossvein usually
curved: see cui present but often detached. Eye flattened or with
Curved of Krivshenko: see C-K vertical shallow furrow. Legs weak, especially
curvi: see cut tarsal joints. Occasionally overlaps wild type.
curvoid: see cur RK2.
cuf: see cf cytology: In region 88A through 88C, based on its
inclusion in the synthetic deficiency with 3R
proximal derived from T(Y;3)PJ02 = T(Y;3)87B2-3
and 3R distal derived from T(3;4)P86 =
T(3;4)88B-C;101 (Bernstein) as well as in the
duplication from T(1;3)O5 = T(1;3)4F2-3;62B-
C;88A-C;92C-D (Lindsley and Grell, 1958, DIS 32:
136).
cv'd: crossveinless-d
location: 3-65.
cv: crossveinless origin: Appeared among progeny of ether-treated
From Weinstein, A., 1920, Proc. Natl. Acad. Sci. U.S. flies.
6: 625-39. discoverer: Duncan, 34c.
cv: crossveinless phenotype: Posterior crossvein absent or reduced
location: 1-13.7. to an oblique fragment or bar parallel to L5.
origin: Spontaneous. Anterior crossvein sometimes detached. RK2.
discoverer: Bridges, 19112. other information: Possibly an allele of cv-b.
references: 1920, Proc. Natl. Acad. Sci. U.S. 6: *cvw; convex wing
660-63. location: 1-58.2.
Weinstein, 1920, Proc. Natl. Acad. Sci. U.S. 6: origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
625-39 (fig.). phenylalanine (CB. 3026).
phenotype: Crossveins absent or traces only discoverer: Fahmy, 1953.
present. Veins L3 and L4 slightly delta at tips. references: 1958, DIS 32: 69.
Classifiable in unexpended wings. Wing effects phenotype: Wings slightly shortened and arched
due to excessive contraction in the pupal period, convexly. Variable and may overlap wild type.
obliterating the cavity which should normally Tergites in some females have serrated edges or
remain between the epithelia to form the vein are grossly deformed. Viability and fertility good
(Waddington, 1940, J. Genet. 41: 75-139). RK1. in both sexes. RK2.
cytology: Salivary-chromosome studies by Demerec cx; cur/ex
and Sutton show locus to lie from 4F1-2 through location: 1-13.6.
5D1-2 inclusive (Demerec, Kaufmann, Fano, origin: Spontaneous,
Sutton, and Sansome, 1942, Carnegie Inst. Wash. discoverer: R. L. King, 1927.
Year Book 41: 191). Region can be narrowed to phenotype: Wings bent upward for posterior two-
4F9 through 5D2 on basis of inclusion of cv + in thirds of length; anterior one-third warped and
Dp(l;l)ybl « Dp(l;l)lB2-3;4F8-9;5D4-5 margin kinked. Wings not spread. RK2.
(Lindsley). cytology: Salivary analysis by Demerec and Sutton
cv-2: crossveinless on chromosome 2 shows that locus lies from 4F1-2 through 5D1-2
location: 2-96.2. inclusive (Demerec, Kaufmann, Fano, Sutton, and
origin: Spontaneous. Sansome, 1942, Carnegie Inst. Wash. Year Book
discoverer: Nicoletti, 62j. 41: 191).
phenotype: Anterior and posterior crossveins absent. cX: see c(3)G
RK1. cx*9; curl ex-twisted genitalia
cytology: Salivary chromosomes normal. origin: Spontaneous,
*cv-b; crossveinless-b discoverer: Curry, 37cl9.
location: 3-65. phenotype: Wings always divergent, usually 45°
origin: Spontaneous. from axis. Basal one-third of wing wavy, but less
discoverer: Bridges, 24k8. so than in cx; posterior two-thirds of wing curled
phenotype: Crossveins reduced or absent. May slightly upward or downward. Genitalia of nearly
overlap wild type. RK3. all males rotated, usually 45° counterclockwise.
MUTATIONS 65
Flies dwarfish. Viability irregular. Male sterile. d: dachs

RK2. location: 2-31.0.
cx-b: see wy2 origin: Spontaneous.
CxD: see In(3LR)CxD discoverer: Morgan and Bridges, 12k22.
CxF,D: see In(3LR)DcxF references: Bridges and Morgan, 1919, Carnegie
Inst. Wash. Publ. No. 278: 216 (fig.).
Cy: Curly Morgan, Bridges, and Sturtevant, 1925, Bibliog.
location: 2-6.1 (removed from In(2L)Cy and located Genet. 2: 212 (fig.), 223.
by Tinderholt). phenotype: Tarsi four jointed instead of five
origin: Spontaneous. jointed. Legs short and held close to body. Leg
discoverer: L. Ward, 20c. effects enhanced by s s a and ss*B (Villee, 1945,
references: 1923, Genetics 8: 276-300 (fig.)- Genetics 30: 26—27). Wings smaller than wild
phenotype: Wings curled upward; rarely overlaps type, narrowed, with L2 and L3 joined near
wild type at 25 , but frequently overlaps at 19 C. anterior crossvein; distance between crossveins
Curvature caused by the unequal contraction of smaller and crossveins sometimes absent. Angle
the upper and lower epithelia during the drying between L2 and L5 greater than normal. Eyes
period following emergence from the pupa case small and rough. Posterior scutellar bristles
(Waddington, 1940, J. Genet. 41: 75-139). Clas- erect. Viability erratic. Frequently sterile. RK2.
sifiable in single dose in triploids. Usually dU see Df(2L)d
homozygous lethal, but may emerge as dwarf with
more extreme wing character. RK1A.
cytology: Ordinarily inseparable from In(2L)Cy =
In(2L)22Dl-2;33F5-34Al, although it has been
separated by Tinderholt (1961, DIS 35: 47).
other information: Cy removed from In(2L)Cy still
causes a local reduction in crossing over in the
ed-cl region (Sederoff).
D: Dichaete
Publ. No. 327: 127.
D: Dichaete
location: 3-40.7 (40.4-41.0).
references: Bridges and Morgan, 1923, Carnegie
phenotype: Wings extended uniformly at 45 from
body axis and elevated 30° above (occasionally
Cy: Curly sharply downcast and dragging). Alulae missing.
From L. Ward, 1923, Genetics 8: 276-300. Dorsocentrals and some other bristles reduced in
number (Sturtevant, 1918, Carnegie Inst. Wash.
Cyd: Curlyoid Publ. No. 264; Plunkett, 1926, J. Exptl. Zool.
location: 3- (rearrangement). 46: 181-244). Head often deformed or split in
discoverer: Jollos. postvertical region. Halteres turned down. Homo-
references: Curry, 1939, DIS 12: 46. zygous lethal. Nearly lethal in combination with
phenotype: Wings curled upward in heterozygote. eyD (Sobels, Kruijt, and Spronk, 1951, DIS 25:
Homozygous lethal. RK2A. 128). Partially suppressed by sc alleles that re-
other information: Associated with an inversion of move postverticals (.sc, sf*. sc6, sc7) but not by
3R; possibly In(3R)P. others (sc^, sc$) (Sturtevant). Ciassifiahle in trip-
cy/: see rA'c>' loids. RK2A.
66 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTEFl
cytology: Inseparable from Jn(3L)D = In(3L)69D3- dark eye: see dke

El;70C13-Dl (Bridges in Morgan, Bridges and dark eye1: see sf3 2e
Schultz, 1937, Carnegie Inst. Wash. Year Book dark red brown: see drb
36: 301). Darkened eye: see Dke
darkener of w/i/fe-eos/n: see dark
origin: Spontaneous; derivative of D. darker legs: see thld
discoverer: Plunkett, 24f. darky: see dar
references: 1926, J. Exptl. Zool. 46: 181-244. daughterless: see da
phenotype: Less extreme than D. Wings extended db: dark body
and lifted; alulae missing. Head effect of D location: 3-44.4.
missing. Bristles usually wild type; occasionally origin: Spontaneous.
outer verticals, upper humerals, presuturals, and discoverer: Chovnick and Talsma, 1965.
anterior postalars absent. Viability of D^/+ references: 1966, DIS 41: 58.
better than D/+. RK2A. phenotype: Body color darker than normal. Male
cytology: In(3L)D present as in D. rarely survives; dies in late pupal stage. RK2.
*DE: Dichaete-Extended DcX: see ln(3LR)CxD
origin: Spontaneous in £>/+ culture; probably a DcxF: see In(3LR)DcxF
modified D. *dd: displaced
discoverer: Sturtevant, 16fll. location: 1-24.3.
references: Bridges and Morgan, 1923, Carnegie discoverer: Bridges, 31d7.
Inst. Wash. Publ. No. 327: 165 (fig.). phenotype: Antennae sunken into shortened head;
phenotype: Wings divergent but not bent at base. eyes also deformed. Females often sterile. RK2.
Bristles and alulae normal. Overlaps wild type. cytology: Locus lies between 7C4-5 and 8C1-2
Homozygous lethal; D/D^ is lethal. RK3(A). (Demerec, Kaufmann, Fano, Sutton, and Sansome,
da: daughterless 1942, Carnegie Inst. Wash. Year Book 41: 191).
location: 2-39.3. Further restricted to 7E1 through 8C2, on the basis
origin: Spontaneous. of its genetic location to the right of oc which is
discoverer: Bell, excluded from Df(l)sn = Df(l)7B2-3;7D22-El
references: 1954, Genetics 39: 958—59. (Hintonand Welshons, 1955, DIS 29: 125-26).
1954, DIS 28: 73. *ddh displacedlike
phenotype: Homozygous da females, mated to any location: 1-27.2.
male, produce normal sons but no daughters. origin: Induced by triethylenemelamine (CB. 1246).
Otherwise, da/da individuals appear wild type. discoverer Fahmy, 1953.
Lethal action of da occurs in egg stage. Counce synonym: dd$.
finds lethal female embryos show consistent references: 1959, DIS 33: 84.
abnormalities in midgut formation. In about half phenotype: Frontal region with antennae sunken
the abnormal embryos, shortening of germ band into shortened head. Eyes deformed. Thoracic
fails and anus and posterior spiracles open on bristles stiff and slightly shortened. Wings fre-
dorsal surface behind head segments. Differentia- quently misheld. Males sterile and viability
tion of almost all other tissues surprisingly slightly reduced. RK2.
normal. RK3. other information: One X-ray-induced allele.
cfe; see dar *de: deacon
dachs: see d location: 1-56.
dachsous: see ds origin: X ray induced.
*dar: darky discoverer: Muller, 26112.
location: 1-0 (no crossovers with sc in 547 flies). references: 1935, DIS 3: 29.
origin: X ray induced. phenotype: Body and wings narrow and rectangular.
discoverer: Fahmy, 1956. Eyes slightly flattened, with oblique cast. RK3.
synonym: da; preoccupied. other information: Possibly an allele of si (1-53.5).
references: 1959, DIS 33: 84, De; Dented
phenotype: Small, heavily melanized flies. Some- location: 2- (between dp and b).
times wings curl upward. Male sterile; viability origin: X ray induced,
about 15 percent wild type; late ©closing. RK2. discoverer: Belgovsky, 36c.
*dark: darkener of white-eosin references: 1937, DIS 8: 7.
location: Autosomal. phenotype: In heterozygote most flies show one or
discoverer: Bridges, 13i23. two indentations on thorax at front. Homozygote
references: 1916, Genetics 1: 148. has two smaller, sharper dents. Wings often
1919, J. Exptl. Zool. 28: 347. raised. RK3.
phenotype: Specific partial suppressor of we. RK3. deacon: see de
dark: see dk deep orange: see dor
dark body: see db defective: see df
dark bubbly: see dkb deflected wing: see dfw
MUTATIONS 67
Deformed: see Dfd folded flat at middle. Eyes sometimes rough and
deformed antennae: see dfa bulging. Wings slightly spread. Bristles tend to
deformed eye: see dfi break; scutellars occasionally doubled. RK3.
deformed terga: see dft Detached: see Dt
deformed wings: see dwg *df: defective
degenerated spermatheca: see dg-a location: 1-32.5.
Delta: see Dl origin: Spontaneous.
delta vein: see thvd discoverer: Bridges, 1513.
delta wing: see dta phenotype: Head bristles around ocelli missing.
deltex: see c/x Viability poor. RK3.
deltoid veins: see c//v *dfa: deformed antennae
Dented: see De location: 1-13.9.
*</ep: depressed origin: Induced by 2-chloroethyl methanesulfonate
location: 1-18. (CB. 1506).
discoverer: Bridges, 13d. discoverer: Fahmy, 1956.
references: Morgan and Bridges, 1916, Carnegie references: 1959, DIS 33: 84.
Inst. Wash. Publ. No. 237: 67 (fig.). phenotype: Wings short, broad, either convex or
phenotype: Wings turned down at tips, flat from concave, and abnormally held. Eyes small, dark,
side to side. Somewhat variable but does not and rough. Bristles short, stiff, occasionally bent.
overlap wild type. RK2. Trident pattern more pigmented. Abnormal
*depl: depressedlike antennae and aristae. Males viable and fertile.
location: 1-23. Females sterile. RK2.
origin: Recovered among progeny of flies treated
with Janus green,
discoverer: Muller, 28e20.
synonym: depr: depressed-roof.
phenotype: Wings droop at sides. Flies dark and
weak; bristles fine. Viability variable, about
20 percent wild type. RK3.
depressed: see dep
depressedlike: see depl Dfd: Deformed
*der: deranged From Bridges and Morgan, 1923, Carnegie Inst. Wash.
location: 1-57.2. Publ. No. 327: 94.
discoverer: Fahmy, 1953. Dfd: Deformed
references: 1958, DIS 32: 69. location: 3-47.5.
phenotype: Thoracic hairs deranged; many point origin: Spontaneous.
toward midline. Wings usually obliquely upheld discoverer: Cattell, 13g.
and twisted, bringing inner margins together. references: Bridges and Morgan, 1923, Carnegie
Overlaps wild type. Good viability in both sexes, Inst. Wash. Publ. No. 327: 93 (fig.).
but female fertility reduced. RK3. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
Genet. 2: 223 (fig.).
phenotype: Eye reduced ventrally and anteriorly, or
furrowed. Vibrissae tufted. Extremely variable,
often overlapping wild type, but usually excellent
character with Ly or at 19°C. Classifiable in
single dose in triploids (Schultz, 1934, DIS 1: 55).
Homozygous lethal, although Muller reported an
occasional homozygote as extreme dwarf with
flimsy wings. RK2.
det: detached cytology: Salivary gland chromosomes appear
From Bridges and Brehme, 1944, Carnegie Inst. Wash. normal.
Publ. No. 552: 54. *Dfd38k
det: detached discoverer: Mossige, 38k30.
location: 3-72.5. references: 1939, DIS 12: 47.
origin: Spontaneous. phenotype: Like Did. Homozygous lethal. RK2,
discoverer: Nichols-Skoog, 35k27. Dfdr; Deformed-recessive
phenotype: Posterior crossveins detached from origin: Spontaneous.
longitudinals at one or both ends and may be discoverer: Bridges, 3014.
absent. Wings occasionally folded back under or synonym: &m: almond.
phenotype: Eye small, narrow, and kidney shaped. dfw: deflected wing
Overlaps wild type in older cultures. Dfdr/Dfd location: 1-21.6.
more extreme than Dfd/+, RK2. origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
*Dfdr2 phenylalanine (CB. 3025).
origin: Spontaneous. discoverer:: Fahmy, 1955.
discoverer: Pierce. references: 1959, DIS 33: 84.
references: 1945, DIS 19: 46. phenotype: Wings slightly divergent and upheld to
phenotype: Eyes smaller and more constantly kidney various degrees, often twisted on their axes. Inner
shaped than in Dfdr. Wings thin, dull, uneven, margins frequently incised; occasionally wing
slightly spread (about 60° from body axis), and membranes separated by fluid. Eyes slightly
drooping. Body slightly smaller and lighter colored smaller. Males viable and fertile. Females
than normal. Bristles shortened and delicate. Last sterile; viability reduced. RK2.
abdominal segment of male may be rotated. Via- other information: One X-ray-induced allele.
bility low. Dtdr2/DfdT slightly more extreme than *dg-a: degenerated spermatheca
Dfdr/Dfdr. RK2. location: 3-75.5.
origin: Spontaneous. discoverer: Collins, 21a.
discoverer: Hollander, 1957. references: Wexelsen, 1928, Genetics 13: 389-400
references: 1960, DIS 34: 51. (fig.).
phenotype: Kidney-shaped eye. Penetrance and phenotype: Adult females show degeneration and
expressivity variable. RK2. pigmentation of epithelial cells of spermathecae
24 hr or more after eclosion. Viability and
origin: Spontaneous. fertility good. Penetrance 100 percent. RK3.
discoverer: Kidwell. dh: see eg2
references: 1961, DIS 35: 46. *di: dimorphos
phenotype: Eyes reduced. Expression varies from location: 1- (near spindle attachment).
absence of both eyes to wild type. Penetrance origin: Spontaneous.
varies from 75 to 100 percent. Penetrance discoverer: Harnly, 32dlO.
increased by selection for reduced eye. About 5 references: 1935, J. Exptl. Zool. 72: 75-99 (fig.).
percent of Dfdr60/+; ey/+ flies exhibit deformed 1940, DIS 13: 49.
phenotype. RK2. phenotype: Specific lengthener of vg wings, espe-
Dfdri-: Deformed-recessive of Luers cially in males (di; vg female much like vg). At
discoverer: Luers. higher temperatures, eyes small and rough and
references: Vogt, 1946, Experientia 2: 313-15. wings of both sexes approach wild type. RK2 in
1947, Biol. Zentr. 66: 81-105 (fig.)- vg male.
phenotype: Like Dfdr. RK2. dibrd: see frdi
*dfi: deformed eye Dichaete: see D
location: 3- (near D). dihedral: see eg2
origin: Recovered among descendants of heat-treated *dil: specific dilutor
flies. location: 2-57.
discoverer: Ives, 32c. origin: Spontaneous,
synonym: rough III. discoverer: Bridges, 32f22.
references: Plough and Ives, 1934, DIS 1: 34. phenotype: Dilutes bw to pale yellowish brown,
1935, Genetics 20: 42-69. and we, we2 and wb* to paler grades. RK3.
phenotype: Eyes roughish, reduced, and misshapen. *dil-3: dilute in chromosome 3
Overlaps wild type. Female sterile, poorly location: 3- (not located).
viable. RK3. discoverer: Bridges, 1519.
*dft; deformed tergi references: Bridges and Morgan, 1923, Carnegie
location: 1-33.7. Inst. Wash. Publ. No. 327: 151.
origin: Induced by 2-chloroethyl methanesulfonate phenotype: Eye color like maroon, overlaps wild
(CB. 1506). type. RK3.
discoverer. Fahmy, 1956, *c///-w°: dilutor of wft/fe-opricof
references: 1959, DIS 33: 84. location: 3- (not located).
phcnotype: Small fly with small, slightly rough discoverer: Weinstein.
eyes. Wings slightly divergent or upheld, abnor- references: Morgan, Bridges, and Sturtevant, 1925,
mally shaped with occasional incision of the inner Bibliog. Genet. 2: 218.
margin. Bristles slightly thinner and shorter with phenotype: Lightens » a . RK3.
one or both postscutellars frequently absent; and dilute in chromosome 3: see dil-3
a dorsocentral occasionally missing. Abdominal dilute ocelli: see po2
segmentation deformed to various degrees; Dilute-1: see
abdominal hairs fewer and deranged. Males poorly Dilute-2: see
fertile, viability about 50 percent wild type. RK2. Dilute-3: see
MUTATIONS 69
Dilute-4: see bwV30ki2 phenotype: Eye color soft, dull, and dark, like sf.
Dilute-5: see bwV3 0ki3 sf/dke is wild type. Flies have 65 percent normal
Dilute-6: see bwV30kis red pigment and 98 percent normal brown pigment
dilutor of white-apricot: see dil-wa (Nolte, 1955, J. Genet. 53: 1-10). RK2.
diminutive: see dm Dke: Darkened eye
dimorphos: see di location: 2- (not located).
Din: Dinty origin: X ray induced.
location: Unknown; associated with a rearrange- discoverer: Hendrix. 1963.
ment. references: 1964, DIS 39: 58.
origin: X ray induced. phenotype: In heterozygotes eye facets roughened
discoverer: Braver, 55a. with black-spotted pigmentation, varying from light
references: 1955, DIS 29: 70. spotting near margin of eye to heavy pigmentation
Pollock, 1963, DIS 38: 50. covering one-half of the eye. A bleached area
phenotype: In male and heterozygous female, sometimes appears adjacent to the pigmentation.
central portion of vein L2 interrupted. Posterior Effect usually symmetrical. Homozygous lethal.
supra-alar bristles absent in 95—99 percent of RK3.
females and 97—99.5 percent of males. Anterior cytology: Salivary chromosomes appear normal
postalar bristles absent in 6—11 percent of (Peacock).
females and 2—6 percent of males. Wings dkl: see thld
divergent. Viable and fertile in male and hetero-
zygous female; homozygous lethal. RK2.
cytology: Associated with T(l;2;3)Din =
T(1;3)3C;63A + T(2;3)39D;73A.
*dis: distorted eye
location: 1-23.
origin: Recovered among progeny of natural-gas-
treated fly.
discoverer: Mickey, 49b5.
phenotype: Whole or part of eye roughened. Some-
times bristles absent or doubled. Wings may be
roughened with nicked margins and plexus veins.
Expressivity variable. RK3.
cytology: Salivary chromosomes appear normal.
Discolored: see bwv?
dishevelled: see dsh
displaced: see dd
displaced!ike: see ddl
disrupted: see dsr
distorted eye: see dis
disturbed segmentation": see dss DI: Delta
divergent: see cfv From Bridges and Morgan, 1923, Carnegie Inst. Wash.
divergent wings: see dvw Publ. No. 327: 197.
divers: see dvr
*dk: dark DI: Delta
location: 3- (not located). location: 3-66.2.
discoverer. Clausen, 20g. origin: Spontaneous.
references: Bridges and Morgan, 1923, Carnegie discoverer: Bridges, 18k30.
Inst. Wash. Publ. No. 327: 235. references: Bridges and Morgan, 1923, Carnegie
Morgan, Bridges, and Sturtevant, 1925, Bibliog. Inst, Wash. Publ. No. 327: 197-201 (fig.).
Genet. 2: 223. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
phenotype: Eye color maroon. Overlaps wild type. Genet. 2: 75 (fig.).
RK3. phenotype: Veins thickened and broadened into
*dkb: dark bubbly deltas at junction with margin. Eyes somewhat
location: 2- (to the left of vg). small and slightly roughened. In extreme cases,
discoverer: Bridges, 38d25. ocelli run together into a crescent that encloses
phenotype: Thorax has dark bubbly longitudinal the ocellar bristles. Hairs on thorax straggly and
streak. RK3. more numerous. Body color dark. Wings small,
dke: dark eye dark, and somewhat spread. Effects of DI neu-
location: 2-73. tralized by H. DI and several of its alleles shown
origin: Spontaneous. to interact synergistically with certain Minutes,
discoverer: Bridges, 38cll. producing extreme phenotype and drastically
lowered viability (Schultz, 1929, Genetics 14: references: 1956, DIS 30: 71.
366-419). Homozygous lethal. RK2. MacDonald, 1957, DIS 31: 84.
cytology: Located in region 91D1-92A2, inclusively, phenotype: Spreading of veins most marked for L2
based on its inclusion within the transposed sec- and L5. Posterior crossvein thickened, coalesc-
tion of Tp(3)bxdU0 = Tp(3)89E2-3;91C7-Dl;92A2-3 ing with the delta of L5. L2 slightly thickened.
(E. B. Lewis) and in Df(3R)Dl» = D((3R)91C6- Expression more extreme at higher temperature,
Dl;92A2-3 (Slizynski). Heterozygous deficiency especially in males. Spreading of veins apparent
for 91D1-92A2 produces the Dl phenotype since in the 40- to 41-hr-old pupa. Suppressed by H.
Tp(3)bKd*l°/+ is D/ + , whereas Df(3R)bxdU0/ + j s Lethal homozygous and in combination with Dl3.
Dl. RK2.
cytology: Salivary chromosomes normal.
origin: Spontaneous. 1; Delta-Barish
discoverer: Bridges, 24110. discoverer: Schultz, 1933.
phenotype: Like Dl but deltas at margin are slight; phenotype: Delta venation and eyes of a broad
longitudinal veins between anterior and posterior heterozygous Bar type. Homozygous lethal. RK2A.
crossveins and crossveins themselves are thick- cytology: Associated with In(3R)DlB =
ened. Spreading of wing slighter than in Dl. In(3R)90A;91A (Schultz).
Better viability than Dl. Homozygous lethal. *DICf-3: Delta-Confluent
*DI4 discoverer: Imaizumi.
origin: Spontaneous. synonym: Cf-3.
discoverer: Bridges, 26g28. references: 1962, DIS 36: 38.
phenotype: Slight deltas at margin; posterior parts phenotype: Like Dl. Homozygote dies as late
of L2 and L3 thickened to delta at margin; L4 and embryo or early larva. RK2.
L5 slight. RK2.
other information: Gives increased crossing over origin: Induced by unspecified chemical (probably
throughout the third chromosome. mustard gas).
Dis discoverer: Auerbach.
discoverer: R. L. King, 32d. phenotype: Homozygous lethal. RK2A.
phenotype: Resembles Dl but deltas are slight. cytology: Associated with Dl(3R)DlH =
Wings occasionally vesiculated; only slightly Df(3R)91C6-Dl;92A2-3 (Slizynski).
spread. Homozygous lethal. RK3. *D/': Delta-lethal
other information: heterozygote shows reduced origin: Spontaneous.
crossing over. discoverer: Bridges, 38clO.
*DI6 synonym: 1(3 )DL
origin: X ray induced. phenotype: Heterozygote normal; homozygote not
discoverer: Schultz, 33a5. tested; thought to die in combination with Dl12.
phenotype: Extreme Dl. Veins at basal part of RK3.
wing thicker than Dl; thickening marked along other information: Presence of a recessive lethal
entire L2, with a striking confluent delta at allele of Dl on the H chromosome inferred from
margin; L3 has basal and distal thickening and failure to observe Dli2/H progeny from cross of
marginal delta; L4 extremely broad, especially h cu H2 ca/In(3R)P, Did ca X Dl^2/? A recessive
beyond posterior crossvein ending in large delta. lethal allele of H on the Dl*2 chromosome seems
Posterior crossvein moderately thick. Wings an equally likely interpretation.
blistered and wrinkled. Veins inhibited from *DlOf: Delta-Overflow
narrowing in late pupal period [Waddington, 1940, origin: Spontaneous.
J. Genet. 41: 75—139 (fig.)]. Homozygous lethal. discoverer: Tsukamoto, 1956.
RK2. synonym: Of.
origin: X ray induced, 1957, DIS 31: 85.
discoverer: Schultz, 33a7. phenotype: Slight deltas at margins; striking con-
phenotype: Like D/*. RK3. fluent effects on longitudinal veins, especially L2
and L5, and near posterior crossvein. Eyes nearly
*DI7P: Delto-7 of Pan shin normal, but with spac**, posterior half of eye
discoverer Panshin, 1935. surface resembles spaPot. Suppressed by H.
references: 1935, Dokl. Acad. Nauk SSSR 4: 85-88. Lethal homozygous and in combination with Dl12.
phenotype: Like Dl$. RK3A. RK1.
cytology: Associated with T(3;4)Dl^p. dlv: deltoid veins
location: 1-25.9.
origin: X ray induced. origin: Induced by S-2-chloroethylcysteine
discoverer: Clark. (CB. 1592).
MUTATIONS 71
discoverer: Fahmy, 1957. dor/dor and dor/+ females accumulate more

references: 1959, DIS 33: 85. isoxanthopterine than wild-type females, but dor
pbenotype: Wings small, abnormal, with margin males contain less than wild-type males. These
occasionally incised, and frequently either diver- differences detectable in prepupal stages. Ommo-
gent or slightly upheld. Extra venation, especially chrome pigments also affected. Reciprocal trans-
at junctions between longitudinal and costal plantation experiments show that eye color is
veins, giving Delta-like formations. In extreme autonomous (Hadorn and Counce). Homozygous
cases, wings grossly deformed and blistered. dor females exhibit same sterility effects as tu
Excess melanization throughout body. Eyes dark, (Merrell, 1947); i.e., dor progeny of dor mothers
small, and slightly rough. Total body size die. dor females crossed with wild-type males
reduced. Both sexes viable and fertile. RK1. produce some dor/+ daughters. Under certain
other information: One allele induced by CB. 1592. conditions dor males sterile as result of excess
dm: diminutive accumulation of preadult fat, which mechanically
location: 1-4.6. prevents union of gonads with rudiments of rest of
discoverer: Nichols-Skoog, 33j9. genital system (Counce). Post-blastulation devel-
references: 1935, DIS 3: 10. opment of dor progeny of dor females abnormal,
phenotype: Bristles and body small and slender. leading to embryonic death (Hildreth and
Viability excellent. Females sterile. RK1. Lucchesi). Cellular degeneration begins before
cytology: Locus placed between 3C9 and 3D2 by gastrulation is ended, and by 16 hr, degeneration
Slyzinska (1938, Genetics 23: 291—99), at 3C9 by of embryo is almost complete. Some embryos die
Schultz, and at 3D1-2 by Demerec, Kaufmann, at early cleavage regardless of sex. Eggs of dor
Fano, Sutton, and Sansome (1942, Carnegie Inst. females contain less than normal amounts of yolk
Wash. Year Book 41: 191). (Counce, 1956). The double mutants dor; ry, and
dm264-S8 dor, ry2 are lethal (Luchessi, 1963, Proc. Intern.
origin: X ray induced. Congr. Genet, 11th. Vol. 1: 169-70). RK1.
discoverer: Demerec, 38d. cytology: Placed in region from 1F1 through 2A2
references: 1940, Genetics 25: 618-27. on the basis of its inclusion in Dp(l;f)1337 =
phenotype: Described only as nonvariegated. RK2A. Dp(l;f)lF4-2A3; 19-20 but not in Dp(l;f)112 =
cytology: Associated with T(1;3)N264-58 ~ Dp(l;f)lE4-Fl; 19-20 (R. F. Grell, Gersh).
T(l;3)3B2-3;3D6-7;80D-F (Sutton).
dn: doughnut origin: Spontaneous.
location: 3-50. discoverer: Hildreth, 61e.
discoverer: Wallbrunn. phenotype: Orange eye color. Poor viability. RK2.
Wright, 1946, DIS 20: 68. dor1: deep orange-lethal
phenotype: Eye of se dn has unpigmented spot (in origin: Spontaneous.
middle or toward posterior) at emergence from discoverer: Bridges, 15al.
puparium. Spot gradually darkens; after 2 days synonym: 1(1)7: lethal(l) 7.
eyes appear sepia. Difficult to detect with wild- references: 1916, Genetics 1: 149.
type eye color; appears as slightly lighter red phenotype: Male larvae die 90—lOOhr after
spot, which disappears after 2 days. Viability hatching. According to Russell [l940, J. Exptl.
low; many die as pupae at 25°C. Viability nearly Zool. 84: 363—79 (fig.)], a gut abnormality appears
normal at 17°C, but character not detectable. at 65 hr resulting in obliteration of gut lumen and
Both sexes highly infertile; testes about one-third loss of all food material. Stark observed me Ian otic
normal length. Spermathecae very small. External tumors [1918, J. Cancer Res. 3: 279-301 (fig.);
genitalia of both sexes often abnormal. RK3. 1919, J. Exptl. Zool. 27: 509-29 (fig.)]. Oftedal
dd: see po2 studied histology of dor1 larvae (1953, Z. Induktive
dor: deep orange Abstammungs- Vererbungslehre 85: 408—22).
location: 1-0.3 [allelic to dor1 (Redfield and Malpighian tubes nearly colorless except at base.
Schultz; Clancy)]. dor1/'dor has orange eyes like dor/dor as well as
origin: X ray induced. the sterility effects (Clancy; Redfield and
discoverer: E. D. King. Schultz). RK2.
references: Merrell, 1947, Am. Naturalist 81:
399_400. origin: X ray induced.
Counce, 1956, Z. Induktive Abstammungs- discoverer Alikhanian.
Vererbungslehre 87: 443-61 (fig.). synonym: 1(1)76.
phenotype: Eye color true orange at 25°C, red- references: Ardoshnikov, 1941, Dokl. Akad. Nauk
orange at 18°. Chromatographic studies (Counce, SSSR 30: 344-46.
1957, Experientia 13: 354) indicate pteridine phenotype: Like dor1. Incidence of dor!2/Y larvae
patterns differ from wild type. Red pigment bearing melanotic masses reduced if one parent
reduced (but increased in heterozygous females); carries a free duplication carrying dor*. RK2.
two, or all of these phenotypes. For example:

origin: Spontaneous. dp° (o) has oblique wings, but no vortices, and is
discoverer: H. W. Lewis. not lethal; dpov (ov) has oblique wings and
references: 1954, J. Exptl. Zool. 126: 235-75 (fig.). vortices, but is not lethal; and dpol,v (olv) has all
phenotype: Like dorK Survival of larvae containing three attributes. The three recessive character-
melanotic masses optimal at 25°C. dorI3/Basc fe- istics o, I, and v complement one another. Thus
male shows more rapid development and higher via- dpol/dpv, dp°v/dpl, and dp°/dplv are phenotyp-
bility than +/Basc female. RK2. ically wild type; dp°/dp°", dpol/dp°v, and
*double: double dp°/dp°lv are o; dp°v/dpv, dplv/dp°v, and
location: 1-0. dpv/dpolv are v; any two lethals are lethal in the
origin: Spontaneous. trans heterozygote.
discoverer: Bridges, 1918, cytology: Located between 24E2 and 25A2 based on
references: Morgan, Bridges, and Sturtevant, 1925, its inclusion in Df(2L)M-zB = Df(2L)24E2-
Bibliog. Genet. 2: 224. F1-25A1-2 (Morgan, Bridges, Schultz, and Curry,
phenotype: Postvertical bristles doubled. Wings 1939, Carnegie Inst. Wash. Year Book 38: 273-77).
very small. Viability somewhat low. RK3. other information: The alleles at the dp locus are
Double Bar: see BB pseudoallelic and have been positioned by Carlson
Double Inhabar: see B'Bi (1959) and Southin and Carlson (1962) (see map).
double sex: see dsx Carlson often puts the superscripts on the line and
Doublet: see Dp(l;l)BSRMG has used an apostrophe instead of dp; dp o / " may
doughnut: see dn be written, therefore, as olv or 'olv.
Im
olm
olvW
olvbm
obm o2 olv cm2 Iv h ov ovS2b olS v2
1 .—• 1 1 1
Map of the dp locus

From Southin and Carlson, 1962, Genetics 47: 1017-
26.
dp: see dp°v

dp*: see dplv?

discoverer: Fogel, 1949.
other information: A series of 13 alleles of diverse
phenotype. Viability data given (Fogel, 1950).
See
rfow: downy dps*1: see

From Bridges and Brehme, 1944, Carnegie Inst. Wash. dpS2b; see
Publ. No. 552: 64. *dpS8
dow: downy discoverer: Fradkin, 1958.
origin: Spontaneous. other information: A series of 14 alleles; not
discoverer: Bridges, 36c28. described or tested for viability.
phenotype: Bristles very short and slender, nearly dp61d
as small as as. Males entirely sterile; testis origin: X ray induced.
shape normal. Viability good, RK2. discoverer: Thompson, 61d.
dp; dumpy cytology: Associated with T(Y;2)dp(>1 d.
location: 2-13.0. b see dpobw
references: Carlson, 1958, Ph.D. Thesis, Indiana *c/pcm: dumpy-comma
Univ. origin: Spontaneous.
1959, Genetics 44: 347-73 (fig.1. discoverer: Bridges, 13b5.
Seuthin and Carlson, 1962, Genetics 47: 1017-26 references: Bridges and Morgan, 1919, Carnegie
phenorype-: Alleles of dp produce three general phenotype: Sharply outlined depressions of comma
phermtypes: oblique wings Co), vortices on thorax shape at anterior edge of thorax. Penetrant in 20
O), and lethal CO. A specific mllele may have one, percent of females but rarely in males. RK3.
MUTATIONS 73
<Jpcm2
effect in dp!^/-h; Me/+ and dpIM +/+ ta. Pheno-
origin: Spontaneous. typically normal in combination with dp°2( dpv?,
discoverer: Meyer, 53c. dp o v , and dpc™2. RK2.
references: 1955, DIS 29: 74. dpL': see
phenotype: Homozygotes nearly normal at 25°C,
with slightly oblique wings in some. Anterior dpLS: see
edge of thorax usually shows a pair of commalike
depressions; wings may occasionally have
blisters. dpcm2/dpov shows good commas in all dpi*: dumpy-lethal vortex
flies, vortices in most, and slightly oblique origin: Spontaneous.
wings. dp<>™2/dpolv is semilethal at 22°C, and discoverer: Bridges, 20c3.
survivors are similar in phenotype to dpov/dpolv. synonym: dptx: dumpy-thoraxate.
Degree of viability of dpcm2 when heterozygous phenotype: Homozygous lethal; no abnormalities
with various dp °'"-like alleles varies, but is when heterozygous witti wild type. dplv/dpov has
usually lower at low temperatures (Carlson and strong volcanolike dorsocentral mounds or pits
Falk, 1962, DIS 36: 59-61). RK3. with brown pigmentation; comma effects striking;
sternopleural bristles turned down; wings normal.
dpD: dumpy-Dominant At higher temperature (28°C), two additional pits
origin: X ray induced. anterior to dorsocentrals occasionally appear.
discoverer: E. B. Lewis, 1962. Lethal in compound with dpo1, dpo1^, and dplM%
synonym: olvD. RK2.
references: Del Campo, 1963, DIS 38: 32. dpi* *
phenotype: dpD/+ has slightly oblique wing and origin: Spontaneous within In(2L)Cy.
moderate vortex and comma effects on thorax. discoverer: Muller.
dpD/dp°v has strong wing and thorax effects, with synonym: dp2.
reduced leg and body size. dpD/dpolv is lethal, phenotype: Similar to dpIv; slight oblique wing
and dpD resembles dpo!v in several ways. effect in dplv2/dpov; comma effects occasionally
dpD/dpv has strongly enhanced thorax effects, seen in d p ^ 2 / + . R K 2 .
and dp°/dpD has reduced wings with possibly *c/p/v57e
enhanced thoracic effect. Homozygous lethal. origin: Ultraviolet induced.
RK2A. discoverer: Meyer, 51e.
cytology: Associated with T(2;3)dpD = synonym: dp^"^5Je: dumpy-thoraxate 51e.
T(2;3)25A;95B-D. references: Meyer and Edmondson, 1951, DIS 25: 72.
dpdei: s e e r f p o d e f phenotype: Similar to dplv. Occasionally
dpdr: see dpovdr dplv51 e/dpov shows blistering of wings but no
*dpG; dumpy of Goldschmidt oblique truncation. RK2.
discoverer: Goldschmidt. dplvl: dumpy-lethal vortex of Ives
references: 1945, Univ. Calif. (Berkeley) Publ. origin: Spontaneous within In(2t,)Cy.
Zool. 49: 511, 520. discoverer: Ives, 39j.
other information: Several alleles; not analyzed in synonym: dptxî dumpy-thoraxate of Ives;
detail for effect.
dph: dumpy-humpylike phenotype: Identical to dplv. RK2.
origin: Spontaneous, dpNov: see
discoverer: Edmondson, 54g.
phenotype: As homozygote, has strongly truncated
wings, four marked thoracic humplike vortices,
strong comma effect, reduced body size, and
small, weak legs. Because of weak legs, flies
have difficulty walking and become stuck in the
food. No body or leg reduction in dph/dpv2 but
vortices and commas present as in dpv2/dpolv. dp°: dumpy-oblique
dph viable with dpo'S, dp°lv, dplv, and dp1^, From Bridges and Brehme, 1944, Carnegie Inst. Wash.
RK2. Publ. No. 552: 65.
dpH: see dpoivH
dplM: dumpy-lethal of Meyer dp°: dumpy-oblique
origin: Ultraviolet induced. origin: Spontaneous.
discoverer: Meyer, 57f. discoverer: Bridges, 17i24.
references: 1958, DIS 32: 83. phenotype: Wings obliquely truncated but nearly
phenotype: Homozygous lethal and lethal in combi- full length; overlaps wild type at 25°C when
nation with dpo!S, dp 0 ' 1 ', and dplv. Slight oblique homozygous and in combination with dp o v . RK3.
dpo2 reduction in females and slight reduction in about

origin: Spontaneous. half the males. RK3.
discoverer: Laemmerts, 1926. *dp°h dumpy-oblique lethal
synonym: clip wing. origin: Spontaneous in dp°.
phenotype: Wings truncated and shortened as in discoverer: Muller, 1919.
dpov, but without vortex or comma effects. No synonym: dpL: dumpy-Lopped.
overlap with wild type as homozygote. Over- phenotype: Homozygous lethal. dpol/dpov shows
lapping frequent at 25°C in compound with dp°v. small body and legs and drastically reduced
Mutant effect more pronounced in females than in wings; thorax normal but occasional comma
males in dpo2/dpov; sexual dimorphism less ex- effects at 26°C or higher; no vortex effect. dp°l/+
treme in dp°2/dp°2. RK1. occasionally shows wing effect. RK2.
*dpoS0c dpotM; dumpy-oblique lethal of Meyer
origin: Spontaneous. origin: Ultraviolet induced.
discoverer: Sobels, 50c7. discoverer: Meyer, 51b.
synonym: dp5°c. synonym: dpLM: dumpy-Lopped of Meyer; dpTSlb:
references: Sobels, Boterenbrood, Faber, and Truncate*1*.
Oppenoorth, 1951, DIS 25: 76. references: 1952, DIS 26: 66.
phenotype: Manifests oblique wing only when Me phenotype: Homozygous lethal. Similar to dp°l but
present. Best penetrance at 25°C; less at 28°C has occasional slight vortex effect and good
and none at 16°C. Manifestation better in females comma effect in compound with dp°v. RK2.
than males. Tests of allelism with dpov have dp°IS; dumpy-oblique lethal of Schalet
been inconclusive. RK3. origin: Spontaneous.
dpoSU discoverer: Schalet, 1955.
origin: Ultraviolet induced. synonym: dpLS; dumpy-Lopped of Schalet; dpLSch.
discoverer: Byers and Meyer, 51e. references: Carlson and Schalet, 1956, DIS 30: 70.
references: Meyer and Edmondson, 1951, DIS 25: phenotype: Homozygous lethal. Heterozygote
72. dp°l$/dpov shows small body and legs and
phenotype: Slight wing effect in dp°v/dp°Sl«; no drastically reduced wings; thorax normal but
vortex effect. Wing effect in dp°Sle/dp°lv o n i y occasional comma effects at 26°C or higher; no
slightly greater. RK3. vortex effect. Lethal with dp°tv, dplv, and dp'M.
Jpobm; dumpy-oblique from bilateral mosaic RK2.
origin: X ray induced. dp°ly: dumpy-oblique lethal vortex
discoverer: Carlson, 1957. origin: Spontaneous.
references: Carlson and Southin, 1959, Genetics discoverer: Morgan, 1923.
44: 502-3. synonym: dpT: dumpy-Truncate.
phenotype: Inviable as homozygote, perhaps from references: Altenburg and Muller, 1920, Genetics
independent lethal nearby, because heterozygotes 5: 1-59 (fig.).
with dpoIv, dp°l, and dp1* are viable. Heterozy- phenotype: Homozygous lethal. dpolv/dpov has
gous dpov/dpobm has oblique wing, slightly reduced body and leg size; wings shortened to
reduced body size, and comma effects, but normal half length and obliquely truncated; thorax shows
dorsocentral region. RK2. two and sometimes four vortices; comma effect
other information: Arose in same fly as dpolvbm; strong; stemopleural bristles turned down; wings
dpolvhm/dpobm i8 virtually lethal. often elevated 45° and sometimes contain black
*dpobw: dumpy-oblique brown blisters. Body reduction also seen in compound
origin: Spontaneous, with dp°2 but not with dpv2. Lethal in compound
discoverer: Honer, 1939. with dp°ts, dptMt a n d dp/v, R K 2 .
synonym: dpbw- dtxmpy-brovm.
references: 1939, Z. Induktive Abstammungs- origin: Spontaneous.
Verebungglehre 77: SOI-15 (fig.). discoverer: Muller, 1913.
phenotype: Homoxygous females have moderately synonym: dpT2f x2.
truncated wings, but males only slightly affected. references: Altenburg and Muller, 1920, Genetics
dpQlvH/dpobw females have reduced body and leg 5: 1-59 (fig.).
size, but males almost normal; variable expression phenotype: Similar to dp°lvM. RK2.
in wing size. Some wings pointed in females. RK3. *dpotv54d
*dp»d*f: dumpy-oblique deformed origin: Ultraviolet induced,
origin: Spontaneous, discoverer: Meyer, 54d.
discoverer. Honer, 1939. synonym: dpT54dw
synonym: dp^®1; dumpy-deformed. references: 1955, DIS 29: 74.
references: 1939, Z. Induktive Abstammungs- phenotype: Like dp°'*. RK2.
Vererbungslehre 77: 501-15 (fig.), *dpolvSSb
phenofype: Males normal, female wings slightly origin: Spontaneous.
oblique. dp@de(/dp°lvH shows moderate wing discoverer: Meyer, 55b.
MUTATIONS 75
synonym: dpTSSb. c/po'vR; dumpy-oblique lethal vortex Ruffled

phenotype; Similar to dpolv, RK2. discoverer: Schultz, 33a25.
*dpolv55c synonym: dp&t-. dumpy-Ruffled.
origin: Ultraviolet induced. phenotype: Similar to dpoly but heterozygous
discoverer: Meyer, 55c. dpolvR has slightly spread wings with uneven
synonym: dpTSSc. surface and ruffled margin. Raffling effect over-
references: 1955, DIS 29: 74. laps wild type at 25°C, RK2A.
phenotype: Like dpolv. RK2. cytology: Associated with In(2L)dpolvR =
*dpolvS7g In(2L)25A;25B3-4.
origin: Ultraviolet induced, dpoIvS; dumpy-oblique lethal vortex of Schalet
discoverer: Meyer, 57g. origin: Neutron induced.
synonym: dpr57g# discoverer: Schalet, 1955.
references: 1958, DIS 32: 78. synonym: dpTSch. dumpy-Truncate of Schalet.
phenotype: Like dpoJv. RK2. references: Carlson and Schalet, 1955, DIS 29: 71.
phenotype: Similar to dpoIv, RK2.
dpolvbm; dumpy-oblique lethal vortex from other information: Arose simultaneously with
bilateral mosaic mutation to y in Dp(l;2)sc1 9, but ed, cl, and
origin: X ray induced. dw-24F not affected; possibly a minute inversion.
discoverer: Carlson, 1957. *c/po'vSn: dumpy-oblique lethal vortex Snub
references: Carlson and Southin, 1959, Genetics origin: Spontaneous.
44: 502-3. discoverer: Bridges, 16bl7.
phenotype: Like dpotv. RK2. synonym: dpTs: dumpy-Truncate Snub.
other information: Recovered from same fly that references: Bridges and Morgan, 1919, Carnegie
produced dpobm. Inst. Wash. Publ. No. 278: 140.
*dpolvD: dumpy-oblique lethal vortex of Duncan phenotype: Similar to dp°lvM. RK2.
origin: Spontaneous. c/po/vW; dumpy-oblique lethal vortex of Williams
discoverer: Duncan, 1914. origin: Spontaneous.
synonym: dpTD: dumpy-Truncate of Duncan. discoverer: Williams, 1957.
references: 1915, Am. Naturalist 49: 575-82. synonym: dpTW: dumpy-Truncate of Williams.
phenotype: Similar to dp°'vM. RK2. references: Carlson, 1956, DIS 30: 70.
*cfpo/vH; dumpy-oblique lethal vortex of Hb'ner phenotype: Similar to dp°tv. RK2.
origin: Spontaneous, *dpou: dumpy-oblique ultraviolet
discoverer. Hb'ner, 1931. origin: Ultraviolet induced.
synonym: dpH: dumpy of Honer, discoverer: Meyer, 54g.
references: 1939, Z. Induktive Abstammungs-
Vererbungslehre 77: 501-15 (fig.). phenotype: Similar to dp°2. RK2.
1939, DIS 11: 45-46.
phenotype: Similar to dpolv. dp°IvH/+ may show
some wing truncation. RK2.
*t/po/vM; dumpy-oblique lethal vortex of Morgan
discoverer: Morgan, lOh.
synonym: dpTO. dumpy-Truncate zero.
Altenburg and Muller, 1920, Genetics 5: 1—59
(fig.).
phenotype: dp°lvM/+ showed oblique truncated
wings in up to 90 percent of flies in maximally
selected lines. RK2.
other information: Altenburg and Muller established
that temperature and modifier genes affect a major
gene at 2-13.0. Later tests with dpo*, dpy, dp°
and other alleles established similarity to dp°lv.
*e/po'vP; dumpy-oblique lethal vortex of Plough
dpov: dumpy-oblique vortex
origin: Recovered among progeny of temperature-
shocked parents.
discoverer: Plough, 40cl5.
synonym: dpTP', dumpy-Truncate of Plough.
t/pov; dumpy-oblique vortex
phenotype: Like dpotr, RK2.
discoverer: Morgan, 18jl6.
synonym: dp. dpy: dumpy-vortex

references: Morgan, 1929, Carnegie Inst. Wash. origin: Spontaneous.
Publ. No. 399: 196. discoverer: Bridges, 1919.
phenotype: Wings reduced one-third, with oblique references: Bridges and Mohr, 1919, Genetics 4:
truncation; angle between veins L2 and L3 283-306 (fig.).
widened and intercrossvein distances shortened; Bridges and Morgan, 1923, Carnegie Inst. Wash.
marginal vein intact; two vortices with whorls of Publ. No. 327: 168 (fig.).
hairs in dorcentral region; dorsocentral bristles phenotype: Homozygote normal; expresses vortices
medially displaced; vortex expression in most only when e(dpv) on the third chromosome is also
flies at 26°C; comma effect in anterior thorax also homozygous. Heterozygotes with dpv2, dp°v,
present in most flies at 26°C. At lower tempera- dplv, and dpolv have good vortices without e(dpv).
tures, wing effect persists but thorax appears RK3.
normal. Body and leg size normal except at
temperatures above 28°C, when they are somewhat origin: Spontaneous.
reduced. RK1. discoverer: Mohr, 20a30.
dpov51f references: 1923, Studia Mendeliana (Brunae)
origin: Spontaneous, pp. 266-87.
discoverer: Mossige, 1951. 1929, Z. Induktive Abstammungs- Vererbungslehre
synonym: dpsli, 50: 113-200.
phenotype: Like dp°*. RK1. phenotype: Thorax shows two vortices in dorso-
Jpev52b central region, frequently pitted with brown
origin: Spontaneous. pigment. Whorls of hairs and medially displaced
discoverer: Edmonds on, 1952. dorsocentral bristles in almost all flies at 26°C.
synonym: dp^^b^ Females more extreme than males. No wing
references: 1952, DIS 26: 60. effect, no body or leg reduction in any compound.
phenotype: Like dp°*. RK1. Commas absent in dpv2/dpv2 and dpv2/dpov, but
present in dpvl/dp1* and dpv2/dp°lv. RK2.
dpovdr; dumpy-oblique vortex drumlins ; dumpy-vortex of Muller
origin: Spontaneous in dpov. origin: Spontaneous.
discoverer: R. C. King, 48b26. discoverer: Muller, 1920.
synonym: dp*; dumpy-dtumlins. synonym: dpvl; dumpy-volcano.
references: Poulson and King, 1948, DIS 22: 54. phenotype: Homozygote has vortices, but less well
phenotype: Has a more pronounced vortex effect developed and less numerous than in dpy2.
than dp°y but otherwise similar. Viability slightly Overlaps wild type. RK2.
reduced. RK1. dp**: dumpy-vortex of Vfaddington
other information: Probably a modifier in or near origin: Spontaneous.
the dumpy locus. discoverer: Waddington.
synonym: dpvo.
dpovN; dumpy-oblique vortex of Novitski phenotype: Similar to dpv2. RK2.
discoverer: Novitski. dpwh dumpy-warped
v
•ynonym: dpN° : dumpy of Novitski. origin: X ray induced.
phenotype: Like dpov. RK1. discoverer: Schalet, 1955.
dp^h see references: Carlson and Schalet, 1955, DIS 29: 71.
dpT; see Carlson, 1958, DIS 32: 117-18.
dpT0: see phenotype: Heterozygote of dpov/dpwl has variable
dpT2; see and asymmetrical expression of vortex and oblique
dpTsib: s e e wing effects. Mutant phenotype completely sup-
dpTS4d: pressed by an extra Y chromosome. Homozygous
dpT55b; lethal but viable in compound with lethal dp
see alleles. dpwl/dptvl have wrinkled opaque wings.
see dpotvsrg RK2A.
see d cytology: Associated with T(2;3)dpwl (breakpoints
not determined).
other information: Appears to be a variegated
dpi position effect.
c/pw2
see
dpTS: see origin: X ray induced.
discoverer: Schalet, 55k.
references: Carlson and Schalet, 1956, DIS 30: 71.
dp**: see c Carlson, 1958, DIS 32: 117-18.
dp**': see phenotype: Similar to dp*!. RK2A.
MUTATIONS 77
cytology: Associated with T(Y;2)dp*2 (break appearance; constant expression. Lethal homozy-
points not determined), gous but survives in combination with DrWe (E. B.
Dp: see Dr Lewis). RK1.
*dpy: dumpoidy other information: DrJAio/DrWe yields rare wild-type
locotion: 3- (right arm near 90). recombinants (Lewis).
origin: Spontaneous. DrWe: Drop-Wec/ge
discoverer: Villee, 40a. origin: Spontaneous.
phenotype: Wings obliquely truncated and reduced discoverer Muller.
in length; marginal vein intact. No vortices or references: 1965, DIS 40: 36.
whorls of bristles on thorax. Suppressed by Cy phenotype: Eyes are small vertical wedges with
and Gla; made dominant by Me, Overlaps wild points downward; much smaller than homozygous
type slightly. RK2. 23. Lethal homozygous and in combination with Dr,
d<j: see Ld(i but DtWe/DiMio survives (E. B. Lewis). RK1.
*dr: droopy other information: Drê/Drîo yields rare wild-type
location: 2-71.2 (to the left of Lobe). recombinants (Lewis).
origin: Ultraviolet induced. drb: dark red brown
discoverer: Meyer and Edmondson, 49c. location: 3-47.7 (may be rearrangement; st—p
references: 1949, DIS 23: 60. crossing over 50 percent of normal),
phenotype: Wings spread wide apart and droop origin: Spontaneous.
downward, like c, often crumple and drag in the discoverer: Rosin, 48b.
food. Alula broad and short. Viability at hatching references: 1951, DIS 25: 75.
fair; females tend to die before males. Penetrance phenotype: Eye color dark red-brown at 18° and
100 percent. Fertility good. RK2. dark red at 28°C. drb/+ darker than wild type at
other information: Not a lie lie to c, 18°, but not at 28°C. RK2(A).
dt: see drw droop wings: see drp
Dr: Drop droopy: see dr
location: 3-99.2. droopy wing: see drw
origin: X ray induced. Drop: see Dr
discoverer: Krivshenko, 54c25. *drp: droop wings
synonym: Dp. location: 1- (rearrangement),
references: 1954, DIS 28: 75. origin: Spontaneous,
phenotype: Heterozygote has 1—10 eye facets, discoverer: Ives, 48f.
which appear dark red. Viability of heterozygote references: 1949, DIS 23: 58.
excellent. Homozygous lethal. RK1. cytology: Associated with In(l)drp - ln(l)12B;20B.
cytology: No visible rearrangement in salivary *drw: droopy wing
chromosomes, location: 1-52.3.
other information: Recombination between Dr*1*0 and origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
DrWe indicates that Dr is a pseudoallelic locus phenylalanine (CB. 3025).
(E. B. Lewis). discoverer: Fahmy, 1953.
*DrA: Drop of Abrahamson synonym: Symbol originally dr, which was
origin: X ray induced. preoccupied.
discoverer: Abrahamson, 60d28. references: 1958, DIS 32: 69.
synonym: DpA. phenotype: Small fly with drooping wings. Chitin
references: Abrahamson and Siegel, 1960, DIS 34: of abdomen irregularly ridged and pigmented.
48. Hairs deranged. Males infertile; viability 10 per-
phenotype: Facet number of heterozygote averages cent wild type. RK3.
four. Homozygote viable and usually lacks eye ds: dachsous
facets. RK1. location: 2-0.3.
*DrL: Drop of Lewis origin: Spontaneous,
origin: X ray induced, discoverer: Bridges, 17kl2.
discoverer. E. B. Lewis. references: Stern and Bridges, 1926, Genetics 11:
phenotype: Like Dr. RK1A. 511.
cytology: Associated with T(2;3)DrL = Mohr, 1929, Z. Induktive Abstammungs-
T(2;3)44;89F-90A + In(3R)89C;95D-96Bl, which is Vererbungslehre 50: 113-200 (fig.),
probably independent of the mutation. phenotype: Wings shorter, blunter, and broader,
>: Drop-Microp/ifhalmia with crossveins uniformly very close together.
origin: Mustard gas induced, Abdomen and legs chunky. Slight dominance of
discoverer: Sobels, 57J22. close crossveins. Strong interaction with d, ij,
synonym: Mlo. and eg; double homozygotes often have excessive
references: 1958, DIS 32: 84. growth of thoracic parts and sometimes conversion
phenotype: Drîo/+ show extreme reduction in eye of one organ into another [e.g., twinning of wings
size; facets coalesce to give a shiny, dark red or antennae or conversion of eyes into antennae,
Waddington, 1943, J. Genet., 45: 44-50 (fig.)]. cytology: Associated with In(2LR)bwvl =
Tarsal shortening enhanced by homozygous s s a and In(2LR)21C8-Dl;60Dl-2 + ln(2LR)40F;59D4-El.
ssaB (Villee, 1945, Genetics 30: 26-27). RK1. other information: ds33k has not been separated
cytology: Analysis by E. B. Lewis (1945, Genetics from the recessive lethal factor of In(2LR)bwVl,
30: 137-66) indicates that ds is located in 21D1-2 and so has not been observed in homozygous
or possibly slightly to the left in the last band of condition.
21C.
discoverer: Waddington, 38k.
references: Curry, 1939, DIS 12: 45.
phenotype: ds38k/ds has close crossveins; fly
short and thick bodied. Homozygote probably like
dsd. RK2.
discoverer: T. Hinton and Bliven, 41b.
references: Hinton, 1942, DIS 16: 48.
phenotype: Wings shorter and extremely blunt.
Less extreme and more varied in male. Viability
and fertility good. RK2.

discoverer: Meyer and Byers, 51a.
references: Meyer and Edmonds on, 1951, DIS 25: 72.
phenotype: Like dsd. RK3.
*dsd: dachsous-dachsoid
discoverer: Sturtevant, 17b9.
ds: dachsous references: Bridges and Morgan, 1919, Carnegie
Edith M. Wallace, unpublished. tost. Wash. Publ. No. 278: 294 (fig.).
phenotype: Wings broader but half normal length,
almost round and widely spread; crossveins
origin: Spontaneous. broken, very close together. Hairs erect on costal
discoverer: Bridges, 25d2. vein near base. Body small and foreshortened.
references: Stern and Bridges, 1926, Genetics 11: Legs stumpy. Viability about 40 percent wild
513. type. RK3.
phenotype: Crossveins closer together, but only *Jsd53h
slight shortening of legs, wings, and abdomen. origin: Ultraviolet induced.
Fully viable and fertile. RK1. discoverer: Meyer, 53h.
s3 references: 1953, DIS 27: 58.
origin: Spontaneous, phenotype: Wings rounded; crossveins close. Legs
discoverer: Bridges, 25k5. have thickened coxae. Viability low; females
references: Stern and Bridges, 1926, Genetics 11: sterile. RK3.
513. ds*: dachsous-Wide
phenotype: Crossveins extremely close, but wings, origin: Spontaneous.
legs, and abdomen shortened only slightly. Via- discoverer Bridges, 29d24.
bility low (40 percent wild type), and females synonym: da2*d.
completely sterile. Males fertile. Emergence references: 1935, DIS 3: 10.
delayed. RK3. phenotype: Heterozygote has uniformly widely
spaced posterior scute liar bristles; wings a bit
see ds* short and blunt; crossveins close together. Via-
ds33k bility and separability excellent. Homozygote
origin: X ray induced at same time as resembles ds rf and dtsd3 with viability 40 percent
ln(2LR}bwVl. wild type. RK1 as heterozygous dominant; RK3
discoverer: Bridges, 33k28. as homozygote.
references: 1935, DIS 4: 7. dsh: dishevelled
j*h#n©ty|H>: dt$33k/+ has phenotype like dsw/+; location: 1-33.5.
s cute liar bristles wide apart, wings shorter and origin: Induced by methyl methanesulfonate
wider than normal, and crossveins close together. (CB. 1540).
d@33k/dsW n^g extreme phenotype like d«d. discoverer: Fahmy, 1956.
RK3A. references: 1959, DIS 33: 85.
MUTATIONS 79
phenotype: Thoracic hairs deranged. One or more has sex combs; however, on basitarsus of forelegs
hairs abnormally curved. Wings usually divergent of each, bristles of last transverse row are
and blistered. Eyes ellipsoid, with some deranged enlarged and rotated toward area comb would
ommatidia. Males viable and fertile; females occupy if present. Internally, the X/X intersex
sterile. RK2. usually has male and female reproductive parts
other information: One allele induced by CB. 3026. with varying degrees of completeness. Well-
developed ovaries present in some. Frequently a
single gonad is attached to both male and female
systems. The X/Y intersex usually has a predom-
inantly male internal reproductive system, but
occasionally both a male and a female system are
present. In all cases, gonads are poorly de-
veloped. RK2.
discoverer: Puro, 601.
dsr: disrupted synonym: Originally called ix-3: intersex on
From Bridges and Brehme, 1944, Carnegie Inst. Wash. chromosome 3.
Publ. No. 552: 7 1 . references: 1964, DIS 39: 64.
phenotype: Like dsx, X/X and X/Y flies have
dsr: disrupted external characteristics of both sexes and are
location: 2-90. similar except for a slight difference in size. No
origin: Spontaneous. sex combs. Last abdominal segments form rather
discoverer: Curry, 38a28. well-developed but rotated male-like terminalia
phenotype: Wings have plexus of extra and doubled with female-like structures of the seventh and
veins at anterior and posterior cross veins and at eighth tergites. Rudimentary gonads usually a
L3 and L4. L3 and L4 spread wide apart. Wing mass of undifferentiated tissue. RK2.
slightly wider and warped. At 25°, overlaps wild cytology: Salivary chromosomes normal.
type; at 19°C, no overlap but viability reduced to other information: Not allelic to tra and no inter-
60 percent wild type. RK3. action with ix in doubly heterozygous compounds.
*dss: disturbed segmentation *Dt: Detached
location: 1-27.3. location: 2-10.
origin: Spontaneous. discoverer: Bridges, 17ell.
discoverer: Fahmy, 1954. references: Morgan, Bridges, and Sturtevant, 1925,
references: 1959, DIS 33: 85. Bibliog. Genet. 2: 224.
phenotype: Extremely abnormal abdomen with phenotype: Vein L2 fails to reach margin in 60
segmentation grossly deformed, very few hairs, percent of flies. Homozygote not known. RK3.
and disturbed pigmentation. Occasionally some other information: Bridges considered this a
bristles shortened. Eyes reduced in size and possible effect of 5 or requiring S as an enhancer
sometimes abnormal in shape. Males fertile; as it was found in a S stock and apparently was
viability about 10 percent wild type. Females never separated from S.
sterile. RK3. *dta: delta wing
dsx: double sex location: 1- (rearrangement).
location: 3-48.1 (to the right of pP). origin: Induced by triethylenemelamine (CB. 1246).
origin: Recovered among progeny of flies fed discoverer: Fahmy, 1953.
tritiated thymidine. references: 1958, DIS 32: 69.
discoverer: Hildreth and Lucchesi, 62c. phenotype: Wings widely outspread, frequently
synonym: Originally called ix62c: intersex-62c, drooping in homozygous female. Viability good
but name and symbol preoccupied. but female sterile. RK2A.
references: 1963, Proc. Intern. Congr. Genet., 11th, cytology: Associated with ln(l)dta =
Vol. 1: 171. In(l)6B2-3;15E7-F2.
Hildreth, 1965, Genetics 51: 659-78 (fig.)- dtv: see thv**
phenotype: Chromosomal females (X/X) and males *du; dunk el
(X/Y) transformed into intersexes. The two types location: 3-47.
are similar. Pigmentation of tergites similar to origin: Spontaneous,
wild-type males. Like females, they have seventh discoverer: Hadorn, 49el5.
tergite with seventh spiracle at its base. Ventral references: Hadorn and Fritz, 1950, Arch. Julius
part of eighth abdominal segment has protuberance Klaus-Stift. Vererbungsforach. Sozialanthropol.
similar to female gonopod; ninth segment bears Rassenhyg. 25: 504-8.
claspers like males. Anal plates situated verti- phenotype: Body color dark, sootylike. Wings
cally as in males. Neither X/X nor X/Y ixttersex blistered. Viability almost normal at 25°, greatly
reduced at 18°C. Males fertile; females sterile. references: 1936, Biol. Zh. (Moscow) 5: 537—40
Ovaries and eggs normal size and morphology. (fig.).
Insemination of females normal (motile sperm in 1937, DIS 8: 81.
spermathecae and receptaculum). Either eggs from phenotype: Has shorter, darker wings; post-
du females not fertilized or zygotes die before scutellars bowed in; body size small; sterility
blastoderm formation, du ovaries behave auton- high; semilethal. In combination with yellow-
omously as implants in normal hosts, and wild- bodied y alleles, gives strongly curled wings with
type ovaries are fertile in du hosts. RK2 at 25°C. slight outward twist. With f, gives crumpled
other information: Not an allele of by or cu. wings. With sc, almost lethal. RK3.
cytology: Salivary chromosome studies by Demerec
dumpoidy: see dpy and Sutton show locus to lie at right of 8D8-9
dumpy: see dp (Demerec, Kaufmann, Fano, Sutton, and Sansome,
dunkel: see du 1942, Carnegie Inst. Wash. Year Book 41: 191).
dusky: see dy
dusky body: see dyb origin: Spontaneous.
discoverer: Curry, 37kl7.
phenotype: Practically wild type. With y2, wings
tightly curled; with y, wings spirally curled. RK2
with y.
*dvrs: divers-subliminal
discoverer: Muller.
synonym: dvri*).
phenotype: Wild type either alone, heterozygous to
dvr-2, or in combination with y. y dvra/y dvt%, on
the other hand, has wings distinctly curly or
wavy, usually as in typical Cy, but other effects
noted in dvr flies not evident. RK3.
*dvw: divergent wings
location: 1-13.3.
dv: divergent origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
From Bridges and Morgan, 1923, Carnegie Inst. Wash. phenylalanine (CB. 3026).
Publ. No. 327: 182. discoverer: Fahmy, 1953.
*dv: divergent phenotype: Sex-limited character. Males late
location: 3-20.0, hatching; wings divergent, occasionally upheld,
origin: Spontaneous, with inner margins frequently cut away to various
discoverer: Bridges, 17fl3. degrees. Bristles short and stiff. Homozygous
references: Bridges and Morgan, 1923, Carnegie females normal. RK1 in males.
Inst. Wash. Publ. No. 327: 182 (fig.). *dw: dwarf
Morgan, Bridges, and Sturtevant, 1925, Bibliog. location: 3-50.
Genet. 2: 58 (fig.). origin: Spontaneous.
Mohr, 1937, DIS 8: 12. discoverer: Bridges, 13kl2.
phenotype: Wings spread, smaller and have slight references: Bridges and Morgan, 1923, Carnegie
venation disturbances. Both sexes rather in- Inst. Wash. Publ. No. 327: 101.
fertile. dv/Dt(3L)Vn progeny of homozygous dv Morgan, Bridges, and Sturtevant, 1925, Bibliog.
mothers practically lethal although the same geno- Genet. 2: 58 (fig.).
type from other crosses survives (Mohr and phenotype: Body weight 76 percent that of hetero-
Mossige, 1943, Avhandl. Norske Videnskaps-Akad. zygous sibs. Females usually sterile (3 of 63
Oslo, I. Mat.-Natur. Kl. No. 7: 1-51). RK2. gave a few offspring). RK3.
cytology: Salivary chromosome locus placed be- dw-24F: dwarf in salivary chromosome section
tween 64C12 and 65E1, on basis of its inclusion in 24F
Df(3L)Vn = Df(3L)64C12~Dl;65D2-El (Mohr, 1938, location: 2-13.
Avhandl. Norske Videnskaps-Akad. Oslo, I. origin: Spontaneous.
Mat.-Natur. Kl. No. 4: 1-7). discoverer: Curry, 39k.
dvr: divers phenotype: Body small; abdomen narrow and mis-
location: 1-28.1 (located using a W ) . shapen. Body surface dull if not properly dried.
origin: Recovered among progeny of iodine-treated Eyes dull in color and smallish. Wings close
male. textured, small, tend to droop; crossveins close.
discoverer: Sacharov, 1932. Bristles slender. Low viability and fertility. RK3.
MUTATIONS
cytology: Located between 24E2 and 25A2 based on origin: Induced by 2-chlorethyl methanesulfonate
its inclusion in Df(2L)M-zB = Df(2L)24E2- (CB. 1506).
Fl;25Al-2 (Morgan, Bridges, Schultz, and Curry, discoverer: Fahmy, 1956.
1939, Carnegie Inst. Wash. Year Book 38: 273-77). references: 1959, DIS 33: 85.
*dw-b: dwarf-b phenotype: Extremely inviable dwarf; wings
location: 3-12. frequently fail to expand completely. Males fertile
origin: Spontaneous. if they survive to breed. RK3.
discoverer: Bridges, 20b5. dwx: dwarfex
references: Bridges and Morgan, 1923, Carnegie location: 1-33.2.
Inst. Wash. Publ. No. 327: 182, 228, 231 (fig.). discoverer: Bridges, 33c31.
phenotype: Flies about 70 percent as heavy as phenotype: Body small. Wing texture coarse;
wild type. RK3. marginal hairs slightly disarranged. Classifica-
tion sometimes difficult. RK3.
*dw-sc: dwarf with scute *dwxm n: dwarfex-manikin
location: 1-0.7. origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
origin: Spontaneous (arose with sc and separated). phenylalanine (CB. 3026).
discoverer: Bridges, 16a22. discoverer: Fahmy, 1954.
phenotype: Small body. Viability erratic. RK3. synonym: mn.
dwarf unexpanded: see dwu references: 1959, DIS 33: 88.
dwarfex: see dwx phenotype: Fly small with narrow abdomen. Reduc-
dwarfish: see dwh tion in size may be bilaterally asymmetrical and
dwarfoid: see dwf affect abdomen and thorax independently. Male
dwarp: see dwp viability reduced; flies rarely survive more than 48
*dwf: dwarfoid hr. Sterile, probably owing to reduced vigor. RK3.
location: 1-13.3. other information: One X-ray-induced allele.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- dx: deltex
discoverer: Fahmy, 1955. origin: Spontaneous.
references: 1959, DIS 33: 85. discoverer: Bridges, 22h26.
phenotype: Flies small. Males fertile; viability references: Morgan, Bridges, and Schultz, 1931,
about 50 percent wild type. Homozygous females Carnegie Inst. Wash. Year Book 30: 410.
show extreme expression; fertility and viability phenotype: Veins show thickenings and terminal
low. RK2. deltas, resembling Dl in third chromosome, but
dwg: deformed wings fully viable, fertile, and easily classified. Nearly
location: 1-1.6. suppressed by su(dx), Su(dx), and Su(dx)2. RK2.
origin: Induced by DL-p-NN-di(2-chloroethyl)amino- cytology: Demerec and Sutton show locus to be
phenylalanine (CB. 3007). between 6A3-4 and 6F10-11 (Demerec, Kaufmann,
discoverer: Fahmy, 1954. Fano, Sutton, and Sansome, 1942, Carnegie Inst.
references: 1959, DIS 33: 85. Wash. Year Book 41: 191).
phenotype: Wings broad, round tipped, and have
occasional marginal incisions; sometimes grossly dxst: deltex-sterile
deformed in shape and venation. Extremely fine origin: Spontaneous change of dx to dxst.
bristles. Eyes small and occasionally rough. discoverer: Bridges, 31a3.
Males late in eclosion; viable but sterile. RK2. phenotype: Veins heavy, confluent, and dilated at
dwh: dwarfish junctions; strong deltas at tips. Wings spread
location: 3- (not located), wide; margins and tips snipped and nicked. Ocelli
origin: Spontaneous. sometimes fused, with disturbance of hairs and
discoverer: Bridges, 30dl6. bristles in the region. Acrostichals irregular.
phenotype: Small body. Wings disproportionately Male sterile. Less abnormal phenotype and fertile
broad; eyes irregularly knobby and somewhat dull with Su(dx). RK2.
in color; legs weak and slightly crippled. RK3. dy: dusky
*dwp: dwarp location: 1-36.2 (to the right of m).
location: 1-0. origin: Spontaneous.
origin: Spontaneous, discoverer: Bridges, 1611.
discoverer: Bridges, 23b20. references: Morgan, Bridges, and Sturtevant, 1925,
references: Morgan, Bridges, and Sturtevant, 1925, Bibliog. Genet. 2: 35 (fig.), 224.
Bibliog. Genet. 2: 225. Slatis and Willermet, 1954, Genetics 39: 45-58.
phenotype: Dwarf with warped wings. Body color Dora and Burdick, 1962, Genetics 47: 503-18.
pale. Nearly always lethal, but survivors fertile. phenotype: Wings smaller than normal but of nearly
RK3. wild-type shape, dusky in color. Cell expansion
*dwu; dwarf unexpanded inhibited in prepupal as well as pupal period
location: 1-58.3. (Waddington, 1940, J. Genet. 41: 75-139). RK1.
cytology: Demerec and Sutton showed that locus phenotype: Similar to other ay's. Fully fertile in
lies just to right of 10E1-2 (Demerec, Kaufmann, both sexes. Shows high complementation with
Fano, Sutton, and Sansome, 1942, Carnegie Inst. m60 and low with dy61a. RK1.
Wash. Year Book 41: 191). dy61a
other information: No reverse mutations among origin: Gamma ray induced.
2 X 10 s progeny of progeny of homozygous females discoverer: Ives, 61a24.
(Pullar). A member of the m-dy pseudoallelic references: 1961, DIS 35: 46.
complex. Dorn and Burdick, 1962, Genetics 47: 503-18.
phenotype: Like dy. Good fertility and fecundity in
both sexes. RK1.
dy62b
discoverer: Burdick, 62b.
phenotype: Similar to other dy's. RK1.
*dy286-9
discoverer: Demerec, 35b.
phenotype: Lethal in male and cell lethal. RK2.
other information: rn + and fw+.
*dyala: dusky-alae
origin: Recovered among progeny of heat-treated
flies.
discoverer: Gottschewski, 34c.
synonym: a/a; ala parvae.
phenotype: Wings about 90 percent as large as dy.
dy: dusky RK1.
Edith M. Wallace, unpublished. *dyb: dusky body
location: 1-44.6.
*dy2 origin: Induced by ethyl methanesulfonate
origin: Spontaneous. (CB. 1528).
discoverer: Bridges, 20a3. discoverer: Fahmy, 1958.
references: Morgan, Bridges, and Sturtevant, 1925, references: 1959, DIS 33: 85.
Bibliog. Genet. 2: 224. phenotype: Dusky body color and browner eyes.
phenotype: Wings not quite so small as dy. RK1. Eye and wing shapes slightly altered. Males
*dy3 viable and fertile; females sterile. RK2.
references: Morgan, Bridges, and Sturtevant, 1925, e; ebony
Bibliog. Genet. 2: 224. location: 3-70.7.
phenotype: Wings slightly larger than dy. Clean origin: Spontaneous,
separation difficult in females. RK2. discoverer: E. M. Wallace, 12bl5.
*dy31d references: Bridges and Morgan, 1923, Carnegie
origin: X ray induced. Inst. Wash. Publ. No. 327: 50 (fig.).
discoverer: Oliver, 1931. phenotype: Body color shining black. Puparia much
synonym: m-like: miniature-like. lighter than wild type. Classifiable throughout
phenotype: Like dy. RK1. larval period by darkened color of spiracle sheaths
dySSk (Brehme, 1941, Proc. Natl. Acad. Sci. U.S. 27:
origin: Spontaneous. 254—61). Viability lowered to about 80 percent
discoverer: Krawinkel, 58k. wild type. Heterozygote has slightly darker body
references: Burdick, 1961, DIS 35: 45. color than normal. For interaction with other body
Dorn and Burdick, 1962, Genetics 47: 503-18. color mutants, see Waddington, 1941, Proc. Zool.
phenotype: Wings shorter than ay, resembling the Soc. London, Ser. A 111: 173-180. Virtually un-
longer-winged m's. Fertile in both sexes. Shows able to incorporate C 1 4 from labeled beta-alanine
more complementation with tn'» than ay. RK1. into puparium or hardening adult integument,
other information: Recotnbines with all m's except whereas normal flies incorporate heavily; e/+ in-
mD but not with ay. corporates intermediate amounts Qacobs and
dy60k Brubaker, 1963, Science 139: 1282-83; Jacobs,
origin: Spontaneous, 1966, Genetics 53: 777-84). RK1.
discoverer Burdick, 60k. cytology: Placed in salivary chromosome region be-
references: 1961, DCS 35: 45. tween 93B7 and 93F9, on the basis of its inclusion
MUTATIONS 83
in both Dt(3R)el 00.1 72 = Df(3R)93B7-10;93F10- phenotype: Like es when heterozygous with e».
94A1 and Df(3R)el 00.256 = Df(3R)93A5-Bl;93F5-9 Homozygous lethal. RK2A.
(Ward and Alexander, 1957, Genetics 42: 42—54). cytology: Associated with Df(3R)el00.256 -
Df(3R)93A5-Bl;93F5-9.
origin: Spontaneous. *e100.265
discoverer: Sturtevant, 17i27. origin: X ray induced.
references: Bridges and Morgan, 1923, Carnegie discoverer: Alexander.
Inst. Wash. Publ. No. 327: 184. references: Ward and Alexander, 1957, Genetics 42:
phenotype: Body color very dark like e, but 42-54.
abdomen lighter. Most viable and generally best phenotype: Like e. RK1A.
of the dark alleles, such as e and ell. c l a s s i - cytology: Associated with In(3R)e*00.265 =
fiable from hatching throughout larval period by In(3R)93B5-6;95E.
darker spiracle sheaths (Brehme, 1941, Proc. *e100.307
Natl. Acad. Sci. U.S. 27: 254-61). RK1. origin: X ray induced.
*e4.39 discoverer: Alexander.
origin: X ray induced. references: Ward and Alexander, 1957, Genetics 42:
discoverer: Alexander. 42-54.
references: 1960, Genetics 45: 1019-22. phenotype: Like es when heterozygous with e«.
phenotype: Like es when heterozygous with e«. Homozygous lethal. RK2.
Homozygous lethal. RK2A. cytology: Induced simultaneously with
cytology: Associated with Df(3R)e4-39 = In(3L)100.307 = In(3L)62E2-4;64C2-4.
Di(3R)93B;93F. other information: Not shown whether lethality
associated with e or the inversion.
origin: Spontaneous. *e300.96
discoverer: Stern, 25a. origin: X ray induced.
references: 1926, Z. Induktive Abstammungs- discoverer: Alexander.
Vererbungslehre 41: 198-215. references: Ward and Alexander, 1957, Genetics 42:
1934, DIS 1: 35. 42-54.
phenotype: Dark allele similar to e and e4. phenotype: Like e but male sterile. RK2(A).
Tyrosinase formed in adults (Horowitz). Classi- cytology: Induced simultaneously with
fiable throughout larval period by dark spiracle In(3R)300.96 = In(3R)89F2-90Al;99B2-4 but prob-
sheaths (Brehme, 1941, Proc. Natl. Acad. Sci. ably separable from it.
U.S. 27: 254—61). Phenylthiocarbamide inhibits e*: ebony-sooty
development of ell homozygotes more than wild origin: Spontaneous.
type; reverse is true for inhibition by silver discoverer: Sturtevant, 13i20.
chloride; heterozygotes intermediate in both references: Bridges and Morgan, 1923, Carnegie
cases. Mixtures of the two inhibitors affect heter- Inst. Wash. Publ. No. 327: 99 (fig.).
ozygotes to a greater extent, thus exhibiting phenotype: Body color darker than wild type but
"negative heterosis" (Kroman and Parsons, 1960, lighter than e. Viability excellent. Most easily
Nature 186: 411—12), Under ordinary conditions, classified as ea/e4. Classifiable from hatching
viability and fecundity of heterozygote superior to throughout larval period by dark color of spiracle
either homozygote (Moree). RK1.
sheaths, although darkening is less than in e, e4,
eiitti: see tu-bwe
or ell (Brehme, 1941, Proc. Natl. Acad. Sci. U.S.
e60h
27: 254—61). Ability to incorporate beta-alanine-
origin: Spontaneous, derived C intermediate between that of e and +
discoverer: Ives, 60h. (Jacobs and Brubaker, 1963, Science 139: 1282-83;
references: 1965, DIS 40: 35. Jacobs, 1966, Genetics 53: 777-84). RK1.
phenotype: Medium e. RK1. *e**: ebony-striped
* e 700.I72 origin: Spontaneous.
origin: X ray induced. discoverer: Villee, 39k.
discoverer: Alexander. references: 1941, DIS 14: 40.
references: Ward and Alexander, 1957, Genetics 42: 1942, Univ. Calif. (Berkeley) Publ. Zool. 49: 137.
42-54. phenotype: Body color similar to es but with
phenotype: Like e* when heterozygous with e*. definite longitudinal stripes on mesonotum.
a at
Homozygous lethal. RK2A. eat/ea resembles e ; e /e resembles e. Viability
cytology: Associated with Dl(3R)el 00.172 m excellent. RK2.
Df(3R)93B7-10;93F10-94Al. ; ebony-Uganda
*9100.2S6 origin: Spontaneous.
origin: X ray induced. discoverer: Zurcher, 1956.
discoverer Alexander. references: 1958, Arch. Julius Klaus-Stift.
references: Ward and Alexander, 1957, Genetics 42: Vererbungsforsch. Sozialanthropol.
42-54. Rassenhyg. 33: 75.
1960, DIS 34: 112. e(dpv): enhancer of dumpy-vortex

1963, Genetics 34: 1-33 (fig.). location: 3-40.4.
phenotype: A light ebony allele. Females have origin: Spontaneous.
dark, extremely wide marginal bands on abdominal discoverer: Bridges, 16h7.
tergites; entire abdominal tergite region darkly synonym: vo-3: vortex in chromosome 3.
shaded. Males only slightly darker than wild type. references: 1919, Bridges and Mohr, Genetics 4:
Viability lower than wild type. RK2. 283-306 (fig.).
E(B): Enhancer of Bar 1923, Bridges and Morgan, Carnegie Inst. Wash.
location: 1-57.3. Publ. No. 327: 168.
origin: Spontaneous. 1925, Morgan, Bridges, and Sturtevant, 1925,
discoverer: Bonnier and Nordenskibld. Bibliog. Genet. 2: 41-43 (fig.).
synonym: i; I: Intensifier of Bar; Eb: Exaggeration phenotype: Normal. In combination with dpv
of Bar. produces one or two pairs of pits or volcanolike
references: 1942, DIS 16: 47. protrusions on thorax; hairs and bristles arranged
Bonnier, Nordenskiold, and Bagman, 1943, in whorls. RK3.
Hereditas 29: 113-33 (fig.). *E(f): Enhancer of forked
Rasmuson, 1948, Proc. Intern. Congr. Genet., 8th. location: 2-86.5.
pp. 645—46. origin: X ray induced,
phenotype: E(B) heterozygous with any B allele, discoverer: Belgovsky, 37c4.
including 23+, produces flies similar in phenotype synonym: I-t: Intensifier of forked.
to homozygotes for that allele. B +/+ E(B) eyes references: 1937, DIS 8: 7.
have 80—90 facets but B E(B)/++ eyes have only 1938, Izv. Akad. Nauk SSSR, Ser. Biol. 1017-36.
40. Homozygous lethal. RK2(A). 1940, DIS 13: 52.
cytology: Salivary chromosomes appear normal, but 1944, Zh. Obshch. Biol. 5: 325-56.
there is occasional indication of deficiency for phenotype: Homozygote has short, twisted bristles
faint bands 16A5 and 6. intermediate between f and Bl; postcutellars often
other information: Reduces B-fu crossing over about pale; viability and fertility reduced. Heterozygote
40 percent. is wild type. f/+; E(f)/+ slightly more extreme than
e(bx): enhancer of bifhorax f. f/f; E(f)/+ has an extreme forked phenotype and
location: 1-1.0. hairs are forked, f/f; E(f)/E(f) rarely survives.
origin: Gamma ray induced. RK3.
discoverer: E. B. Lewis, 53b. cytology: Salivary chromosomes normal.
synonym: en-bx. *e(g): enhancer of garnet
phenotype: Recessive enhancer of most bithorax discoverer: Payne and Denny, 1921.
alleles. By itself, the only abnormal phenotype is synonym: m(g): modifier of garnet.
a slight variegated eye color. Most readily references: 1921, Am. Naturalist 55: 377-81.
classified in combination with bx34e +/+ Ubx. phenotype: Apparently wild type, but in combination
Enhances development of dorsal metanotum with g produces a more orange eye than g alone.
of latter from a few hairs to a broad band of hairy RK3.
tissue somewhat like that seen in homozygous *E(H): Enhancer of Hairless
fox3*8. Enhances bx3 +/+ pbx from wild type to a location: 2-50.5.
slight postbithorax phenotype (slight wing-like discoverer: Nash.
modification of posterior portion of halteres). references: 1963, Proc. Intern. Congr. Genet., 11th.
RK2A. Vol. 1: 174-75.
cytology. Associated with In(l}e(bx) = In(l)3A;4F. Nash, 1965, Genet. Res. 6: 175-89.
Locus within Dp(l;l)w = Dp(l;l)3A;3C, probably phenotype: E(H); H flies nearly devoid of chaetae.
in section 3A. Majority of bristle sites are vestiges. RK2.
e(bx)2 *E(M3g): Enhancer of Minute(3) g
origin: X ray induced, location: 3- (near spindle attachment).
discoverer: E. B. Lewis, 55h, origin: Spontaneous.
synonym: m2~bx. discoverer: Bridges.
references: 1959, DIS 33: 96. phenotype: Specific intensifier of shortness of
phenotype: Similar to e(hx). RK2. bristles of M(3)g. RK3.
cytology: Salivary chromosomes normal. *e(N*): enhancer of Notch-8
E(bx): Enhancer of biihorax location: 3- (not located).
location: 3- (to the left of R). origin: Spontaneous.
origin: X ray induced. discoverer: Mohr, 181.
discoverer. E. B. Lewis. references: 1923, Z. Induktive Abstammungs-
synonym: En-bx. Vererbungslehre 32: 108-232 (fig.).
ph*notyp«: Enhances expression of bx34m, bx3, and phenotype: Produces slight nicking of wings.
Ubx/'*. Lethal homozygous. RK2. Enhances Ns. RK3.
MUTATIONS 85
*e(S): enhancer of Star Variegated position effects do not respond uni-

location: 3- (between 0 and 10; perhaps an allele of formly to E(var)7. RK2(A).
ru or R). cytology: May be small abnormality in 25A (Schultz).
origin: Spontaneous. E(w°): Enhancer of white-apricot
discoverer: Bridges, 16kl8. location: 2- (not located).
synonym: S-i: intensifier of Star. origin: Spontaneous.
references: Bridges and Morgan, 1923, Carnegie discoverer: Scandlyn.
Inst. Wash. Publ. No. 327: 175 (fig.), phenotype: Heterozygote dilutes wB to pale yellow.
phenotype: By itself, homozygous e(S) has normal Homozygote with wa is white. Not tested with
eyes. S/+; e(S)/e(S) has eyes smaller and rougher other white alleles. No effect on eye color in
than S/+, although overlapping somewhat; abdomen presence of w+. Homozygous sterile. RK3.
bulbous, body color darkish. RK3. *efw«); enhancer of white-eosin
E(S): Enhancer of Star location: 1-32.
location: 2-6 [claimed to lie between left break of origin: Spontaneous.
In(2L)Cy and locus of Cy\. discoverer: Green, 55b21.
discoverer: Bridges, 30a27. synonym: en-we.
phenotype: E(S)/+ normal, E(S)/B(S) gives slight references: 1957, DIS 31: 81.
roughening of eye. B(S)/+ strongly reduces size 1959, Heredity 13: 303-15.
and increases roughness of S/+ and S2/+ eyes; phenotype: Enhances intermediate alleles at the
imparts dominance to ast/+, ast%/+, ast3/+, and fourth recombinational site of the w locus, e.g.,
a s ^ / + ( L e w i s , 1945, Genetics 30: 137-66). we, we2, WBE, wh, and wXi6 to produce nearly
S +/+ E(S) occasionally emerges as a late-eclosing white eye color. No enhancement of tested inter-
giant. RK3A. mediate alleles at other w subloci, e.g., wa, w"2)
cytology: Arose in In(2L)Cy = wa3, w*"t, wbi, wch2, wco, wcoi, or wsat> Also
In(2L)22Dl-2;33F5-34Al. suppresses f. e(we) flies occasionally have px-
E(sd): Enhancer of scalloped like venation, or shortened wings, or both. Homo-
location: Autosomal. zygous females sterile. RK2.
origin: Spontaneous. eagle: see eg
discoverer: R. M. Valencia, 1963. eb: ebonized
phenotype: Almost completely removes wings of origin: Induced by ethyl methanesulfonate
sdBP; not tested with other alleles of sd. No inter- (CB. 1528).
action with Bx or BxT. RK2. discoverer: Fahmy, 1956.
E(spl): Enhancer of split references: 1959, DIS 33: 86.
location: 3- (near ro). phenotype: Heavily pigmented fly, with trident
origin: Spontaneous. pattern and scutellum very dark. Wings slightly
discoverer: Green. shorter; membrane often slightly concave; wing
synonym: En-spl. tips occasionally truncate. Males viable and
references: Welshons, 1956, DIS 30: 157-58. fertile; females sterile. RK2.
Von Halle, 1965, DIS 40: 60. Eb: seeE(B)
phenotype: Both E(spl)/+ and E(spl)/E(spl) cause ebonized: see eb
spl/+ to resemble spl/spl and cause spl/spl and ebony: see e
spt/Y to have an extreme phenotype. ec: echinus
spl/+/Dp(l;2)51b; E(spl)/+ less extreme than location: 1-5.5.
spl/+; E(spl)/+. E(spl) homozygote viable and origin: Spontaneous,
fertile. RK2. discoverer: Bridges, 1516.
e(tu-K): enhancer of tumor K phenotype: Eyes large and bulging. Eye surface
location: 3- (not located). rough; facets large. Wings rather short and broad.
origin: Spontaneous. Body thickset. Tends to remove dorsocentrals
discoverer Bur net and Sang. (posterior more than anterior) and posterior
references: 1964, Genetics 49: 223—35. notopleurals; may also add dorsocentrals anterior
phenotype: Homozygote produces a significant in- to anterior dorsocentrals whether or not posterior
crease in the penetrance of tu-K in both untreated bristles removed (Sturtevant). ec is visible in
flies and those treated in ways known to increase +/ec/ec triploids (Gersh). RK1.
tumor incidence in tu-K. RK3. cytology: Locus placed at 3F1-2 by Demerec and
E(var)7: Enhancer of variegation Sutton (Demerec, Kaufraann, Fano, Sutton, and
location: 2- (not located). Sansome, 1942, Carnegie Inst. Wash. Year Book
origin: X ray induced. 41: 191).
discoverer: Schultz. echinoid: see ed
phenotype: E(ver)7/+ has no phenotype of its own echinus: see ec
but enhances variegation, e.g., w**>* is made much ed: echinoid
lighter and variegation for tmt appears in males. locotion: 2-11.0.
origin: Spontaneous. el: elbow

discoverer: Bridges, 31al6. location: 2-50.0.
phenotype: Eyes large and rough. Easily classified, origin: Spontaneous.
although not so extreme as ec. RK1. discoverer: E. M. Wallace, 35dl.
cytology: Placed between 24D2 and 24F1 on the phenotype: Wings extended and bent backward (as
basis of its inclusion in Df(2L)M'Zc = Df(2L)24D2- in bt and bat), often warped, shortened; sometimes
5;25A2-3 but not in Df(2L)M-zB = Df(2L)24E2- blistered or nicked. Alula always reduced and
Fl;25Al-2 (Morgan, Bridges, Schultz, and Curry, fused with main wing blade. Venation reduced by
1939, Carnegie Inst. Wash. Year Book 38: 273-77). terminal shortening of L5 and of crossveins.
*e/: elfin Balancers reduced, especially in third segment.
location: 1- (rearrangement). Eye size decreased, varying with other manifesta-
origin: Induced by triethylenemelamine (CB. 1246). tions. RK2.
discoverer: Fahmy, 1952. cytology: Placed in region between 34E5 and 35D1
references: 1959, DIS 33: 86. on the basis of its inclusion in Df(2L)64j =
phenotype: Small fly with slightly excess melaniza- Df(2L)34E5-Fl;35C3-Dl (E. H. Grell).
tion. Wings proportionally smaller, slightly altered elfin: see ef
in shape, and warped. Abdominal tergites often Ellipse: see Elp
broken and abnormally pigmented. Males viable elliptical rough: see e/r
but sterile. RK3A. Elp: Ellipse
cytology: Associated with T(l;2)ef = location: 2-99 (to the left of px).
T(1;2)14C8-D1;2R. origin: Spontaneous.
eg: eagle discoverer: E. H. Grell, 57b.
discoverer: Morgan, 1930. phenotype: Eyes of heterozygotes rough and more
phenotype: Wings extended. Hairs on thorax some- oval than wild type. Homozygotes have small
what disarranged. Dark pattern on thorax. RK2. eyes with fused facets; resemble homozygous ast.
eg' Homozygotes have low viability. RK1 as hetero-
discoverer: Bridges, 33J16. zygote.
synonym: dh; dihedral. *e/r; elliptical rough
phenotype: Wings extended and uplifted. Females location: 1-25.1.
often sterile. RK2. origin: X ray induced.
eg 57c discoverer: Fahmy, 1956.
discoverer: Nicoletti, 57c. phenotype: Eyes slightly elliptical and rough.
references: 1957, DIS 31: 84. Wings slightly broader. Both sexes viable and
phenotype: Wings held out at 45-90° angle from fertile. RK2.
midline. Excellent viability and fertility. RK1. other information: Two other alleles: one induced
by X rays, one by CB. 3025.
en: engrailed
location: 2-62.0.
discoverer: Evang, 26k7.
references: Eker, 1929, Hereditas 12: 217—22
(fig.).
Bridges, Skoog, and Li, 1936, Genetics 21: 788—
95.
Brasted, 1941, Genetics 26: 347-73 (fig.).
phenotype: Longitudinal cleft extends from rear
border of scutellum forward; may be reduced to
median nick or posterior flattening of scutellum.
Bristles often javelin- or hooked-like. Wings
larger, broader, and thin textured, with spatulate
end; venation always disturbed (like that of cf),
with gap in L4 and L5 and branching plexus of
extra veins. In males, extra sex comb often
present, smaller than normal and in mirror-image
position on outer side of tarsus. Action of en on
differentiation of secondary sex comb on male
foreleg is autonomous LTokunaga, 1961, Genetics
el: elbow 46: 157—76 (fig.)J. Male genitalia may be malformed
From Bridges and Brehme, 1944, Carnegie inst. Wash. and rotated, resulting in sterility (Curry, 1941,
Pub!. No. 552: 75. DIS 14: 50). eg en/+ + characterized by slight
MUTATIONS 87
degree of L4 interruption and thinning in males at may be segmented and have hairs. Less extreme
low temperature. The triple heterozygote expression produces derangement of central or
eg en/+ +; ci/+ has about 50 percent expression anterior-central facets. Eruption may occur as
of L4 interruption (House, 1961, Genetics 46: encroachment of chitin with bristles and hairs into
871). For other interactions of en with ci alleles anterior edge of eye. RK2.
andfl, see House (1953, Genetics 38: 199-215, other information: Alleles of at least five different
309-27). RK1. strengths present in different wild stocks.
Present in many wild stocks in suppressed
condition.
*Er: Erect
location: 3-50.
discoverer: Neel, 41c9.
phenotype: Posterior scutellars at greater than
normal angle with body; vary from slight effect to
condition in which bristles stand at right angles
to scute Hum. In latter case, bristles usually
appear warped and twisted. Wings incompletely
expanded and crinkled to varying degrees. RK3.
erupt: see er
es: ether sensitive
location: 2- (not located).
discoverer: Tinderholt.
references: Kidd, 1963, DIS 37: 49.
phenotype: Hypersensitive to diethyl ether and
chloroform. Homozygotes killed by exposure to
doses of these agents harmless to normal flies.
en; engrailed Sensitivity probably affected by modifiers. A male
From Eker, 1929, Hereditas 12: 217-22. sterility factor seems to be associated but may be
separable. Viability of homozygote about 70 per-
en-; see e( ) cent that of es/SMl and remains low in strains
En-: see E( ) selected for less sensitivity. Not sensitive to
engrailed: see en carbon dioxide. RK3.
enhancer: see e( ) esc; extra sex combs
Enhancer: see E( ) location: 2-54.9 (based on location of escD).
eq: equational producer origin: Spontaneous.
location: 1- (to the right of car — probably hetero- discoverer: Slifer, 40e2.
chromatic). references: 1942, J. Exptl. Zool. 90: 31-40 (fig.).
origin: X ray induced, phenotype: Sex combs may be present on all six legs
discoverer: Schultz, 33a2. of male; at least one extra sex comb present in
references: Morgan, Bridges, and Schultz, 1934, majority of males. Expression affected by culture
Carnegie Inst. Wash. Year Book 33: 280. conditions. When expressivity high, extra trans-
phenotype: Produces 1—2 percent equational non- verse bristle rows appear between sixth and eighth
disjunction of X's in male, producing both X/X longitudinal rows of bristles, mainly on distal por-
and nullo-X, nullo-Y sperm. Original eq male tion of basitarsus and tibia of second and third
when crossed to attached-X female produced legs in both sexes; accompanied by shortening of
89/289 equational exceptional X/X daughters. affected leg segments. Sex comb development
Claimed to generate 66-deficient Y chromosomes. autonomous in mosaics produced by somatic cross-
RK3. ing over [Tokunaga and Stern, 1965, Develop. Biol.
cytology: Both salivary and mitotic chromosomes 11: 50—81 (fig.)]. For interactions with Pc and
appear normal. Sex see Hannah-Alavah, 1958, Genetics 43: 878—
er: erupt 905 (fig.). Males and females sterile. RK2.
location: 3-70.7 (60.7 to 80.7; not an allele of k). cytology: Arose in chromosome carrying In(2L)t »
origin: Spontaneous. In(2L)22D3-El;34A8'9, but mutant and inversion
discoverer: Glass, 1941. separable (Tokunaga).
references: 1943, DXS 17: 50. esc 2 : see escD
1944, Genetics 29: 436-46. *esc°: extra sex combs-Dominant
1957, Science 126: 683-89 (fig.), origin: Spontaneous,
phenotype: Exhibits eruption of underlying hypo- discoverer: Stromnaes, 53f.
dermis in center of one or both eye*. Eruption synonym: ©sc-2; Esc2.
references: Hannah and Stromnaes, 1955, DIS 29: system. Est-CF/Est-CF produces a rapidly
121-23. migrating esterase C and Est-CF/Est-Cs produces
phenotype: esc&/+ similar to esc/esc. Homozygous two bands on starch gel; no evidence for hybrid
lethal. RK2. enzyme in heterozygotes. Est-Cs Est-6S/Est-CF
Est-6F: Esterase 6-Fast Est-6F homogenates produce four electrophoretic
location: 3-36.8 (10.3 units to the right of h and 6.4 bands in place of the two found in homozygotes.
units to the left of th). RK3.
origin: Naturally occurring allele. Est-Cs: Esterase C-S/ow
discoverer: T. Wright, 61h. origin: Naturally occurring allele.
references: 1963, DIS 37: 53. discoverer: Beckman and Johnson.
1963, Genetics 48: 787-801 (fig.). references: 1964, Hereditas 51: 212-20 (fig.).
phenotype: Esterase 6 is one of ten positively phenotype: Produces esterase C with slow electro-
migrating esterases demonstrable histochemically phoretic mobility on starch gel. RK3.
with (X-naphthyl acetate and Fast Blue BB after ether sensitive: see es
starch gel electrophoresis of imaginal homoge-
nates. Readily identified in zymograms as the
most heavily staining esterase. Migrates about
5 cm when exposed to a voltage drop of 2.0—2.5 v
per cm for 17 hr at 4°C in a starch gel prepared
with 0.05 M tris buffer, pH 8.7. Flies homozygous
for Est-6F have a single esterase 6 band that
migrates faster than a similar, single esterase 6
band found in flies homozygous for Est-6S. The
enzyme specified by Est-6F is completely inac-
tivated by exposure to 60°C for 10 min (Wright,
1964, DIS 39: 60). Heterozygotes, Est-6F/Est-6S,
exhibit both the fast and slow esterase 6 bands and
do not contain a hybrid esterase 6 with an interme-
diate electrophoretic mobility. No morphological
difference between Est-6F and Eat-6S homozygotes
is apparent. Both have been found together in wild
populations and in numerous wild type and mutant
stocks. Esterase 6 of Wright corresponds to es-
terase D of Beckman and Johnson. RK3.
ex; expanded
discoverer: Maclntyre, 63d. From Stern and Bridges, 1926, Genetics 11: 503-30.
references: Wright, 1964, DIS 39: 60.
phenotype: Produces an esterase 6 with the same ex; expanded
electrophoretic mobility as that produced by location: 2-0.1.
Est-6F but that is stable to treatment with 60°C origin: Spontaneous.
for 10 min. RK3. discoverer: Bridges, 17k21.
Est-6S: Esterase 6-Slow references: Stem and Bridges, 1926, Genetics 11:
origin: Naturally occurring allele. 514 (fig.).
discoverer: T. Wright. phenotype: Wings extremely wide and large, some-
references: 1963, DIS 37: 53. times curved and divergent. Effect produced in
1963, Genetics 48: 787-801 (fig.). prepupal wing, probably by influence on cell
1964, DIS 39: 60. division (Waddington, 1940, J. Genet. 41: 75-139).
phenotype: Produces slowly migrating esterase 6 Eyes slightly reduced in size and roughish. Body
that is stable to treatment with 60°C for 10 min. large. RK.2.
RK3. cytology: Salivary chromosome location in or near
Est-C?: Esterase C-Fast 21C3 (Lewis, 1945, Genetics 30: 137-66).
location: 3-49 (based on 2/68 crossovers with Aph
and 9/43 with Est-6). origin: Recovered among progeny of flies treated
origin: Naturally occurring allele. as larvae with natural gas.
discoverer: Beckman and Johnson. discoverer. Mickey, 48k.
reference*: 1964, Hereditas 51: 212-20 (fig.). references: 1950, DIS 24: 60.
phenotype: Esterase C is one of six postively phenotype: Slightly more extreme than ex. RK2.
migrating esterases that can be demonstrated with Exaggeration of Bat: see E(B)
a-aapiithyl acetate and Fast Blue RR after starch exi: exiguous
gel electrophoresis of single fly homogenates for location: 1-51.5,
3 hr at room temperature at a voltage gradient of origin: Induced by 2-choroethyl methaneaulfonate
6—8 v per cm, using Poulik's discontinuous buffer (CB. 1506).
MUTATIONS 89
discoverer: Fahmy, 1956. 1946, DIS 20: 88) and its presence on Df(4)ll =
references: 1958, DIS 32: 70. Df(4)102E2-10;102F2-10 (Hochman).
phenotype: Small fly with rather dusky body color.
ey-
Not easily classified. Viability and fertility good origin: Spontaneous.
in male, slightly reduced in female. RK3. discoverer: Nonidez, 1919.
expanded: see ex references: Bridges, 1935, Biol. Zh. (Moscow) 4:
*exr: extra venation 401-20 (fig.).
location: 1- (associated with In(l)exr). phenotype: Eye reduced to one-half to one-fourth
origin: Induced by triethylenemelamine (CB. 1246). wild-type area, with relatively little variability.
discoverer: Fahmy, 1952. Cephalic complex smaller than wild type at 25 hr
references: 1958, DIS 32: 70. after hatching from egg at 27°C; subsequent
phenotype: Eyes slightly rough and smaller than growth rate same as wild type [Medvedev, 1935,
normal. Wings have irregularly distributed extra Z. Induktive Abstammungs- Vererbungslehre 70:
vein tissue. Males viable and fertile; females 55-72 (fig.); 1935, Tr. Inst. Genet. Akad. Nauk
viable but sterile. RK3A. SSSR 10: 119-51; Steinberg, 1944, Proc. Natl.
cytology: Associated with In(l)exr = Acad. Sci. U.S. 30: 5-13]. RK1.
In(l)l 2E8-10; 15D1 -3. other information: Most often used ey allele.
*ext: extended
location: 2- (not located),
discoverer: Stroher, 1958.
references: Mainx, 1958, DIS 32: 82.
phenotype: Wings held out at about a 75° angle from
body axis, are wavy and gradually curve down-
ward. Distal parts of wings often crumpled and
folded. Halteres normal. Function of wings ey4; eyeless-4
reduced. Viability and fertility good. RK3. Edith M. Wallace, unpublished.
Ext: Extras
location: 1-15.2.
discoverer Schultz, 3318. origin: Spontaneous.
phenotype: Heterozygous female has thickened, discoverer: Li, 25fl0.
branched, and extra veins. Overlaps wild type. references: Bridges, 1935, Biol. Zh. (Moscow)
Lethal in male. RK3. 4: 401-20.
ext-b: see bat phenotype: Eye size and variability intermediate
Ext-sct-3: see Su(sc) between ey and ey2. Optic disk of mature larvae
extended: see exf about 20 percent size of wild-type disk (Chen,
extended-b: see bat 1929, J. Morphol. 47: 135—99). Expression and
Extra sex comb: see Sex fertility of both males and females highly suscep-
extra sex combs: see esc tible to modification of genetic background
extra venation: see exr (Spofford, 1956, Genetics 41: 938-59). RK2.
Extras: see Exf *ev34g
ey; eyeless origin: Occurred among progeny of cold-treated fly.
location: 4-2.0 (located in diplo-4 triploids by discoverer: Gottchewski, 34g26.
Sturtevant, 1951, Proc. Natl. Acad. Sci. U.S. 37: references: 1935, DIS 4: 15.
405-7). phenotype: Strong allele of ey, like ey2. RK1.
discoverer: Hoge, 14e. origin: Spontaneous.
references: 1915, Am. Naturalist 49: 47—49. discoverer: Spencer, 36e25.
Bridges, 1935, Biol. 2h. (Moscow) 4: 401-20 (fig.), references: 1937, DIS 7: 8.
phenotype: Eye reduced to three-fourths to one-half phenotype: Like ey, expression variable; viability
normal area, but varies from no eyes to extensive poor. RK3.
overlapping of wild type. Less extreme at low *eY46l
temperatures. Optic ganglia reduced (Richards origin: X ray induced.
and Furrow, 1922, Proc. Oklahoma Acad. Sci. 2: discoverer: King and Poulson, 461.
41—45). Variability in size of eye affected by references: 1948, DIS 22: 54.
environmental and genetic conditions (Morgan, phenotype: Eye smaller than ey2, conical in pro-
1929, Carnegie fast. Wash. Publ. No. 399: file, may be notched or partly covered with
139-68; Baron, 1935, J. Exptl. Zool. 70: cuticle. Sometimes bristle-covered palps protrude
461-90). RK2. from border of eye. Palps often found on bucca.
cytology: Placed between 1O2C15 and 102E10 on Often dark smudges on gena. Extra vibrissae and
basis of the absence of ey + from the 2LD4^P ele- buccal bristles. Anterior verticals usually
ment of T(2;4)h~ T(2;4)25E;102C15'Dl (Morgan, doubled; posterior verticals thickened, shortened,
and bent or split at tips. ey46i/ey2 intermediate eyK; eyeless from Oregon-K
between the two homozygotes. Viability fair. origin: Spontaneous.
RK2. discoverer: Sang and McDonald.
*eyAD: eyeless of A. Das references: 1954, J. Genet. 52: 392-412 (fig.).
origin: Spontaneous. Sang and Burnet, 1963, Genetics 48: 1683—1700.
discoverer: Das, 63a7. phenotype: Eyes reduced in varying degrees from
references: Sarkar, 1963, DIS 38: 28. eyelessness to wild type. Supernumerary antennae
phenotype: Bye varies from absence to normal. occur in a small proportion of flies, especially in
RK2. flies with no eyes. Eye size depends on diet.
RK2.
eyopt: eyeless-ophthalmoptera
origin: Spontaneous derivative of ey2. On chromo-
some 4; possibly separable from ey2.
discoverer: E. Goldschmidt.
references: Goldschmidt and Lederman-Klein, 1958,
J. Heredity 49: 262-66 (fig.).
phenotype: Eyes small with variable outgrowths.
RK2.
ey R : eyeless-Russian
discoverer: Sacharov, 23hl3.
references: Serebrovsky and Sacharov, 1925, Zh.
eyD: eyeless-Dominant Eksperim. Biol. 1: 75—91.
Left: head. Right: first pair of legs. Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20.
From Patterson and Muller, 1930, Genetics 15: 495- phenotype: Eye reduced to one-half or one-fourth
577. normal area, reduction occurring chiefly in
anterior part. Similar to ey2. RK1.
eyD; eyeless-Dominanf *eyfu; eyeless-fumorous
origin: X ray induced. origin: Spontaneous,
discoverer: Muller, 27k. discoverer. Datta, 63c26.
references: Patterson and Muller, 1930, Genetics synonym: tu-h^^: tumor head-63.
IS: 495-577 (fig.). references: Sarkar, 1963, DIS 38: 28.
Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20. phenotype: Abnormal growths in various regions of
1935, Tr. Dinam. Razvit 10: 463-73. the head; protuberances on or near eyes; small
phenotype: Eyes small, outline irregular, displaced eyes; aggregations of hairs in antennal region.
toward top and rear. Head large, often with dupli- RK2.
cated antennae or ocelli. Sex combs always very other information: Allelism shown by Fahmy.
large. Second joint of tarsi shortened to give *eyW59: eyeless of Wfc/fe
lumps, sometimes very conspicuous. Fully origin: Spontaneous.
dominant in triplo-4 flies (Sturtevant, 1936, discoverer: White, 59f.
Genetics 21: 448). Eye size of B; ey-*V+ males references: Meyer, 1959, DIS 33: 97.
larger than of B alone. Produces extreme pheno- phenotype: Eyes small or absent. More extreme at
type in combination with D, JD/+; eyD/+ almost higher temperatures. Good viability and fertility.
completely lethal (Sobels, Kruijt, and Spronk, RK1.
1951, DIS 25: 128). Homozygous lethal. Homo- *Eye: Eyeless dominant in chromosome 2
zygotes die during larval period and many eyD/+ location: 2-62.7.
flies die as pupae (Hochman). RK2. origin: Probably ultraviolet induced.
cytology: Salivary chromosomes show duplication discoverer: Edmonds on, 51g.
of about a dozen bands inserted into middle of synonym: ey-HD.
fourth chromosome as a reversed repeat. Source references: 1952, DIS 26: 60.
of duplication unknown (Bridges, 1935). phenotype: Eyes may be greatly reduced in size,
other information: May not be an allele of ey. with frequent doubling of antennae. Overlaps wild
*6yD39k type, especially in old vials. Recessive in
origin: X ray induced. triploids. Eye/+'f &yD/+ has smaller eyes than
discoverer: Suttoo, 39k. either alone. Homozygous lethal. RK3.
references: Hinton, 1940, DIS 13: 49. eye gone: see eyg
phenotype: Eyes appear as a knot or two separated eyeless: see ey
knots. Homozygote overlap* wild type in 75 per- Eyeless dominant in chromosome 2: see Eye
cent of flies. Heterozygote overlaps wild type in Eyeluf. see Eyl
only 50 percent of flies (T. Hinton, 1942, Am. eyes reduced', see eyr
Naturalist 76: 219-22). RK3. eyg: eye gone
cytology: Salivary chromosomes normal. location: 3-35.5.
MUTATIONS 91
origin: Spontaneous. affected, but this is visible only at high magnifi-

discoverer: Ives, 40g20. cations. Treatment with methylurea causes normal
references: 1942, DIS 16: 48. bristle formation (De Marinis). Developmental
phenotype: Eyes and head much smaller than studies [Lees and Waddington, 1942, Proc. Roy.
normal. Considerable pupal mortality, probably Soc. (London) Ser. B 131: 87-110 (fig.); Lees and
from inability to push open pupa cases. Adults Picken, 1945, Proc. Roy. Soc. (London), Ser. B
normal in viability and productivity. Character 132: 396—423 (fig.)] show nature of pupal bristle
subject to genetic modifiers and possibly environ- secretion is affected. Suppressed by su(f). RK1.
mental influences. Expression varies from com- cytology: According to Demerec and Sutton, bands
plete absence of facets to formation of about 100 15F1-5 are involved (Demerec, Kaufmann, Fano,
facets. RK2. Sutton, and Sansome, 1942, Carnegie Inst. Wash.
*Eyl: Eyeluf Year Book 41: 191). This can be narrowed to
location: 1-18. 15F1-3 on the basis of Df(l)f2S7-S -
origin: Spontaneous. Dt(l)l 5E7-F1.-15F2-4.
discoverer: Marzluf. other information: Green (1955, Proc. Natl. Acad.
phenotype: One or both eyes reduced in size. Sci. U.S. 41: 375-79; 1956, Proc. Natl. Acad.
Expression varies from slight reduction to Sci. U.S. 42: 73—77) showed the forked mutants
absence of eye. Sometimes extraneous materials can be assigned to either of two pseudoallelic
protrude through eye; frequently one or more series, f is a member of the right series. Back
duplicated antennae present. Penetrance incom- mutations to f+ occur spontaneously and their
plete; viability good. In aged and crowded cul- incidence is not increased by X rays (Green,
tures, both penetrance and expressivity increased. 1959, Proc. Natl. Acad. Sci. U.S. 45: 16-18;
Third chromosome carries important modifiers Lefevre and Green, 1959, Genetics 44: 769—76).
affecting penetrance, and different wild-type and
mutant stocks carry different modifiers. Pene-
trance lower at 18°C than at 25°C. RK3.
eyr: eyes reduced
location: 3-103.
origin: Found among flies grown on food containing
copper sulfate.
discoverer: Edwards and Gardner, 1962.
1966, Genetics 53: 785-98.
phenotype: Eyes vary from normal to absence of
ommatidia. Shows some degree of dominance;
many heterozygotes have some eye abnormality,
usually a nick in anterior region of one or both
eyes; an abnormal growth of wing tissue may be
associated with the nick, eyr; ey4 flies have very
small heads, usually without ommatidia. Viability
greatly reduced. RK2.
X4 XI
X3 h h forked
X2 36 a Edith M. Wallace, unpublished.
3N 5
3 I
l I
discoverer: Bridges, 19kl4.
Map of the / locus references: 1938, DIS 9: 46.
Drawn from Green, 1956, Proc. Natl. Acad. Sci. U.S. Green, 1955, Proc. Natl. Acad. Sci. U.S. 41:
42: 73-77. 375-79.
phenotype: Slight allele of /. Ends of bristles
f: forked twisted or split. Not suppressed by su(f). RK2.
location: 1-56.7. other information: A member of the left pseudo-
origin: Spontaneous. allelic series.
discoverer: Bridges, 12kl9. f3N
references: Morgan and Bridges, 1916, Carnegie origin: Spontaneous derivative of /.
Inst. Wash, Publ. No. 237: 58 (fig.), discoverer: Green.
phenotype: Bristles shortened, gnarled, and bent, references: 1955, Proc. Natl. Acad. Sci. U.S. 41:
with ends split or sharply bent. Hairs similarly 375-79.
1959, Proc. Natl. Acad. Sci. U.S. 45: 16-18. cytology: Associated with In(l)i257"4 =
Lefevre and Green, 1959, Genetics 44: 769—76. In(l)15F2-16Al;16D2-El.
phenotype: Expression similar to /. Unlike f, does f2S7-5
not respond to su(f). RK1. origin: X ray induced.
other information: At the left f sublocus (Green). discoverer: Demerec, 33k.
Reverts spontaneously, and X rays delivered to phenotype: Lethal in male and cell lethal. RK2A.
oogonia and oocytes cause high incidence of cytology: Associated with Dt(l)f257-5 -
reversion. Irradiation of postmeiotic stages in the Df(l)15E7-Fl;15F2-4 (Sutton).
male does not cause back mutation (Green 1959; *{2S7-6
Lefevre and Green 1959). orjgin: Spontaneous; recovered originally as a Bar
{5 reversion.
origin: Spontaneous. discoverer: Bridges, 14i25.
discoverer: Bridges, 21b. references: 1917, Genetics 2: 445—65.
references: 1938, DIS 9: 47. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
phenotype: Bristles and hairs much curled and Genet. 2: 164, 226.
twisted. A rather extreme allele of f. Suppressed Sutton, 1943, Genetics 28: 99.
by su(f). RK1. phenotype: Lethal and cell lethal. RK2A.
other information: A member of the right f pseudo- cytology: Associated with Dt(l)P57-6 =Df(l)15E4-
allelic series (Green, 1955). F1;16A7-1.
*f34b other information: Identity of Dt(l)f2*7-6 w i t h the
origin: X ray induced. Bridges f B deficiency claimed in the original
discoverer: Stone, 34b. edition, but this does not seem well established.
references: 1935, DIS 4: 63. Bridges's case was the first recorded deficiency;
phenotype: Subliminal allele of f. Wild type in in 1925 it was claimed to be lost. Di(l)f257-6
males and homozygous females, but gives a weak obtained by Demerec from Pasadena in 1933 as a
forked phenotype when heterozygous with / or f5. t B deficiency.
RK3. *f2S7-9
*(34» origin: X ray induced; recovered with simultaneous
origin: X ray induced. reversion of BiB*.
discoverer Oliver, 34e4. discoverer: Demerec, 34c.
references: 1939, DIS 12: 48. phenotype: Lethal in male and cell lethal. RK2A.
phenotype: Like f. RK1. cytology: Associated with Df(l)f2S7-9 =
{36a Df(l)l 5E7-Fl;16D2-4.
discoverer: Ives, 36a27. *f2S7-J5
phenotype: The most extreme f allele. Hairs and origin: X ray induced.
bristles extremely crooked. RK1. discoverer: Demerec, 35a.
other Information: A member of the right / pseudo- phenotype: Lethal. RK2A.
allelic series (Green). cytology: Associated with T(l;2)f257mls =
*{42 T(l;2)13E9-10;15E2-3;24F (Sutton).
discoverer: Anderson, 42c30. /257-I9
references: Oliver, 1942, DIS 16: 53. origin: X ray induced.
phenotype: An extreme allele like is. RK1. discoverer: Hoover, 35h.
*fSla phenotype: Lethal, extreme forked. RK2.
origin: X ray induced. cytology: No major band missing (Kaufmann).
discoverer: Green, 51a. *f2S7-22
references: Lefevre and Green, 1959, Genetics 44: origin: X ray induced in BiB* X chromosome.
769-76. discoverer: Demerec, 36c.
phenotype: An extreme f like f36a. Not suppressed phenotype: Lethal. RK2A.
by au{f). RK1. cytology: Associated with T(l;2)t2^7~22 -
other information: Not observed to revert sponta- T(1;2)4D2-3;8F;15E4-F1;39E;41F-42A (Kaufmann).
neously; reversions not induced by X rays. *f2S7-24
/56« origin: X ray induced.
origin: Spontaneous. discoverer: Demerec, 36e.
discoverer; Williams, 56e. phenotype: An extreme /. Lethal. RK2.
references: 1956, DIS 30: 79. cytology: No major band missing (Kaufmann).
phenotype: Like i. RK1. *{257-27
origin: X ray induced. discoverer: Demerec, 381.
discoverer: Demerec, 33j. phenotype: Lethal- RK2A.
phwitotyp*: Bristles and hairs strongly forked. cytology: Associated with Df(l)l357-27 - Df(l)14F6-
Viability and fertility good. RKIA. 15Al;l5FS-6 (Sutton).
MUTATIONS 93
*f257-28 fB27
origin: X ray induced. origin: X ray induced in BM2 male.
discoverer: Sutton, 40h. discoverer: Belgovsky, 361.
phenotype: Lethal. RK2A. references: 1940, DIS 13: 47.
cytology: Associated with Df(l)f257-28 = Df(l)15E7- phenotype: Males have mostly normal bristles; a
F1;16E5-F1 (Sutton). few reduced like a Minute; rarely forked. fB27/f
*/257-29 are mosaic for forked. fB27/fB27 females rarely
origin: X ray induced. survive, those that do sometimes have reduced
discoverer: Bishop, 401. bristles or notched wings or both, and are sterile.
phenotype: Very slight f effect. Male sterile. RK2A. More extreme in combination with E(f). RK3A.
cytology: Associated with T(l;3)f257-2 9 -
T(1:3)15F5-16A1;64 (Sutton). * / « : forked of Hexter
*{257-3 0 origin: Spontaneous.
origin: X ray induced. discoverer: Hexter.
discoverer: Bishop, 41a. synonym: f*3.
phenotype: Forked bristles; viability and fertility references: Green, 1956, Proc. Natl. Acad. Sci.
good. RK1. U.S. 42: 73-77.
cytology: Salivary chromosomes apparently normal phenotype: Like f but not suppressed by su(f). RK1.
(Sutton). other informotion: A member of the right f sublocus.
*f2S7-31 /*"; see f+ih
origin: X ray induced. fa*: see fx
discoverer: Bishop, 41a.
phenotype: Lethal. RK2A. fx: forked from X irradiation
cytology: Associated with Df(l)f257-31 =Df(l)15E7- origin: X ray induced, simultaneously with f+il*.
F1.-J5F5-6 (Sutton). synonym: f"; forked-mottled = fx / + ^ .
discoverer: Muller.
f+//»; forked-wild type in heterochromatin references: 1946, DIS 20: 88-89.
origin: X ray induced simultaneously with fx. 1947, DIS 21: 71.
synonym: P": forked-mottled = fx f+if>. Muller and Oster, 1957, DIS 31: 141-44.
discoverer: Muller. Oster, Ehrlich, and Muller, 1958, DIS 32: 144-45.
references: 1946, DIS 20: 88-89. phenotype: A medium /• Suppressed by su(f). RK1.
1947, DIS 21: 71. cytology: Salivary chromosomes appear normal
Muller and Oster, 1957, DIS 31: 141-44. (J. I. Valencia),
Oster, Ehrlich, and Muller, 1958, DIS 32: 144-45. other information: Located to right of f3-. Does not
phenotype: f +ih with any f allele has normalizing cross over with /. Is not induced to revert by X
effect. Patches of bristles and occasionally whole rays. See last two references for relation between
fly is wild type. An extra Y chromosome enhances fx and
the normalizing effect. RK2A.
cytology: Salivary chromosomes appear normal *fX7
Q. I. Valencia). origin: X ray induced.
other information: Apparently, £**** is all or part of discoverer: Green.
the normal allele of f transposed to the proximal references: 1956, Proc. Natl. Acad. Sci. U.S. 42:
heterochromatin of the X chromosome, where it 73-77.
variegates. phenotype: Like f but not suppressed by su(f). RK1.
*fB; forked of Belgorsky other information: In right f sublocus.
origin: X ray induced in In(l)scs or Jn(l)BM2. *fX2
discoverer: Belgovsky, 1936-1937. origin: X ray induced.
references: 1937, DIS 8: 7. discoverer: Green.
1938, Izv. Akad. Nauk SSSR, Ser. Biol., 1017-36. references: 1956, Proc. Natl. Acad. Sci. U.S. 42:
1940, DIS 13: 47-48. 73-77.
other information: A series of several dozen / phenotype: Like f but not suppressed by su(f). RK1.
alleles of different strengths. Analyzed genet- other information: In left f sublocus.
ically but not cytologically. +fX3
fBIS origin: X ray induced.
origin: X ray induced in B ^ 2 male, discoverer: Green.
discoverer Belgovsky, 361. references: 1956, Proc. Natl. Acad. Sci. U.S. 42:
references: 1940, DES 13: 47. 73-77.
phenotype: Shows variegated expression of f. More phenotype: Like /but not suppressed by su(f). RK1.
extreme in combination with E(f). RK2A. other information: In left sublocus of forked.
cytology: G#netic data indicate that the mutation *fX4
is associated with a reinversion of the BM3 origin: X ray induced.
inversion. BMS phenotype reverted. discoverer Green.
references: 1956, Proc. Natl. Acad. Sci. U.S. 42: references: Muller and Altenburg, 1921, Anat.
73-77. Record 20: 213.
phenotype: Like f but not suppressed by su(f). RK1. Muller, 1935, DIS 3: 30.
other information: In left sublocus of forked. phenotype: fa^/fa resembles fa/fa; not notched.
fa: facet Homozygous lethal. RK2.
location: 1-3.0.
discoverer: Bridges, 14b.
references: Morgan and Bridges, 1916, Carnegie
phenotype: Eyes of all males moderately rough
owing to irregularity in size, shape, and arrange-
ment of facets. Eyes of females less rough, with
about 10 percent overlap of wild type. Eye
abnormality caused by overgrowth of secondary
pigment cells, which compresses cones and
causes overlying corneal facet to bulge
(Waddington and Pilkington, 1942, DIS 16: 70).
Wings have apical nicks in 0.25 percent of males
and 0—5 percent of females. N/fa has rough eyes
of fa as well as a Notch phenotype; that is, fa has
a pseudodominant effect with Notch. RK2.
cytology: Salivary chromosome locus placed at 3C7
by Slizynska (1938, Genetics 23: 291-99).
Salivary chromosomes normal (Welshons). fa": facet-notched
other information: A member of the pseudoallelic From Glass, 1933, J. Genet. 27: 233-41.
series at the Notch locus (Welshons and
Von Halle, 1962, Genetics 47: 743-59; Welshons, fa": facet-notched
Von Halle, and Scandlyn, 1963, Proc. Intern. origin: X ray induced in In(l)dl-49.
Congr. Genet., 11th, Vol. 1: 1-2), located discoverer: Glass, 1929.
between JV 55eJjr and fano. references: 1933, J. Genet., 27: 233—41 (fig.).
phenotype: Wings have apical nicks or notches in
fa3: see spl 90—100 percent of males, but only about 8 percent
*fado-vg: facet-dominigene for vestigial of homozygous females. Eyes not rough. fa/fan is
origin: Spontaneous. wild type. Viability and fertility excellent. RK2
discoverer: Goldschmidt. in male.
references: 1935, Z. Induktive Abstammungs- cytology: fa" is on an In(l)dl-49 chromosome but
Vererbungslehre 69: 38-131 (fig.)- should be separable from the inversion.
1935, Biol. Zentr. 55: 535-54 (fig.). fano; facet-notchoid
Gardner, 1942, Univ. Calif. (Berkeley) Publ. location: About 0.05 unit to right of fa.
Zool. 49: 85-102. origin: X ray induced.
phenotype: By itself it is wild type. fad°-v6/fa discoverer: Bauer, 1943.
shows rough-eye character of fa. fado~v6/fado"vi', references: 1943, Z. Induktive Abstammungs-
v&/+ produces some wing notching. RK3. Vererbungslehre 81: 374-90 (fig.).
other information: Presumed by Goldschmidt to Welshons, 1958, Proc. Natl. Acad. Sci. U.S. 44:
enhance dominance of vg and thus termed a 254-58.
' * d ominigene''. 1958, Cold Spring Harbor Symp. Quant. Biol. 23:
fa9: facet-glossy 171-76.
origin: Spontaneous. Welshons and Von Halle, 1962, Genetics 47:
discoverer: Pratt, 1962. 743-59.
phenotype: Eyes have facets more irregular than fa, phenotype: Wings of both sexes notched at ends of
but surface is smoothed, giving a glossy effect. L3 and LA veins; other longitudinal veins
Pigment distribution may be uneven, contributing enlarged and show deltas. Somewhat less extreme
to an impression of altered eye color. No wing ef- at elevated temperatures. Up to 5 percent of fano
fect. Eyes of fa*//a intermediate between the two males from aged cultures show hyper- and hypo-
. Complementary with apt, fano, nd, developed external genitalia (Kroeger, 1960,
(Vtm Halle and Welshons). RK.1. J. Morphol. 107: 227—32). Heterozygote shows
cytology: Salivary dtarotBOSoroe* normal (Welshons). extremely weak dominance. fetno/N almost com-
other infaraMrttoa: Loc@t©d to the left of fe**0; not pletely lethal. Rare survivors have exaggerated
y«t ••pttnted from fa (Wmlshoas,). Notch phenotype. taao/fa closely resembles wild
*hh fwet-htlml type; nicks in wings appear infrequently. RK1.
dJ«c«v«r*r: Matter, 19b. cytology: Salivary chromosomes normal (Welshoos).
MUTATIONS 95
other information: A member of pseudoallelic series normal eggs also laid. Hatchability reduced and
at the Notch locus, located between fa and variable. RK3.
N264-40, fg: see spdl&
facetious: see rgp *fi: frail
/as/: see rgp location: 1-53.
fat: see ft origin: Recovered among progeny of flies treated
faulty chaetae: see fc with Janus green.
*fb: fine bristle discoverer: Muller, 28e20.
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- phenotype: Wings nearly as small as tn, thin and
phenylalanine (CB. 3026). frail. Bristles fine. Fly weak. Viability 10—30
discoverer: Fahmy, 1954. percent wild type. RK3.
references: 1958, DIS 32: 70. *fil: fine lash
phenotype: Thin, slightly shortened bristles. location: 1-56.8.
Occasional scalloping of wing margins. Delayed origin: Induced by L-p-NN-(2-chloroethyl)amino-
emergence. Good viability and fertility both phenylalanine (CB. 3025).
sexes. RK3. discoverer: Fahmy, 1953.
*fc: faulty chaetae references: 1959, DIS 33: 86.
location: 1-0.9. phenotype: Thin, slightly shorter bristles. Eyes
origin: Induced by DLT3-NN-di-(2-chloroethyl)amino- reduced in size; posterior border very close to
phenylalanine (CB. 3007). orbital bristles. Both sexes viable and fertile.
discoverer: Fahmy, 1954. RK3.
references: 1958, DIS 32: 70. other information: Two alleles induced by CB. 1528.
phenotype: Short, thin bristles. About one-third of fin: finer
flies show either absence or duplication of one location: 1-29.6.
scute liar bristle. Viability and fertility good both origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
sexes. RK2- phenylalanine (CB. 3026).
*fd: furled discoverer: Fahmy, 1954.
location: 1- (rearrangement). references: 1959, DIS 33: 86.
origin: Induced by P 3 2 . phenotype: Fly slightly smaller than normal, with
discoverer Bateman, 1949. shorter, thinner bristles. Delayed eclosion. Males
references: 1950, DIS 24: 54. viable but sterile. RK3.
1951, DIS 25: 77. fine bristle: see fb
phenotype: Like vestigial, but with immovable fine chaetae: see fnc
mouth parts and fully extended proboscis. Dies fine lash: see fil
early, perhaps owing to failure to ingest. Viability fine macros: see fm
at eclosion good. RK3A. finer: see fin
cytology: Associated with T(l;3)fd= T(1;3)7A;86E +
In(3R)89C;96A (Darby).
Female lethal: see Fl
female sterile: see fs( )
Female sterile(2) Dominant: see Fs(2)D
fern: see is(2)B

fes(2)K: see is(2)K
ff: fluff
location: 1-57.7.
origin: Induced by 2-chloroethyl rnethanesulfonate
(CB. 1506).
phenotype: Extremely fine short bristles. Wings fj: four jointed
slightly rounded at tips. Males and females viable Second and third legs.
and fertile; eclosion delayed. RK3. Edith M. Wallace, unpublished.
other information: One allele induced by CB. 1414.
*fft: fused filament //; four jointed
location: Not located, location: 2-81.
discoverer: Robertson and Reeve. discoverer: Schultz, 31dl.
references: 1954, DIS 28: 78. phenotype: Tarsi four instead of five jointed. Legs
phenotype: Chorionic filaments of eggs laid by fit short and stocky. Enhanced by s s a and ss*B
female usually fused into a single structure. A few (Villee, 1945, Genetics 30: 26-27). Development
96 GENETIC VARIATIONS OF DROSOPHILA MELANOGAS
similar to that of dachs LWaddington, 1943, J. phenotype: Fertility of Fls/+ females may be
Genet. 45: 29-43 (fig.)]- Wings shorter, broader, reduced. Fls/Fl* females sterile. Fl/Fls fema
with crossveins conspicuously closer together; lethal. RK3.
veins diverge at greater angle. Effect visible in fla: flat eye
prepupal wing (Waddington, 1940, J. Genet. 41: location: 1-2.4.
75—139). Eyes smaller, ellipsoid, coarse origin: Induced by L-p-NN-di-(2-chloroethyl)amin
textured; head foreshortened. RK2. phenylalanine (CB. 3025).
discoverer: Ives, 40e. phenotype: Smaller fly, with smaller and less-
references: 1941, DIS 14: 39. curved eyes. Wings extremely variable, from
phenotype: Venation irregularities more extreme normal, through incised margins, to crumpled
than those described for fj, but variable and may vestigial stumps. Not easily classified. Viabil
overlap wild type. Production of fj*Oe flies low and fertility good in males but reduced in femal
in competition with wild-type flies in crowded RK3.
cultures, but normal in a homozygous stock. flap wing: see flw
Developmental period about 20 percent longer flateye: see fla
than wild type. RK2. flipper: see flp
*///: ftyless
discoverer: Cercos, 41gl5.
references: Andres, 1943, DIS 17: 48.
phenotype: Wings apparently normal, but fly cann
keep them spread and cannot fly more than a fex
inches. RK3.
*flp: flipper
//: fluted location: 2-30.
Edith M. Wallace, unpublished. origin: Spontaneous.
discoverer: Mohr, 18h5.
fl: fluted references: Bridges and Mohr, 1919, Genetics 4:
location: 3-59.9. 304.
origin: Spontaneous, phenotype: Wings fail to expand; remain compact,
discoverer: Redfield, 211. very dark, extended, and curved slightly down-
phenotype: Wings creased lengthwise and dark. ward. Fly a wizened dwarf. Body surface dull
Overlaps wild type slightly at 25° but not at and dark. Both sexes sterile, RK3.
19°C. RK3. flp: see flw
*f!2 fluff: s e e ff
origin: Spontaneous. fluted: s e e fl
discoverer: Spencer, 36dl5. *flw: flap wing
phenotype: Like tl, RK3. location: 1-31.
Fl: Female lethal discoverer: Waletzky, 1937.
location: 1-19.1 (based on crosses with F/ s ). synonym: flp, a preoccupied symbol.
origin: Spontaneous. phenotype: Wings held out; consistently concave
discoverer. Muller and Zimmering, 1960. upward; slightly pointed. Darkened longitudinal
references: 1960, Genetics 45: 1001—2. stripe along thorax, underneath chitin; occasion,
phenotype: Viability of F1/+ females varies from bubbling, with maximum pigmentation in anterior
zero to normal, depending on maternal genotype. part of stripe. Anterior scutellars sometimes
Where viability is low, surviving females often missing or doubled. Eyes bulging; slightly
show diverse morphological abnormalities. Fl/Fl roughened. Head compressed anteroposteriorly.
females lethal; die as early embryos (Oster). Third antennal joint shortened. RK2.
tnftrm has no effect on survival of Fl/Fl or F1/+
(Zimmering and Muller, 1961, DIS 35: 103-4). Ft/Y flyless: see HI
males have normal viability and phenotype. RK2A. *fm: fine macros
cytology: Present in X containing In(l)dt-49 = location: 1-66.1.
ln(l)4D7-El;llF2-4. origin: Induced by 2-chloroethyl methansesulfonate
(CB. 1506).
Fl*: Female lethal-sterile discoverer: Fshmy, 1956.
discovmmr, Zimmering and Muller, 1961. phenotype: Small fly with narrow abdomen and
synonym; F!%. extremely short, thin bristles. Males fertile;
references: 1961, DIS 35: 103-4. viability about 50 percent wild type. RK3.
MUTATIONS 97
*fnc: fine chaetae margin irregularity. Viability variable, from 16—90

location: 1-34.9. percent wild type. Females rarely fertile. Char-
origin: Induced by S-2-chloroethylcysteine acter less extreme at low temperature. RK3.
(CB. 1592). *fr°: fringed-zero
references: 1959, DIS 33: 86. discoverer: Bridges, 15a20.
phenotype: Extremely fine short bristles. Body references: Bridges and Morgan, 1919, Carnegie
parts disproportionately reduced; reduction least Inst. Wash. Publ. No. 278: 257 (fig.).
marked on head and most marked on abdomen. phenotype: Wing margins have bare spots;
Wings broad and slightly rounded at tips, remaining hairs frayed and irregular. Wings
occasionally with incisions of margin. Eyes smaller, discolored, somewhat divergent. Like fr
slightly brighter red than normal. Males viable but but less extreme. RK3.
sterile. RK3. other information: Lost before fr, fr2, and frdi were
fo: folded found.
location: 1-63. fr2
discoverer: Grossman, 1932. origin: Spontaneous.
references: 1934, DIS 1: 30. discoverer: Novitski, 37a22.
phenotype: Wings remain unexpanded in a varying synonym: tan: trimmed.
percentage of flies. Balancers shriveled and post- references: 1937, DIS 8: 10, 13.
scutellars bent forward. Overlaps wild type. RK3. Lewis, 1938, DIS 10: 55-56.
Fo; Forkoid phenotype: Wings scalloped and fringed on all
location: 2-107 (between or and sp). margins, slim and coarse textured, often divergent
origin: X ray induced. and uplifted. Eyes rough. Some extra bristles
discoverer. Mohler, 5 8 c l 8 . present. Abdomen slightly offset, as in fr.
references: 1960, DIS 34: 52. Female sterile. Classification at 25°C excellent;
phenotype: Heterozygote shows reduction in size of characters less extreme at 19°C. Viability
bristles and weak forking of head and posterior generally good, but erratic. RK2.
thoracic bristles. Using Dp(2;3)P, it may be shown *frdl; fringed-dibro
that the expression of +/+/Fo <+/Fo <+/Fo/Fo; origin: Spontaneous.
+/F0/F0 shows extreme forking of all bristles and discoverer: Bridges, 17kl9.
is sterile. Homozygous lethal. Fo interacts with f synonym: dihro.
alleles to produce extreme f bristles. RK1. references: Lynch, 1920, Genetics 4: 527—28.
cytology: Located between 58E3 and 60B10, on phenotype: Spread wings with scalloped margins.
basis of its inclusion in Dp(2;3)P = Dp(2;3)58E3- Eyes rough. Very inviable; both sexes sterile.
F2;60D14-E2;96B5-Cl but not in Df(2R)Px = RK3.
Df(2R)60B8-10;60Dl-2 (Mohler) or in the deficiency frail: s e e fi
for the tip of 2R derived from T(l;2)Bld =
T(1;2)1C3-4;6OB12-13 (Armentrout). Frd: Freckled
location: 2-103.1 (Nicoletti).
focal melanosfs: see me origin: X ray induced.
*fol: folded wings discoverer: M. G. Davis, 1961.
location: 2-39. references: Erlich, 1963, DIS 37: 47.
origin: Spontaneous. Barigozzi, 1963, Proc. Intern. Congr. Genet., 11th.
discoverer: Goldschmidt, 1937. Vol. 1: 207.
phenotype: Expanded wing folded. Overlaps wild 1965, DIS 40: 64.
type. RK3. phenotype: Pupa and young fly characterized by ac-
folded: see fo cumulation of dark pigment; in older fly, pigment
folded wings: see /b/ becomes concentrated in black specks scattered
forked: see f throughout body, head, and legs. Homozygous
Forkoid: see Fo lethal. RK2.
four jointed: see // other information: Barigozzi claims it has an extra-
chromosomal or episomal component in its trans-
fr: fringed mission.
location: 2-80. fringed: see fr
origin: Spontaneous. frizzled: see fz
discoverer: Bridges, 22c30. fs 2.: see fs(2)E
references: 1938, DIS 9: 48. fs(l)N: female sterile (1) of Nasrat
phenotype: Wings often spread; wing margins location: 1-0.0 (closely linked to me).
snipped and bristles irregular and fringelike. origin: Induced by an unspecified chemical
Eyes small and rough. Midline of abdomen at mutagen.
slight angle to ioagitudinal axis of fly. Much discoverer: Nasrat, 1952.
variability in expression; safest criterion is wing synonym: /®f2)" ae .
98
phenotype: Females sterile but no other distin- origin: Ultraviolet induced.

guishing characteristics. Males fertile. Develop- discoverer: Edmondson, 50j.
mental study by Counce and Ede [l957, J. synonym: fs2.1.
Embryol. Exptl. Morphol. 5: 404-21 (fig.)]. Eggs of references: Meyer and Edmondson, 1957, DIS 25:
ta(l)N/i8(l)N females will not support develop- 72.
ment of normal embryos. About half the eggs phenotype: Sterile females do not lay eggs. Gonads
contain little or no yolk; development may or may rudimentary. RK3.
not begin in such eggs but never progresses *fs(2)E2
beyond a highly abnormal cleavage. In eggs location: 2-22.0.
containing more yolk, major effect is on synchrony origin: Ultraviolet induced.
of cleavage and blastoderm mitoses. Twenty per- discoverer: Edmondson, 1951.
cent of these embryos cease development before synonym: fs2.2.
blastoderm formation. The remainder have references: 1952, DIS 26: 61.
abnormal blastoderms and aberrant gastrulation. phenotype: Females produce eggs that appear
Final pattern of damage determined by degree of normal but do not hatch. Fertile in heterozygotes
abnormality of earlier stages, but some embryos with ms(2)E3 (2-28.0). RK3.
show larval differentiation. A few of the least *fs(2)E3
abnormal embryos may emerge but never move location: 2-47.5.
about or feed. Formation of polar granules origin: Ultraviolet induced.
abnormal. RK3. discoverer: Edmondson, 1951.
cytology: No detectable chromosomal rearrange- synonym: fs2.3.
ments (Slizynska). references: 1952, DIS 26: 62.
fs(2)5*6: see is(2)E9 phenotype: Female sterile; narrow curved wings.
ts(2)adp: see adpte No eggs laid. Fertile in heterozygotes with
h(2)B: female sterile (2) of Bridges ms(2)E4 (2-47.9), fs(2)E4 (2-48.5), fs(2)E5 (2-50.4),
location: 2-5. fs(2)E6 (2-54.4), ms(2)E5 (2-54.8), ms(2)E6
origin: Spontaneous. (2-54.8), ms(2)E7 (2-54.8), ts(2)E7 (2-55.2),
discoverer: Bridges, 29c2S. ms(2)E8 (2-55.6), and ms(2)E9 (2-57.0). RK3.
synonym: fes, *fs(2)E4
references: King, Sang, and Leth, 1961, Exptl. Cell. location: 2-48.5.
Res. 23: 108-17 (fig.). origin: Ultraviolet induced.
King, Koch, and Cassens, 1961, Growth 25: 45-65 discoverer: Edmondson, 1951.
(fig.). synonym: is2.4.
Koch and King, 1964, Growth 28: 325-69 (fig.). references: 1952, DIS 26: 62.
phenotype: External morphology normal. Male fer- phenotype: Very few eggs laid; female sterile. Fer-
tile; female sterile. Ovarioles of homozygous fe- tile in heterozygotes with fs(2)E3 (2-47.5), ms(2)E4
male subdivided into a series of sausage-shaped (2-47.9), fs(2)E5 (2-50.4), fs(2)E6 (2-54.4), ms(2)E5
cell aggregates, each surrounded with an ill- (2-54.8), ms(2)E6 (2-54.8), ms(2)E7 (2-54.8),
defined follicular epithelium and filled with hun- is(2)E7 (2-55.2), ms(2)E8 (2-55.6), and ms(2)E9
dreds to thousands of mitotically active oogonia- (2-57.0). RK3.
like cells [King, Burnett, and Staley, 1957, Growth *fs(2)E5
21: 239—61 (fig.)]. These cells occasionally dif- location: 2-50.4.
ferentiate into cells resembling nurse cells, which origin: Ultraviolet induced.
may have polytene chromosomes, and rarely into discoverer: Edmondson, 1951.
oocytes. la(2)B ovaries transplanted into wild-type synonym: fs2.5.
hosts in late larval stages and reciprocal trans- references: 1952, DIS 26: 62.
plant® develop autonomously (Clancy and Beadle, phenotype: Female semisterile. A few larvae hatch
1937, Biol. Bull, 72: 47-56; Bodenstein and King, normally, but most embryos degenerate. Fertile in
1963, DIS 37: 65; 1965, Z. Naturforsch. 20b: 292- heterozygotes with fs(2)E3 (2-47.5), ms(2)E4
97). RK3. (2-47.9), fs(2)E4 (2-48.5), fs(2)E6 (2-54.4), ms(2)E5
Fs(2}D: Female sterile(2) Dominant (2-54.8), ms(2)E6 (2-54.8), ms(2)E7 (2-54.8),
location: 2- (not located). ts(2)E7 (2-55.2), ms(2)E8 (2-55.6), and ms(2)E9
origin: Induced by ethyl methanesulfonate. (2-57.0). RK3.
discoverer: E. H. Grell, 65e. *fs(2)E6
pfieraotype: Heterozygous female entirely sterile, location: 2-54.4.
with underdeveloped ovaries. Heterozygous male origin: Ultraviolet induced.
ba» normal fertility. Bristles short; thorax broad discoverer: Edmondson, 1950.
sad flattened with air bubbles under cuticle. RK3. synonym: f&2.6.
eth*r information: Sterility makes genetic mapping references: 1952, DIS 26: 62.
phenotype: Female produces normal-appearing
*fs(2)Eh female sterile (2) of Edmondson eggs, which do not hatch. Fertile in heterozygotes
location: 2-57.6. with tm(2)E3 (2-47.5), ms(2)E4 (2-47.9), ts(2)E4
MUTATIONS 99
(2-48.5), fs(2)E5 (2-50.4), ms(2)E5 (2-54.8), origin: X ray induced.

ms(2)E6 (2-54:8), ms(2)E7 (2-54.8), fs(2)E7 discoverer: Gill, 59a.
(2-55.2), ms(2)E8 (2-55.6), ms(2)E9 (2-57.0), and synonym: fs(3)2*9*.
is(2)E8 (2-62.6). RK3. references: 1960, Anat. Record 138: 351.
*fs(2)E7 1961, Ph.D. Thesis, Yale Univ.
location: 2-55.2. 1962, DIS 36: 37.
origin: Spontaneous. 1963, J. Exptl. Zool. 152: 251-78 (fig.).
discoverer: Edmondson, 1950.
phenotype: Females almost sterile; produce rare
synonym: fs2.7.
surviving progeny. Oogenesis incomplete; usually
stops in early phases of vitellogenesis. Most (89
phenotype: Females produce collapsing eggs.
percent) follicles contain 32 cells instead of
Apparently, there is a weakness in the vitelline normal 16 as a result of an extra oogonial divi-
membrane, since these eggs cannot be success- sion. The 32 cells of an incipient cyst enclosed
fully dechorionated; when chorion is removed, egg in two chambers in 6 percent of the cases. Posi-
contents flow out. Fertile in heterozygotes with tion of oocyte in follicle abnormal in 28 percent of
fs(2)E3 (2-47.5), ms(2)E4 (2-47.9), fs(2)E4 (2-48.5), cases. Males partially sterile. Viability low.
fs(2)E5 (2-50.4), fs(2)E6 (2-54.4), ms(2)E5 (2-54.8), RK3.
ms(2)E6 (2-54.8), ms(2)E7 (2-54.8), ms(2)ES
(2-55.6), ms(2)E9 (2-57.0), and fs(2)E8 (2-62.6). fs(3)G3
RK3. location: 3-25.
*fs(2)E8 origin: X ray induced.
location: 2-62.6. discoverer: Gill, 59a.
origin: Ultraviolet induced. synonym: fs(3)3^9a.
discoverer: Edmondson, 49k. references: 1960, Anat. Record 138: 351.
synonym: fs2.8. 1961, Ph.D. Thesis, Yale Univ.
references: 1952, DIS 26: 62. 1962, DIS 36: 37.
phenotype: Females do not lay eggs. Fertile in 1963, J. Exptl. Zool. 152: 251-78 (fig.),
heterozygotes with fs(2)E6 (2-54.4), ms(2)E5 phenotype: Oogenesis incomplete; most follicles
(2-54.8), ms(2)E6 (2-54.8), ms(2)E7 (2-54.8), stop development during yolk deposition (after
fs(2)E7 (2-55.2), ms(2)E8 (2-55.6), ms(2)E9 stage 9). Males fertile. RK3.
(2-57.0), ms(2)E10 (2-66.5), ms(2)Ell (2-68.0), and
ms(2)E12 (2-68.2). RK3. fs(3)G5
*fs(2)E9 location: 3-49.
origin: Ultraviolet induced. discoverer: Gill, 59a.
discoverer: Edmondson, 1958. synonym: fs(3)5S9a.
synonym: fs(2)ss&. references: 1960, Ar"'. Record 138: 351.
references: 1960, DIS 34: 49. 1961, PhJD. Thesis, Yale Univ.
phenotype: Numerous infertile eggs produced. 1962, DIS 36: 37.
Viability normal. Males fertile. RK3. 1963, J. Exptl. Zool. 152: 251-78.
*fs(2)K: female sterile (2) of Kikkawa phenotype: Oogenesis incomplete; ovarioles
location: 2-100. contain excessive numbers of follicles, which
origin: Spontaneous, usually stop developing at or before stage 9.
discoverer Kikkawa, 1960. Males fertile. RK3.
synonym: ies(2)K. ft: fat
phenotype: Female sterile. Male fully fertile. RK3. origin: Spontaneous.
fs(3)Gh female sterile (3) of Gill discoverer: Mohr, 2Obi5.
location: 3-47. references: 1923, Studia Mendeliana (Brunae)
origin: X ray induced. pp. 266-87.
discoverer: Gill, 59a. 1929, Z. Induktive Abstaimmngs- Vererbungslehre
synonym: fs(3)l*9». 50: 113-200 (fig.).
references: 1960, Anat. Record 138: 351. phenotype: Abdomen short and fat. Thorax broad.
1961, Ph.D. Thesis, Yale Univ. Wings short and broad, with crossveins much closer
1962, DIS 36: 37. together than normal. Scutellutn shortened;
1963, J. Exptl. Zool. 152: 251-78 (fig.). scutellar bristles far apart. Viability good. Second-
phenotype: Eggs die in early cleavage stage. and third-instar larvae, particularly when there is
Meiosis precocious in 2 percent of oocytes; first little yeast in the food, show vacuoles in cyto-
meiotic spindle parallel to egg surface. Males plasm of salivary gland cells. Two waves of
fertile. RK3. vacuole formation. Cells with vacuoles exhibit a
slight puff in 24D-E, which is negatively corre-
fs(3)G2
lated with puffing in 21 and 22. Tip of X disfigured.
location: 3-11.
TOO GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER
possibly as a result of several small puffs inter- In(l)ClLy4R = Jn(l)4A5-Bl;17A6-Bl^lA8-

mingled with hard nonpuffed bands. In about 1 per- B1;18A3-4R] are more extreme than those in fu
cent of larvae, salivary glands distally expanded homozygote (King, 1959, DIS 33: 142-43). fu/fu
and crooked [Slizynski, 1964, Cytologia (Tokyo), ovaries transplanted into fu + hosts develop auton-
29: 330-36 (fig.)]. RK1. omously in regard to fertility (Clancy and Beadle,
cytology: Placed between 24D2 and 24F1 on the 1937, Biol. Bull. 72: 47-56; Sobels, 1950, Ex-
basis of its inclusion in Di(2L)M-zc = Df(2L)24D2- perientia 6: 139—40) and tumor formation (Boden-
5;25A2-3 but not in Df(2L)M-zB = Df(2L)24E2- stein and King, 1963, DIS 37: 65). The few normal-
Fl;25Al-2 (Morgan, Bridges, Schultz, and Curry, appearing eggs that are laid by fu/fu females
1939, Carnegie Inst. Wash. Year Book 38: 273-77). produce adults only if they have been fertilized by
ftj+-bearing sperm (Lynch, 1919, Genetics 4: 501 —
33). Eggs fertilized by fu- or F-bearing sperm de-
velop into embryos that become abnormal 5-5V2 hr
after fertilization. A general asymmetry in germ
layers is responsible for many ensuing abnormali-
ties. Such embryos never hatch but survive long
after normal embryos have become larvae LCounce,
1956, Z. Induktive Abstammungs- Vererbungslehre
87: 462-81 (fig.)], fu eggs from fu/+ mothers de-
velop normally. Heterozygous daughters from homo-
zygous mother have high incidence of abnormal ab-
dominal segmentation and, as embryos, have
abnormal musculature. This is a maternal effect
not found in the reciprocal cross, and it is temper-
ature sensitive (Armstrong and Sobels). RK1.
cytology: Salivary chromosome location of fu
appears to be in 17D or E (Green, 1953, Genetics
38: 91—105; 1953, Z. Induktive Abstammungs-
Vererbungslehre 85: 435—49).

discoverer: Edmonds on, 51e.
fu: fused references: Meyer and Edmondson, 1951, DIS 25: 72.
Edith M. Wallace, unpublished. phenotype: Like fu, but possibly slightly more
extreme. RK1.
fu; fused fvS7a
location: 1-59.5. origin: Induced by azo-mustard.
origin: Spontaneous. discoverer: Purdom, 57a.
discoverer: Bridges, 12k4. references: King, Burnett, and Staley, 1957, Growth
references: Morgan and Bridges, 1916, Carnegie 21: 239-61.
Inst. Wash. Publ. No. 237: 55-58 (fig.). phenotype: Veins L3 end L4 fused from origin to
Lynch, 1919, Genetics 4: 501-33. beyond anterior crossvein. Ocelli and ocellar and
phenotype: Veins L3 and L4 fused from base to anterior scutellar bristles sparse or absent. Length
beyond anterior crossvein, with elimination of of scute Hum generally reduced. Female sterile as
anterior crossvein and first basal cell; L3 and L4 with hi. Ovaries tumorous but to a lesser extent
fused at tip; this fusion may reach back to basal than with fu. Wings not outstretched but held in
cell. Wings usually extended. Ocelli reduced or normal position. RK1.
absent; bristles of ocellar region small or absent.
Eyes small and slightly rough. Anterior scute liar *fu57f
bristle® reduced in number, and scute Hum short- origin: Induced by azo-mustard.
ened. Female late to eclose and has decreased discoverer: Purdom, 57f.
longevity. Ovaries histologically normal at eclo- references: King, Burnett, and Staley, 1957, Growth
sioo but with half the normal number of ovarioles 21: 239-61.
(BMtty, 1949, Proc. Roy. 8oc. Edinburgh, B 63: phenotype: Wing veins L3 and L4 often completely
249—70); fecundity 7 percent normal. Dewioping fused. Wings outstretched. Ovaries tumorous.
egg chambers may fuse or become timorous with RK1.
at® [King, Boraett, and Staley, 1957, Growth 21:
239—61 (fig.)]. Proportion of tumorou* egg chambers origin: Spontaneous.
increases by 6 percent pan day. Female raised at discoverer: R. F. Grell, 1959.
1S°C stews only 10 percent th« tumor development
references: King and Smith, 1963, DIS 37: 49.
of that rmlaed at 25°. Ovarian effects i& female
phenotype: Wings like fu. Rate at which ovarian
cairying fa and • deficiency for §u [i.e.,
tumors develop is the lowest of all alleles tested;
MUTATIONS 101
number of tumorous egg chambers increases by 1

percent per day. RK1. origin: X ray induced.
*fu6lfl discoverer: MuHer, 31a.
origin: Gamma ray induced. phenotype: Extreme fw. Female sterile. RK2A.
discoverer: Fahmy, 62f. cytology: On a chromosome containing In(l)dl-49.
references: Smith and King, 1963, DIS 38: 39. fw34e
phenotype: Like fuS7a. RK1. discoverer: Duncan, 34e20.
* fa 62 i2 phenotype: Originally showed eye surface medium
origin: Gamma ray induced. folded; bristles much gnarled. Schultz and Curry
discoverer: Fahmy, 62f. report that stock in 1940 showed gnarled bristles
references: Smith and King, 1963, DIS 38: 39. and eye small but no vertical fold. RK2.
phenotype: Wings and ocelli show typical abnormali- fw49c
ties. Ovaries develop tumors. RK1. origin: Induced by P^2.
fu62f3 discoverer: R. C. King, 49c28.
origin: Gamma ray induced. references: 1949, DIS 23: 62.
discoverer: Fahmy, 62f. phenotype: Eyes furrowed; distal portions of aristal
references: Smith and King, 1963, DIS 38: 39. branches hooked; wings divergent and often
phenotype: Like fa. RK1. stringy; scutellar groove reduced. Bristles split,
*fuff: fused-formalin food bent, and often erect; acrostichal hair pattern dis-
origin: Induced by formaldehyde. tributed with whorls and naked areas. Late
discoverer: Auerbach, 1951. hatching, poorly viable, and mostly sterile.
references: Counce, 1956, Z. Induktive f w 49c/f w phenotypically intermediate but more like
Abstammungs- Vererbungslehre 87: 462—81. fw/fw than fw49c/fw49c^ R K 3
King, Burnett, and Staley, 1957, Growth 21:
239-61.
phenotype: Like fu but wings not outstretched. origin: X ray induced.
Ovaries contain tumors. Development of lethal discoverer: Garcia-Be Hid o, 59i21.
embryos roughly similar to that of fu (Counce, references: 1963, Genet. Iberica 15: 1—102.
1956). RK1. phenotype: Eyes rough and creased; facets irregular,
*fu9: fused-glider 15 percent fewer than normal. Eyes browner than
origin: Found among heat-treated flies, normal; pterine concentration reduced in the eyes
discoverer Grossman, 1932. and, except for isoxanthopterine, increased in
references: 1934, DIS 1: 30. testis sheath. Riboflavin accumulates in Mal-
phenotype: Like fu except wings more divergent and pighian tubules. Large bristles of head and thorax
females more fertile, fu^/fu females 40 percent short, thick, angled, and blunt; occasionally re-
fertile. fu&/fu6 females 20 percent fertile. In duced to stumps. Arista thick with contorted and
general, pattern of development of lethal embryos supernumerary branches. Scute Hum small with
is like that of fu (Counce, 1956, Z. Induktive groove between it and thorax reduced. Hatchability
Abstammungs- Vererbungslehre 87: 462—81). RK1. and larval development normal; larval anal plates
furled: see fd swollen and surrounded by melanotic halo. Mela-
furrowed: see fw notic anal region persists in pupa; pupa also has
fused: see fu melanotic spots elsewhere that may result in non-
fused filament: s e e fft pigmented areas on the imaginal integument. Ex-
fuzzy: see fy trusion of anterior and posterior spiracles in pre-
fw: furrowed pupa incomplete. Many fn>5 9 flies die either after
location: 1-38.3. 24-30 hr of pupal development or at the time of
origin: Spontaneous, eclosion. Fecundity of female reduced owing to
discoverer: Duncan, 14k. reduced number of ovarioles. RK2.
references: 1915, Am. Naturalist 49: 575—82.
Morgan and Bridges, 1916, Carnegie Inst. Wash. origin: X ray induced.
Publ. No. 237: 80. discoverer: Garcia-Bellido, 60k8.
Nachtsheim, Z. Induktive Abstammungs- references'. 1963, Genet. Iberica 15: 1-102.
Vererbungslehre 20: 118-56. phenotype: Like fw59 but with lower penetrance and
phenotype: Eyes with vertical fold and furrows. expressivity. RK2.
Head and scute Hum shortened. Bristles gnarled
and shortened, especially the postscutellars. Best *fww; furrowed-weak
classification character is short, blunt noto- origin: Spontaneous.
pleurals. RK2. discoverer Ives, 43b24.
cytology: Placed between 10E2 and 11A7 on the references: 1946, DIS 19: 46.
basis of its being to the right of dy, which is to phenotype: Affects only bristles, particularly the
the right of 10E2 and to the left of the X break- scutellars and postaiars. Eyes normal. Normal
point of T(l;4)A8 » T(l;4)llA6-7. fertility and viability. RK2.
fwwr: {urrowed-wrinkled discoverer: Bridges, 15bl9.

origin: X ray induced in In(l)scslLscSR+dl.49. synonym: salmon.
discoverer R. M. Valencia, 1959. references: 1916, Genetics 1: 151.
synonym: wr, Chovnick, 1958, Proc. Natl. Acad. Sci. U.S. 44:
references: 1959, DIS 33: 100. 333-37.
1965, DIS 40: 36. 1961, Genetics 46: 493-507.
phenotype: Eye surface in folds. Some bristles phenotype: Eye color deep purplish ruby, like pr;
shortened, thickened, or curved; many doubled and pinkish in young and brownish in old flies. Eyes
may be fused. Viability low. RK2. have 38 percent normal red and 56 percent normal
*fy: fuzzy brown pigment (Nolte, 1959, Heredity 13: 233—41).
location: 2-33. RK1.
origin: Spontaneous. cytology: Placed in salivary chromosome region
discoverer: Ives, 39a. 12B9-12C7 by deficiency analysis (J. I. Valencia).
references: 1940, DIS 13: 49. other information: A pseudoallelic locus composed
phenotype: Hairs on abdomen and thorax irregular of four demonstrated sites; g2 occupies the left-
and directed toward midline. Hairs on wing margins most, gSOe £he second, g and g3 the third, and gsi
erect. Resembles fz. Fertility and viability below the right-most site (Chovnick, 1961).
normal. RK2.
50e sf
_J I
Map of the g locus

From Chovnick, 1961, Genetics 46: 493-507.
discoverer: Bridges, 18c28.
Bibliog. Genet. 2: 226 (fig.).
Sturtevant and Beadle, 1939, An Introduction to
Genetics, Saunders, p. 64 (fig.).
phenotype: Eye color translucent yellowish ruby,
somewhat lighter than g. Eyes contain 16 percent
normal red pigment and 32 percent normal brown
pigment (Nolte, 1959, Heredity 13: 233-41). Mal-
pighian tubes very pale yellow at base (Beadle,
fz: frizzled 1937, Genetics 22: 587-611). Eye color auton-
From Bridges and Brehme, 1944, Carnegie Inst. Wash. omous in transplant into wild-type host (Beadle
Pobl. No. 552: 85. and Ephrussi, 1936, Genetics 21: 230). With st,
gives dark yellow to orange color; with bw, gives
fz: frizzled reddish yellow to brownish rose (Mainx, 1938,
location: 3-41.7. Z. Induktive Abstammungs- Vererbungslehre 75:
origin: Spontaneous. 256—76). Dominant in females homozygous for
discoverer: Bridges, 38bl8. car, according to Schultz; i.e., pigmentation of
phenotype: Hairs on thorax directed irregularly g2 car/g2 car — g2 car/+ car < car/car. RK1.
toward midline. Thoracic bristles also in turned other information: Occupies left-most pseudoallelic
and often wavy. Postverticals may turn outward. site of the g locus (Chovnick, 1961, Genetics 46:
Hairs on wing edge and feet nearly erect. Wing 493-507).
may be reduced. Sex combs may be irregular. Eyes 1
always rough. Resembles in. RK2. origin: Spontaneous.
*fz46f discoverer. Bridges, 22d22.
origin: Spontaneous. references: Morgan, Bridges, and Sturtevant, 1925,
discoverer: Ives, 46120. Bibliog. Genet. 2: 226.
references: 1946, DIS 20: 65. phenotype: Like g, but gives lighter orange in
phenotype: Like fit. RK2. combination with v. Has 21 percent normal red
pigment and 47 percent normal brown pigment
9; qornmt (Nolte, 1959, Heredity 13: 233-41). Malpighian
location: 1-44.4. tubes extremely pale yellow (Brehme and Demerge,
origin: Spontaneous. 1942, Growth 6: 351-56). RK1.
MUTATIONS 103
other information: Occupies third pseudoallelic site g53d
from left of g locus (Chovnick, 1961, Genetics 46: origin: Spontaneous.

493-507). discoverer: Hexter, 53d.
94 references: 1958, Proc. Natl. Acad. Sci. U.S. 44:
origin: X ray induced in In(l)dl-49. 768-71.
discoverer: Glass, 1929. 1956, DIS 30: 72.
references: 1934, DIS 2: 7. 1963, Proc. Natl. Acad. Sci. U.S. 50: 372-79.
phenotype: Eye color, like g2, has 22 percent phenotype: Eye color orange, like wB. RK1.
normal red pigment and 23 percent normal brown other information: £53d, when heterozygous with g,
pigment (Nolte, 1959, Heredity 13: 233-41). g2, g3, or g4, yields wild-type progeny, some
Malpighian tubes extremely pale yellow (Brehme associated with crossing over in a manner con-
and Demerec, 1942, Growth 6: 351-56). RK1. sistent with ordering, and some not associated
cytology: Probably inseparable from In(l)dl-49. with recombination 6f outside markers. A double
mutant is not produced by the event that yields
g17Ba6 wild types.
origin: X ray induced in In(l)scslL'sc8ridl-49. *aSSk
9
discoverer: Muller, Valencia, and Valencia, 1946-53. origin: Spontaneous.
references: Valencia, 1966, DIS 41: 58. discoverer: Williams, 55k.
phenotype: Male viable but sterile. RK1A. references: Muller, 1956, DIS 30: 80.
cytology: Associated with In(l)gx 7Bfl<* = In(l)12B14- phenotype: Eyes translucent yellowish ruby. RK1.
15;19F (J. I. Valencia). *a271-2
*g26-10 origin: X ray induced,
origin: X ray induced. discoverer: Demerec, 1933.
discoverer: Sobels, 57j. phenotype: Male lethal, cell lethal. RK2.
references: 1958, DIS 32: 84. other information: ty but not s, wy, or pi also
phenotype: Eye color darker than g2. RK1. affected.
g26-75 *a271-6
origin: Induced by mustard gas. origin: X ray induced.
discoverer: Sobels, 57j. discoverer: Demerec, 34a.
references: 1958, DIS 32: 84. phenotype: Male lethal and cell lethal. RK2.
phenotype: Eye color like g2. RK1. other information: ty but not s or wy also affected.
*g26-41 *g27J-9
origin: Induced by mustard gas. origin: X ray induced,
discoverer: Sobels, 57j. discoverer. Hoover, 35h.
references: 1958, DIS 32: 84. phenotype: Male lethal. RK2.
phenotype: Weak garnet allele. Eye color strikingly other information: ty+.
different from wild type in newly emerged flies. *g271-T0
RK2. origin: X ray induced.
*g28-10 discoverer: Hoover, 35h.
origin: Induced by mustard gas. phenotype: Lethal. RK2.
discoverer Sobels and Jansen, 1957. other information: ty+, s + .
references: Sobels, 1958, DIS 32: 84. g(+): s e e g™
phenotype: Eye color like g2 but darkens more with
age. RK1. origin: Appeared among progeny of cold-treated
g28-40 flies.
origin: Induced by mustard gas. discoverer Gottschewski, 34gl7.
discoverer Sobels, 1957. references: 1935, DIS 4: 8, 15.
references: 1958, DIS 32: 84. phenotype: Eye color of males like g2, of females
phenotype: Eye color darker than g2. RK1. probably lighter than g*. RK1.
g49h 9**: garnet of Schalet
origin: Induced by P 3 2 . origin: Spontaneous,
discoverer: R. C. King, 49h. discoverer: Schalet.
references: 1950, DIS 24: 58. references: Chovnick, 1961, Genetics 46: 493—507.
phenotype: Eye color light purple. Viability of male other information: Occupies right-most pseudoallelic
normal, female 60 percent normal. RK2. site of the g locus.
gSOm gw: garnet-wild
references: Chovnick, 1958, Proc. Natl. Acad. Sci. origin: X ray induced,
U.S. 44: 333-37. discoverer: Muller.
1961, Genetics 46: 493-507. synonym: |f( + ).
phenotype: Eye color orange. RK1. references: 1946, DIS 20: 67.
other information: Occupies second pseudoalleiic Chovnick, 1958, DK 32: 88.
site from left of g locus. 1961, Genetics 46: 493-507.
104
phenotype: Homozygote and hemizygote indistin- phenotype: Does not show G phenotype. Allelic to
guishable from wild type; however, gw/g2 (and ft, but does not exaggerate ft. Lethal in combina-
probably other g alleles) is brownish. RK2A. tion with G. RK2.
cytology: Associated with In(l)gw (breakpoints un- g, Inh: see g*
known), which lies between dy and f. gap: see gp
garnet: see g
gx: garnet from X irradiation *Gd: Gulloid
origin: X ray induced. location: 3-78.
discoverer: Muller. origin: Spontaneous in Dp(2;3)P.
synonym: g, Inh. discoverer: Bridges, 22g26.
references: 1946, SIS 20: 67. phenotype: Gd/+ wings shorter, blunter, slightly
phenotype: Like g2. Male fertile; homozygous fe- more spread, and have crossveins closer together
male has low fertility. RK2A. than wild type. Homozygous lethal. RK3A.
cytology: Associated with /n(i;g* = In(l)12;19-20. cytology: Inseparable from Dp(2;3)P =
Dp(2;3)58E3-F2;60D14-E2; 96 B5- Cl.
*ge: genitalless
location: 1-0.1.
origin: Induced by methyl methanesulfonate
(CB. 1540).
phenotype: External male genitalia absent or
grossly deformed. Bristles fine; wings often small
and deformed. Tergites abnormal; abdomen
frequently contains melanotic tumors. Males viable
but sterile. RK3.
gespleten: see gs
*99: goggle
location: 1-23.1 (no crossovers with oc among 4300
flies).
G: Gull origin: Spontaneous.
From Mohr, 1929, Z. Induktive Abstammungs- discoverer: Nichols-Skoog, 34el4.
Vererbimgslehre 50: 113-200. phenotype: Eyes protruding and bulging; placed far
back on a narrow head. Facets very large in rough
G: Gull areas. Wings smaller, with fringed marginal hairs;
location: 2-12.0. dusky; pebbly appearance caused by large cells.
origin: Spontaneous. Bristles coarse and irregular; hairs sparse and
discoverer: Mohr, 19k23. irregular, especially on abdomen. Body small in
references: 1923, Studia Mendeliana (Brunae), late counts. Viability 20 percent wild type.
pp. 266-87 (fig.). Females usually sterile; males usually fertile.
1927, Proc. Intern. Congr. Genet., 5th. Vol. 2: RK3.
1136. gg2
1929, Z. Induktive Abstammungs- Vererbungslehre discoverer: Waletsky, 371.
501 113-200 (fig.). phenotype: Like gg, but many bristles on posterior
pti*nofyp«: Wings Urge, held out from sides at 45— lateral margin of head missing; others on head and
90°«gl«!, cttnred downward, and somewhat pointed. thorax reduced or missing. Females sterile; at
Vein LI thickened; crosaveins closer together, 19 C, ovaries small and contain no oocytes; at
s©«teti»es broken. Thoracic and vertical bristles 25°C ovaries contain a few eggs, but none are laid
tfoplicated in majority of flies. G/ft has exagger- (Beatty, 1949, Proc. Roy. Soc. Edinburgh, B 63:
ated It pfamotypt. Partially inhibited by d«/+ and 249-70). RK3.
ja»ei» inhibited by dc/rfc. Homozygous lethal. RK2. 99 3
cy*@J®fy: Placed between 24D2 und 24F1 on the origin: Spontaneous.
b*fti* of its inclasioo in Dt(2L,yt~zc = Df(2L)24D2- discoverer R. F. Grell, S3d.
&2SA2-3 bat »ot fa Df2L>f-x« *D§(2L)24E2~ references: 1953, DIS 27: 59.
Pli25Al'2 {**!»&„ Bridges, Scfaults, and Curry, phenotype: Similar to gg, but both sexes fertile.
1939, Otraegic last. Wash. Y®ar Book 38: 273-77). Viability low. RK2.
®f§*«? tafor»#tie«: Causes local shortening of map by
99*
abet* 1.1 units. I* a d^fk-iesicy for or an allel* of
it.
discoverer: Mohler, 54J28.
!te » of 0.
phenotype: Similar to gg3. Varies from eyelessness
in first flies to emerge to nearly normal in aging
MUTATIONS 105
cultures. Always s e p a r a b l e from wild type on phenotype: Like gl3, perhaps with eye color more
basis of p e b b l y a p p e a r a n c e of wings. RK2. normal. RK1.
giant: see gt
Giant: see Gt location: 3-64.
giantoid: see gtd origin: Spontaneous.
gl: glass discoverer: Oliver, 41el.
location: 3-63.1. synonym: gl-l: glass-like.
discoverer: Muller, 18b. phenotype: Eye texture smooth to rough. Eye
references: Bridges and Morgan, 1923, Carnegie smaller than wild type. Eye color orange; lighter
Inst. Wash. Publ. No. 327: 188 (fig.). around rim of eye. Viability and fertility good.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. RK2.
Genet. 2: 214 (fig.), 226. *glS1k
phenotype: Eyes reduced to one-half normal area; origin: Spontaneous.
diamond shaped with glassy texture from fused discoverer Edmondson, 51k.
facets and irregular surface. Eye color blotchy, references: 1952, DIS 26: 60.
ranging from scarlet to orange in males and orange phenotype: Like gl. RK2.
to apricot in females; colorless rim and eroded *glS4g
patches, especially in female. Malpighian tubes of origin: Spontaneous.
larva paler than wild type (Brehme). RK2. discoverer: Hexter, 54g.
cytology: Located in region between 90C2 and 91A3 references: 1956, DIS 30: 72.
on basis of its inclusion in Df(3R)P14 = phenotype: More extreme than gl. Facet area less
Df(3R)90C2-Dl;91A2-3 (E. B. Lewis). than one-half of normal surface; very irregular.
gl2 Eye color white with some orange specks in both
origin: Spontaneous. sexes. Ocelli colorless. Malpighian tubes normal.
discoverer: R. L. King, 1927. RK1.
phenotype: Slightly less extreme than gl. Eyes gi62d
reduced to two-thirds normal size; ovoid; glassier origin: Spontaneous.
and smoother than gl. Eye color blotchy scarlet in discoverer: Tano, 62d.
both sexes, with rim and eroded patches of color- references: Burdick, 1963, DIS 37: 47.
less material. Ocelli colorless and papillose. phenotype: Less extreme than gl2. Viability normal.
Larval Malphigian tubes normal yellow (Brehme). RK1.
Cephalic complex smaller than wild type at 36 hr *gl63a14
after hatching (27°C); growth rate subsequently origin: Spontaneous.
normal [Medvedev, 1935, Z. Induktive discoverer: Ashburner and Hudson, 63al4.
Abstammungs- Vererbungslehre 70: 55—72 (fig.)J references: 1966, DIS 41: 60.
1935, Tr. Inst. Genet. Akad. Nauk SSSR 10: 119- phenotype: Similar to gl, but eye color darker and
5l]. RK2. pigmented area larger. Male eyes darker than fe-
9I3 male. Malpighian tubules wild type, gl63*14/gl
origin: Spontaneous. has more eye pigment than either homozygote. Via-
discoverer: Stern. bility and fertility good. RK1.
synonym: rh: rauhig. gl63d
references: Csik, 1929, Biol. Zentr. 49: 419—21. origin: Gamma ray induced.
phenotype: Eye small and elliptical, with surface discoverer: Ives, 63d29.
less rough than gl or gl2. Eye color homogeneous references: 1965, DIS 40: 35.
scarlet in both sexes. Ocelli colorless. Viability phenotype: Eyes small, nearly colorless, with a
and fertility good. Malpighian tubes of larvae wild typically glassy surface. Viability of homozygote
type (Brehme). RK1. good except in competition with other genotypes.
*gt4 RK2A.
origin: Spontaneous. cytology: Associated with T(2;3)gl63d; breakpoints
discoverer: Villee, 40d. unknown.
references: 1941, DIS 14: 40. *qJ63f6
1942, Univ. Calif. (Berkeley) Publ. Zool. 49: 137. origin: Spontaneous.
phenotype: More extreme than gl. Eyes reduced to discoverer: Ashburner and Hudson, 63f6.
less than one-half normal area; ovoid; narrower, references: 1966, DIS 41: 60.
glassier, and smoother than gl. Eye color white to phenotype: Like gl63t**4 and may be identical. RK1.
apricot in females, pink in males. Ocelli colorless. Gl: Glued
Viability about 65 percent wild type. RK2. location: 3-41.4 [0.9 unit from Ly (Mossige, 1935,
*gl40h DIS 4: 59; 1938, Hereditas 24: 110-16)1.
origin: Spontaneous. origin: Recovered among progeny of heat-treated
discoverer: Ives, 4Qh. flies,
references: 1941, DIS 14: 39. discoverer: Ives, 31 f5.
references: 1934, DIS 1: 34. references: Eyes rough but of normal size; facets
1934, DIS 2: 35. and hairs irregular. RK2.
Plough and Ives, 1935, Genetics 20: 42—69 (fig.) Go; Gold tip
phenotype: Eyes rough, smaller, and oblong; facets location: 2-64.3 (57.5 to 71.1; between en and eg).
rounded; surface smooth and shiny like gl. origin: Spontaneous.
Bristles generally shortened slightly and straighter discoverer: Sturtevant, 1948.
than normal. Viability and fertility good. references: 1948, DIS 22: 55.
Homozygous lethal. RK1. phenotype: Tips of many bristles and hairs pale and
cytology: Salivary chromosomes normal (Bridges in curved. Bristles often short (tips broken off ?).
Morgan, Bridges, and Schultz, 1937, Carnegie Inst. Wild-type bristles sometimes have pale tips, thus
Wash. Year Book 36: 301). interfering with positive classification. Lethal
gl-1: see gl*i« when homozygous. Expression best at low temper-
atures. RK2.
Gla: Glazed goggle: see gg
location: 2- (rearrangement). Gold tip: see Go
references: Morgan, Bridges, and Schultz, 1936, gouty legs: see gy
Carnegie Inst. Wash. Year Book 35: 293.
phenotype: Eye reduced to one-fourth normal area
and narrowed to a point ventrally. Eye color gen-
erally diluted but with some black patches. Omma-
tidia coalesce into gleaming, smooth sheet. Mal-
pighian tubes of larva somewhat lighter than wild
type; difficult to classify (Brehme and Demerec,
1942, Growth 6: 351—56). Homozygous lethal.
RK2A.
cytology: Associated with In(2LR)Gla =
In(2LR)27D;51E, superimposed on In(2L)Cy =
ln(2L)22Dl-2;33F5-34Al or In(2L)t = gp- gap
In(2L)22D3-El;34A8-9. From Bridges and Morgan, 1919, Carnegie Inst. Wash.
glass: see gl Publ. No. 278: 209.
glass-like: see gl41 e
Glazed: see Gla gp; gap
gleam: see gm location: 2-74.
*gli: glide origin: Spontaneous.
location: 1-38.0. discoverer: Bridges, 12alO.
origin: Induced by DL~p-NN-di-(2-chlorethyl)amino- references: Bridges and Morgan, 1919, Carnegie
phenylalanine (CB. 3007). Inst. Wash. Publ. No. 278: 208 (fig.).
discoverer: Fahmy, 1954. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
references: 1958, DIS 32: 70. Genet. 2: 212 (fig.), 226.
phenotype: Wings held horizontally at right angles phenotype: Vein L4 weak or has section missing
to body. Pigmentation of tergites frequently inter- beyond posterior crossvein. Overlaps wild type
rupted along mid-dorsal line; tergites occasionally when homozygous; semidominant as heterozygote.
show a nick in the posterior border. Males sterile; RK3.
viability about 70 percent wild type. RK2. *gr: gracile
glisten: see gn location: 1-36.4.
glossy: see Iz origin: Induced by L-p-NN-di-2-chloroethyl)amino-
Glued: see Gl phenylalanine (CB. 3025).
gly: see Iz6 discoverer: Fahmy, 1953.
gm: gleam references: 1959, DIS 33: 86.
location: 3- (not located), phenotype: Small fly with narrow abdomen. Wings
origin: Spontaneous, frequently held atypically, either upward or down-
discoverer Bridges, 27cl. ward. Very inviable, many dying less than 24 hr
phenotype: Eyes small and rough; irregular hairs after eclosion; males sterile. RK3.
and facets cause glints. Body small. Viability *gre: green body color
about 10 percent wild type but variable. RK3. location: 1- (not located).
cytology: Associated with In(3L)P, according to origin: Spontaneous.
Bridges (Morgan, Bridges, and Schultz, 1937, discoverer: Bridges, 13e.
Carnegie Inst. Wash. Year Book 36: 301). references: Morgan and Bridges, 1916, Carnegie
*gm: glisten Inst. Wash. Publ. No. 237: 73.
Socotlon: 3-67.3. phenotype: Body color tinged greenish black, with
origin: Gamsaa my induced. marked trident pattern. Overlaps wild type. RK3.
r. WcUbrun, 61i6.
other Information: Possibly an allele of ptg.
MUTATIONS 107
gro: groucho gland chromosomes of double thickness in some

location: 3-90 (no recombinants with Pr obtained). cells (Bridges, 1935, J. Heredity 26: 60-64). RK3.
origin: Spontaneous. other information: Used by Bridges (1935) in the
discoverer: E. H. Grell, 64k. construction of salivary chromosome maps.
phenotype: Clumps of extra bristles above each eye *Gt-2: Giant in chromosome 2
give impression of bushy eyebrows; also extra location: 2- (not located),
bristles on humerus. Top of head tends to be mal- origin: Spontaneous,
formed; ocelli often enlarged and run together. In discoverer: Bridges, 14128.
selected stocks, penetrance approaches 100 per- phenotype: Heterozygote normal but, in presence of
cent but is low in unselected stocks. Probably no homozygous gt-3, gives giant male-sterile flies.
gross chromosomal rearrangement since recombina- Homozygous lethal. RK3.
tion is normal in vicinity of mutant. RK2. *gt-3: giant in chromosome 3
location: 3-64.
grooved: see gv origin: Spontaneous.
groove/ess: see gvl discoverer: Bridges, 14i28.
groucho: see gro references: Bridges and Morgan, 1923, Carnegie
gs: gespleten phenotype: Body size much larger than normal. Late
location: 3-35.1. hatching. Entirely sterile in male. Giant character
origin: Spontaneous. produced only in flies homozygous for gt-3 and
discoverer: Smelink-den-Hollander, 561. heterozygous for Gt-2. RK3.
references: 1957, DIS 31: 85. gt-4
phenotype: Thorax cleft by medial groove. Eyes location: 2-24.0.
small; sometimes missing. Hair pattern on thorax origin: Spontaneous,
disturbed. Viability and fertility excellent. RK1. discoverer Bridges, 30bl4.
other information: Probably an allele of gv (3-36.2). phenotype: Giant flies hatch very late. Viability
variable but around 15 percent wild type. RK3.
*gtd: giantoid
location: 1-0.5.
discoverer: Bridges, 21cl2.
references: Bridges and Gabritchevsky, 1928, Z.
Induktive Abstammungs- Vererbungslehre 46: 232
(fig.).
phenotype: Body size larger, especially head. Late
hatching. Viability erratic, about 50 percent wild
type. Separation difficult in females, easier in
males. RK3.
Gull: see G
Gulloid: see Gd
*gv: grooved
gt: gianf location: 3-36.2.
Left: wild-type female. Right: giant female. origin: Spontaneous,
From Bridges and Gabritschevsky, 1928, Z. Induktive discoverer Ives, 43128.
phenotype: A longitudinal medial groove in thorax;
in extreme individuals, thorax nearly cleft. Eyes
gt: giant
reduced. Irregular and often extra alar bristles.
location: 1-0.9.
Viability good. RK1.
other information: Probably allelic to &*.
discoverer: Gabritschevsky, 25i2.
references: Bridges and Gahritschevsky, 1928, Z.
Induktive Abstammungs- Vererbungslehre 49: gvl: groove/ess
location: 4-0.2 fin diplo-4 triploids (Sturtevant,
231-47 (fig.).
1951, Proc. Natl. Acad. Sci. U.S. 37: 405-7)].
Gabritschevsky and Bridges, 1928, Z. Induktive
discoverer: Bridges, 33elO.
phenotype: Larval development 4 days longer than
references: 1935, Biol. Zh. (Moscow) 4: 401-20.
normal, resulting in giant larvae, pupae, and
phenotype: Sharp transverse groove between scu-
imagos. Adult weight 1.7 times normal. But not all
tellum and thorax is nearly eliminated; no overlap
genetically giant flies show the character; the rest
of wild type. Black scars appear on scutellar
have normal size. Distribution sharply bimodal.
groove at si«4e&, in pleural region, and behind
Percentage giant greatest in well-fed cultures;
stemopletirais. Viable and fertile. RK1.
also raised by modifying action of bh11. Salivary
gy: gouty legs (Stern, 1944, DIS 18: 56). Shown byboth Sturtevant
location: 4- (not located). and Rasmussen to recombine with and He to the
origin: Spontaneous. right of h2.
discoverer: Muller. h2
phenotype: Legs shortened and thickened, espe- discoverer: Bridges, 28d23.
cially the metatarsi of the hind legs, which are phenotype: Extra hairs on wings, but fewer than in
often swollen. Usually classifiable; viability and h. Extra hairs not present on scutellum, pleurae,
fertility good. gy/eyD is gy. RK2. or top of head. Bristles cylindrical, with javelin
heads. Some abnormal abdomen effect. RK2.
other information: Shown by both Sturtevant and
Rasmussen to recombine with and lie to the left of
h.
*h!00.12
discoverer: Alexander.
references: Ward and Alexander, 1957, Genetics 42:
42-54.
phenotype: Homozygous viable. RK2A.
cytology: Associated with In(3L)h100-12 =
In(3L)61A2'3;66D.
*/,700.239
references: Ward and Alexander, 1957, Genetics 42:
42-54.
phenotype: Homozygous semilethal. RK2A.
cytology: Associated with In(3L)h100'239 =
In(3L)66Dll-12;80C.
*hl00.271
h: hairy origin: X ray induced.
From Bridges and Morgan, 1923, Carnegie Inst. Wash. discoverer: Alexander,
PubJ. No. 327: 202. references: Ward and Alexander, 1957, Genetics 42:
42-54.
h: hairy phenotype: Homozygous lethal. RK2A.
location: 3-26.5. cytology: Associated with T(2;3)ht00.271 =
origin: Spontaneous. T(2;3)41;66D14'E1.
discoverer. Mohr, 18111. */,7OO.39O
references: 1922, Z. Induktive Abstammungs- origin: X ray induced.
Vererbungslehre 28: 17. discoverer: Alexander.
Bridges and Morgan, 1923, Carnegie Inst. Wash. references: Ward and Alexander, 1957, Genetics 42:
Publ. No. 327: 202 (fig.). 42-54.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. phenotype: Homozygous lethal. RK2A.
(tenet. 2: 214 (fig.), 227, cytology: Associated with Df(3L)h100'390 =
Neel, 1941, Genetics 26: 52-68 (fig.), Di(3L)66D2'5;66D14-El.
pfoenotype: Extra hairs cm scutellum, along veins, ft*; hairy-subliminal
on pleorae, and on top of head. Additional hair- origin: X ray induced.
forming cells present in 19-hr pupa (Lees and discoverer: Green.
Waddiagtoa, 1942, DIS 16: 70). Interacts synergis- references: 1960, Proc. Natl. Acad. ScL U.S. 46:
tically with ppd and Hw or Hw variegateds [e.g., 524-28.
JTnf/)*e*]; In{l)sc7 suppresses h (Steinberg, 1942, phenotype: Homozygote nearly lethal but has no h
DIS 16: 68; Neel, 1941). RK1. phenotype. Heterozygote with h and h2 also wild
ey*of*§y: Pla-eed is aalivsry chromosome region be- type. he/+ has extra hairs on wings, head,
tween 66D2 and 66E1, on basis of its inclusion in pleurae, halteres, and occasionally on scutellum if
DK3LftteO.3f0 mDI(3L)6$D2-5;66DI4'E1 (Ward also heterozygous for certain X-chromosome inver-
•ad AUxmndm, 1957, Genetics 42: 42-54). sions that variegate for Hw, including In(l)scs,
e*h«r in§ormotion: As with c/ + , expression of h + may In(l)acS*, and ln(l)y3P. Presence of y+Y also in-
b® altered i© tht direction of ft by rearrangements duces extra hairs, RK3.
with bmstkm in the vicinity of the h locus [Dubinin H: Hairless
@«j Sldorov, 1934, Biol. Zh. (Moscow) 3: 307-31 ]. location: 3-69.5.
Unlike tike ci case, however, rearranged ft chrotno* origin: Spontaneous.
do act sfeow evidence of altered gene action discoverer: Bridges, 16c4.
MUTATIONS 109
references: Bridges and Morgan, 1923, Carnegie references: Plunkett, 1926, J. Exptl. Zool. 46:
Inst. Wash. Publ. No. 327: 161 (fig.). 181-244.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. phenotype: Bristle effect more extreme than in H
Genet. 2: 170 (fig.), 227. and more easily separated from wild type. Venation
Nash, 1965, Genet. Res. 6: 175-89. effect slighter than in H. Interactions with mutants
phenotype: Bristles, especially postverticals and at the N locus similar to those of H (Von Halle).
abdominals, missing. Bristle sockets present at Homozygous lethal. H2/H lethal. RK1.
some sites, not at others. Veins L4 and L5 do not H3
reach wing margin; occasionally true of L2 also. discoverer: Sturtevant.
Eyes larger than wild type; body color somewhat phenotype: Like H. RK1.
paler. Lees and Waddington [l942, Proc. Roy. Soc. 4
(London), Ser. B. 131: 87-110 (fig.)J show that origin: Spontaneous.
trichogen cell forms a socket instead of a bristle at discoverer: Bridges, 30b20.
some sites. Interactions with other mutants studied phenotype: Like H. RK1.
by House (1953, Genetics 38: 199-215, 309-27;
1959, Genetics 44: 516; 1955, Anat. Record 122: origin: Gamma ray induced.
471; 1959, Anat. Record 134: 581-82). H sup- discoverer: Ives.
presses wing notching of /V, fa, fano, and nd; en- references: 1959, DIS 33: 95.
hances Ax, and enhances eye effect of spl; H re- phenotype: Extreme bristle effects; anterior lateral
moves more bristles in combination with spl acrostichals removed. L5 incomplete distally.
(House, Von Halle). Shows some superadditive in- RK2A.
teraction with en, ci, ciw, and ci® relative to de- cytology: Shown genetically to be associated with
gree of L4 interruption. L2 interruption augmented
in combinations with ve and ri; L3 interruption aug- other information: Allelism to H inferred from
mented in combinations with ve and tt. Triploid, phenotype.
H/+/+, intermediate between wild type and H/+. Hairless of Dobzhansky
H/H/+ most extreme type, with bristles absent origin: X ray induced,
from head, thorax, and abdomen LGowen, 1933, Am. discoverer: Dobzhansky, 1930.
Naturalist 67: 178—80 (fig.)J. Homozygous lethal. phenotype: Slight allele of H with no shortening of
RK1. L4 or L5. Bristle effect slighter, particularly on
cytology: Salivary chromosomes normal. abdomen. Homozygous lethal. RK2.
*HP2: Hairless from P32

origin: Found among descendants of male fed P 3 2 .
discoverer: Bateman, 1949.
phenotype: Bristle effect like H, but venation quite
distinct. Veins not interrupted, but knotted. Homo-
zygous lethal. RK1.
*ha: hair bristles
location: 1-22.7.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
discoverer. Fahmy, 1954.
phenotype: Small fly with extremely fine, short
bristles. Males viable and fertile. Females less
viable and highly infertile. RK3.
Hairless: see H
hairy: see h
Hairy wing: see Ww
Haltere mimic: see Hm
hdp: holdup
location: 1-59.5.
origin: Induced by DL-p-NN-di-{2-chloroethyl)amino-
H: Hairless phenylalanine (CB. 3007).
From Bridges and Morgan, 1923, Carnegie Snst. Wash. discoverer: Fahmy, 1954.
Publ. No. 327: 161. references: 1958, DIS 32: 70.
phenotype: Wings held up to various degrees. May
overlap wild type. Viability and fertility good in
origin: Spontaneous. both sexes. RK2.
discoverer: Stwrtevant. other information; One X-ray-induced allele.
no GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER
*/i</prwg; heldup-reduced wings roughened. Wings usually divergent and may be

origin: Induced by 2-chloroethyl methanesulfonate smaller. Body sometimes small and chunky. Less
(CB. 1506). extreme expression at 19°C, especially the wing
discoverer: Fahmy, 1956. character, but classification reliable. RK2.
synonym: rwg. cytology: Salivary chromosome locus between 37B2
references: 1958, DIS 32: 74. and 40B2 (Schultz and Curry).
phenotype: Wings short; upheld or outspread to
various degrees. Small regions of deranged wing
hairs, giving shaded streaks parallel to longitu-
dinal veins. Male infertile; viability about 70 per-
cent normal. RK2.
heavy vein: see hv
holdout', see ho
heldup: see hdp
Henna: see Hn
Hermaphrodite: see traD
*hi: high
origin: Found in Florida natural population.
discoverer: Ives, 1943.
1950, Evolution 4: 236-52.
phenotype: Male homozygous for hi produces sperm
containing 10 times normal frequency of mutations.
Heterozygous hi/+ causes a mutation rate 2—7
times normal. Ratio of sex-linked lethal to visible
mutations about 8 to 1. Inversions associated with
about 5 percent of mutations. RK3.
other information: Homozygous hi constructed by hk: hook
crossing two balanced lethal stocks, 11 M/CyX From Mohr, 1927, Hereditas 9: 169-79.
12 hi/Cy. Since these stocks have developed a hk2
common lethal, it is now difficult to obtain hi origin: Spontaneous.
homozygotes. discoverer: Bridges, 33a31.
*Hi: Hirsute phenotype: Bristles cylindrical, javelinlike, or bent,
location: 3- (rearrangement). but effect less extreme than hk. RK2.
origin: X ray induced. Hm: Haltere mimic
discoverer: Bishop, 1939. location: 2- or 3- (rearrangement).
phenotype: All bristles except postscutellars and origin: X ray induced.
postdorsocentrals multiplied, especially on head discoverer Slatis, 49b5.
and anterior thorax. Eyes smaller and facets phenotype: Wing resembles a large haltere, but stalk
irregular. Homozygous lethal. RK2A. is more winglike. In combination with vg, wing is
cytology: Associated with In(3LR)Hi - further reduced and haltere almost completely
In(3LR)71A;91F. eliminated. Homozygous lethal. RK3A.
Hia: Hiatus cytology: Appears to be associated with complex
location: 2- (not located), translocation, T(2;3)Hm.
origin: Spontaneous. tin31: s e e UnT<*
discoverer: Bridges, 2%12. Hn; Henna
phenotype: Terminal interruption of L2. More location: 3-23.0.
obvious in heterozygous male than in heterozygous origin: X ray induced.
female. Homozygous viable. RK3. discoverer: Van Atta, 30k.
high: see hi references: 1932, Am. Naturalist 66: 93—95.
Hirsute: see Hi 1932, Genetics 17: 637-59.
hk: hook phenotype: Eye color homogeneous dark, dull brown.
location: 2-53.9. Classification difficult; best in aged flies. Homo-
origin: Spontaneous. zygous lethal. Heterozygotes viable and fertile.
discoverer; Mohr, 24a4. RK2A.
references: 1927, Hereditas 9: 169-79 (fig.)- cytology: Placed in 66A-B, on basis of its associa-
pltttnotyp*: Bristles nearly ail hooked at tip or tion with Df(3L)Hn = Df(3L)66A;66B (Lewis, 1956,
blunted; some bent at right angles. Scute liars and DIS 30: 130). Hn was induced together with, but is
verticals especially affected. Acrostichal hairs separable from T(2;3)Hn = T(2;3)53E-
fewer mad outer rows separated. Eyes slightly 54A;77A;94F;96A.
MUTATIONS in
*HnS3k references: Novitski and Rifenburgh, 1938, Proc.
origin: Ultraviolet induced. Indiana Acad. Sci. 47: 256-60.
discoverer: Meyer and Verderosa, 53k. phenotype: Wings extended at right angles to body.
references: Meyer, 1954, DIS 28: 76. RK1.
phenotype: Heterozygote has brick-red eye color, cytology: Located in or near 22E (Lewis, 1945,
but classification difficult. Homozygote viable; Genetics 30: 137-66).
eye color dark brown like sepia. RK1 as recessive.
Hnr: Henna-recessive origin: X ray induced.
origin: Spontaneous. discoverer: E. B. Lewis, 1940.
discoverer. Bridges, 33c20. synonym: In-ho.
references: Mohr, 1937, DIS 8: 12. references: 1945, Genetics 30: 137-66.
phenotype: Eye color dull, dark brown, like cl. Best phenotype: Wings reduced to tiny stubs. Eyes
separability in aged flies. Hnr/Hn more extreme reduced, with anterior indentation. Male lacks
than Hnr/Hnr. Eye color autonomous in transplant genitalia and anal apparatus; female fertile.
into wild-type larval host (Beadle and Ephrussi, hoÔ/ho resembles ho/ho. RK2A.
1936, Genetics 21: 230). Larval Malpighian tubes cytology: Associated with In(2L)ho'*0 =
bright yellow as in wild type (Beadle, 1937, In(2L)21D4-El;22E2-3.
Genetics 22: 587—611). Eyes have 58 percent hook: see hk
normal red pigment and 120 percent normal brown Hooked veins: see Hv
pigment (Nolte, 1954, J. Genet. 52: 127-39). RK2. hp: humped
location: 3- (rearrangement).
discoverer: Nordenskiold, 39b9. origin: Spontaneous.
synonym: bu; brunette; hn^. discoverer: Bridges, 31a22.
references: 1937, DIS 7: 18. phenotype: Thorax shortened and strongly humped,
phenotype: Eye color darker than HnT in young flies. with thoracoscutellar groove almost absent. Eyes
RK2. sharply reduced; may be absent at 19°C. Bristles
Hnr3 Minutelike and occasionally missing. Viability 10
origin: Spontaneous. percent wild type. RK3A.
discoverer: Weinstein, 1927. cytology: Associated with In(3R)hp.
synonym: sed: sepiaoid. *hpa: hyperantenna
phenotype: Eye color dull chocolate. Classification location: 1-50.1.
easier than for Hnr. 79 percent normal red pigment, origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
100 percent normal brown pigment (Nolte, 1955, J. phenylalanine (CB. 3007).
Genet. 53: 1—10). Eye color autonomous in trans- discoverer: Fahmy, 1954.
plants of optic disk into wild-type hosts (Beadle references: 1959, DIS 33: 86.
and Ephrussi, 1936, Genetics 21: 230). Larval phenotype: Antennae enlarged or have duplicated
Malpighian tubes bright yellow as in wild type parts, sometimes an extra antennal base near the
(Beadle, 1937, Genetics 22: 587-611). RK2. eye. Grossly deformed head and eyes. Wings have
other information: Allelism by Lewis (1956, DIS 30: rounded tips and incised inner margins. An occa-
130). sional bristle absent or shorter. Phenotype vari-
*Hn'53/ able and minimal expression slightly altered eye
origin: Ultraviolet induced. shape and blunt wing tips. Males viable and
discoverer: Meyer, 53j. infertile; females sterile. RK3.
references: 1954, DIS 28: 76. Ht: see traD
phenotype: Similar to se. RK1. Hu: Humeral
*Hnrh Henna-recessive of Ives location: 3-51 (48-54).
origin: Spontaneous. origin: X ray induced.
discoverer: Ives, 45J17. discoverer: Ruch, 1931.
references: 1946, DIS 20: 65. phenotype: Extra bristles on humeral patches of
phenotype: Eye color brownish at hatching, darkens Hu/+. Humeral bristles more numerous in homozy-
to black. Resembles se. Viability good. RK1. gote, with a streak of hairs below humerus toward
*HnrN: Henna-recessive from Nebraska base of first leg. Hu/Hu has viability 90 percent
origin: Spontaneous. wild type. RK2A as heterozygote.
discoverer: Williamson, 53j. cytology: Associated with In(3R)Hu = In(3R)84B2-
references: 1955, DIS 29: 75. 3;84F2-3;86B4-C1.
phenotype: Indistinguishable from se. Larval Mal- humped: see hp
pighian tubules somewhat darker yellow than wild humpy: see hy
type. RK1. hv: heavy vein
ho: heldout location: 2-104.0.
location: 2-4.0. discoverer: Curry, 36115.
origin: Spontaneous. phenotype: Veins thick and knotty, especially at
discoverer Novitski, 35g. ends of crossveins; posterior crossvein oblique
and may show break in middle; extra cross veins Hw*

sometimes present. Wings broad, thick, dark, origin: Spontaneous derivative
warped, divergent, and droopy. Eyes small and discoverer: Nichols-Skoog, 35a9.
bulging. Posterior scutellars blunt, short, and phenotype: Females homozygous for Hw2 show only
crossed. Overlaps wild type at 25°C but useful at occasional extra hairs along wings. Overlaps wild
19°C. RK2. type. RK3A.
*Hv: Hooked veins cytology: Salivary chromosome analysis by Schultz
location: 1-66. (Morgan, Schultz, and Curry, 1941, Carnegie Inst.
discoverer: Tanaka, 35a4. Wash. Year Book 40: 284) shows small inversion
references: 1935, DIS. 4: 16. of the region from 1A3 through 1B1 of the first
1936, D1S 5: 8. 1B1-2 doublet of Dp(l;l)Hw; i.e., associated with
1937, DIS 8: 11. In(l)Hw2 =In(l)lA2-3;lBl-2.
phenotype: Heterozygous female shows small Hw49c
branches from posterior crossvein and L5. Eyes origin: Induced by P32 simultaneously with sc45<=.
small and rough. Homozygous female lethal. discoverer: R. C. King, 49c21.
RK3A. references: Poulson and King, 1949, DIS 23: 62-63.
cytology: Associated with In(l)Hv. phenotype: More extreme than Hw. Homozygous
female has doubling and tripling of many bristles;
extra wing veins; gap in posterior crossvein; extra
hairs on vein L2 and in wing cells. Heterozygous
female has normal bristles but extra hairs on L2
and L3 and in wing cells; often an extra free vein
from posterior crossvein. Hw4^c male much like
homozygous female, but bristle duplication less
extreme. Male and heterozygous female fertile;
homozygous female sterile. RK1.
Hx: Hexaptera
discoverer: Herskowitz, 47j.
references: 1949, Genetics 34: 10-25 (fig.).
phenotype: Expression same in Hx/-f and Hx/Hx;
varies from absence of a detectable difference from
normal, through various intermediate types, to
presence of large appendage on prothorax. Entire
abnormal structure may remain beneath exoskeleton.
Appendage varies from small amorphous mass to
highly differentiated wing. May also produce
haltere- and leglike appendages. Penetrance same
Hw: Hairy wing in homozygote and heterozygote; enhanced by
Edith M. Wallace, unpublished. crowding and by high temperature (20°C, male 1.5
Hw: Hairy wing and female 3.3 percent; 25°, male 6.5 and female
location: 1-0.0. 24.2 percent); affected by genotype, e.g., sup-
origin: Spontaneous in stock containing y and never pressed by In(2L+2R)Cy and by ln(2LR)bwvl.
separated. RK3.
discover**; Bridges, 23cl2. hy; humpy
reference*: N«el, 1941, Genetics 26: 52-68 (fig.). location: 2-93.3.
pHenotyp*: M®ie has extra bristles along wing vein, origin: Spontaneous.
©a btftd (especially occipitals), and on thorax. discoverer: Bridges, 18j22.
Ala©, extra hairs on wing veins, back of head, and references: 1937, Cytologia (Tokyo), Fuji! Jub. Vol.
SMKopletvae. Homoxygous female sterile, expres- 2: 745-55.
sion men mximmm than male, and has 40—80 per- phenotype: Thorax strongly ridged, with commas
cent wiid-typ® viability. Heterozygous female has anteriorly and two pairs of vortices. Wings
good viability. Phenotype similar to male. Classi- obliquely truncated to one-half normal length. An
fiable an stagte do«# in triploids (Schultz, 1934, irregular contraction of larval muscles at time of
DB 1: 55). Interacts synergistically with ft and pupation (Waddington, 1941, Proc. Zool. Soc.
Pf4 (M«®!( 1941). Suppressed by su(Hw) and London Ser. A 111: 181-88). Viability low and
mu(Hw)2. REtA as heterozygous female and as erratic. Both sexes highly infertile. RK2.
cytology: Placed in region 57 on basis of its being
cyteleoy: Salivary chromosome analysis by Denserec to the right o(In(2R)NS =In(2R)52A2-Bl;56F9-13
cad Hoover shows repeat for doublet 1B1-2, i.e., and to the left of Df(2R)M'l = Di(2R)57Fll-
Dp(lilti»l\9*9, Gteetics 24: 68). 58A1;58F8-59A1 (Bridges, 1937).
MUTATIONS 113
apex. Eyes smaller and sometimes slightly rough.

Bristles thin. Males small, late eclosing; viability
reduced. Female sterile. RK3.
in: inturned
location: 3-47 (left of centromere).
discoverer: Bridges, 26k20.
phenotype: Hairs and bristles on thorax directed
irregularly toward midline. Marginal hairs of wing
stand out from wing margin; wings slightly spread
and tend to be long and narrow. RK1.
cytology: Tentatively placed in salivary region
77B-C (Hannah and*Arajarvi).
//?-/: see E(f)
In-ho: see ho40
*inb: incised balloon
location: 2-55.
hy: humpy origin: Spontaneous.
Edith M. Wallace, unpublished. discoverer: Neel, 41d9.
hyperantenna: see hpa phenotype: Wings held at 45° angle to body. Wing
margins incised, varying from slight nicks to
i: see E(B) extreme reduction to small fluid-filled sacs. RK2.
I: see E(B) cytology: Salivary chromosomes normal.
Ic D: see bwVD *lnd: Indented
*if: inflated location: 2-63.
location: 1-55. origin: Spontaneous.
origin: Spontaneous. discoverer Cole, 40e.
discoverer: Weinstein, 1916. references: Whittinghill and Parker, 1945, Genetics
references: 1918, Genetics 3: 157 (fig.). 30: 27-28.
phenotype: Wings inflated with lymph and smaller Whittinghill, 1947, DIS 21: 72.
than normal; venation defective. Wings later dry phenotype: Eye usually kidney shaped with indenta-
and blistered. Viability and fertility poor. RK3. tion anteriorly; shape sometimes normal, but facets
if* irregular. Often indented posteriorly as well as
origin: Spontaneous. anteriorly, sometimes dividing eye into two spots,
discoverer: Curry, 38b. or with only upper lobe persisting. Rarely eyeless.
references: 1939, DIS 12: 45. More extreme at 28° than at 25°C. RK2.
phenotype: Resembles if. Longitudinal veins inflated: see if
thickened, especially at wing base. Anterior infra bar: see B'
cross vein thickened. In most cases, wings inflated infrabar Bat: see BBi
or blistered. RK2. intensifies: see e( )
If: Irregular facets Intensifiet: see E( )
location: 2-107.6 (0.6 unit to the right of sp, ac- interrupted margin: see im
cording to Ives). Interruptus: see ciw
origin: Spontaneous. inter sex: see ix
discoverer: Casey, 65116. intersex on chromosome 3: see dsxôi
phenotype: In heterozygote, eye area about one-half intersex-62c: see dsx
of normal; narrow and pointed ventrally; facets ir- inturned: see in
regular and often missing across middle of eyes, Irregular facets: see If
sometimes fused or absent in ventral portion. In It: see ciw
homozygote, eyes are narrow slits, with smooth ix: intersex
glossy surface. Viability and fertility good. RK1. location: 2-60.5.
*im: interrupted margin origin: Spontaneous.
location: 1-3.1. discoverer: L. V. Morgan, 1943.
origin: Induced by 2-chloroethyl methanesulfonate references: Morgan, Redfield, and Morgan, 1943,
(CB. 1506). Carnegie Insst. Wash. Year Book 42: 171-74.
discoverer: Fahmy, 1956. Kroeger, 1959, Arch. Entwicklungsmech. Organ.
references: 1959, DIS 33: 86-87. 151: 301-22 (fig.).
phenotype: Wing margin nicked to various degrees, phenotype: Females changed into sterile inters exes
with costal vein frequently interrupted. Extra wing with a set of reduced male and a set of irregular
venation often present, and occasionally anasto- female external genitalia. Gonads also mixed.
moses, giving a plexus, particularly at the wing They have no sex combs; pigmentation of abdomen
intermediate between male and female. A large references: 1927, Biol. Zentr. 47: 361-69.
mass of chitinized tissue protrudes from vaginal 1934, DIS 1: 35.
opening. Males not affected. RK2. phenotype: Wings upturned but less extreme than ;.
iV Curl of wings dependent on hatching and unfolding
origin: Ultraviolet induced. at 25°C or higher. RK3.
discoverer: Meyer, 50k. *j49j
synonym: torn: tomboy. origin: Spontaneous.
references: Meyer and Edmonds on, 1951, DIS 25: 73. discoverer: Mossige, 49J18.
Meyer, 1958, DIS 32: 83. references: 1947, DIS 25: 69.
phenotype: Females homozygous for ix2 have male- phenotype: Less extreme than ;. Some overlapping
like pigmentation of posterior tergites, rudimentary at 21°, none at 30°C. Viability and fertility good.
ovaries, and are sterile. Expression extreme and RK2.
viability reduced at 27°C; at 17°C, expression iSO*
less extreme but viability greater. Homozygous origin: Spontaneous.
males appear normal but have nonmotile sperm. discoverer: Mossige, 5Oe5.
RK2. references: 1951, DIS 25: 69.
other information: The possibility that the male phenotype: Like j49i. RK2.
sterility is at another locus has not been excluded. *j58i
ix62c: s e e fax origin: Spontaneous.
ix-3: see discoverer: Andrew, 58i.
phenotype: Expression variable, although pene-
trance complete at temperatures above 25°C. In
most-extreme cases, wings bend sharply upward in
region of anterior crossvein. A small dark blot
occurs near L3 vein at level of the deflection.
Anterior crossvein partly or wholly absent in some
cases. RK2.
J: Jammed
location: 2-41.0.
discoverer: Bridges, 23d3.
phenotype: Wings often compressed into narrow
strips; sometimes filled with fluid. Alula larger
and square tipped, with clumped bristles and bare
regions. Alula modification is characteristic least
likely to overlap wild type. Completely overlaps
wild type at 19°, almost never at 28° or 30°C. Not
lethal when homozygous; viability, as in heterozy-
j: jaunty gote, about 70 percent wild type. Classifiable in
From Bridges and Morgan, 1919, Carnegie Inst. Wash. single dose in triploids (Schultz, 1934, DIS 1: 55).
Publ. No. 278: 148. RK1 at 28°-30°C; RK2 at 25°C.
cytology: Salivary chromosomes apparently normal.
/: (Bridges in Morgan, Bridges, and Schultz, 1937,
location* 2-48.7. Carnegie Inst. Wash. Year Book 36: 301).
origin: Spontaneous, J34*
A'scovemr. Bridges, 11111. origin: Spontaneous,
rtfwrmcos: Bridges mnd Morgan, 1919, Carnegie discoverer*. Duncan, 34e3.
fast. Wash. Pttbl. No. 278: 160 (fig.). phenotype: Resembles / but produces more vigorous
C!a»se<ra, 1924, J. Exptl. Zool. 38: 423-36. homozygous stock. RK1 at 28°C.
Stem, 1927, Biol. Zeatr. 47: 361-69.
phmmw«- Distal half of wing upturned. Curling is
s t r « f if wteg unfolds at 25-30°C, but weak or
wild type if wing unfolds below 25QC.
Placed to region between 34E5 and 35DI

un th* basis of its iaclaaioo in D§(2L)$4j «
Df'2Lt34E5-Fl,35C3-m (E. H. Gmll).
Jag; Jagged
<Ii£c#Y#f#r Stern. 25431.
MUTATIONS 115
Jag: Jagged origin: Spontaneous.

location: 2-54.9 (0.1 unit from Bl). discoverer: Bridges, 12f26.
discoverer: L. V. Morgan, 34b20. references: Bridges and Morgan, 1923, Carnegie
phenotype: Jag/+ has end of wing cut off; better in Inst. Wash. Publ. No. 327: 72 (fig.).
early counts and above 25°C. Jag/Jag has reduced Morgan, Bridges, and Sturtevant, 1925, Bibliog.
and roughened eyes and extremely jagged wings. Genet. 2: 214 (fig.), 227.
RK2 as heterozygote; RK3 as homozygote. phenotype: Eye size reduced by indentation of front
Jammed: see J margin. Tuft of vibrissae and hairs below eye.
jaunty: see / Variable; overlaps wild type. RK3.
jaunty x: see jyx *fc2
javelin: see jv origin: From progeny of heat-treated flies.
javelinlike: see jvl discoverer: Goldschmidt, 1927.
*je: jelly references: Gottschewski and Ma, 1937, Z.
location: 3-46. Induktive Abstammungs- Vererbungs lehre 73:
origin: Spontaneous; arose simultaneously with mu 584_97.
(3-50). phenotype: Eye reduced in size. Penetrance 50—80
discoverer: Mohr, 37121. percent. Expression variable. RK3.
references: Mossige, 1939, DIS 12: 47.
phenotype: Dark pinkish eye color. RK1. origin: Spontaneous.
jv: javelin discoverer: Gottschewski, 1937.
location: 3-19.2 (0.9 unit to left of dv). references: Gottschewski and Ma, 1937, Z.
discoverer: Mohr, 31j29. Induktive Abstammungs- Vererbungs lehre 73:
references: 1937, DIS 8: 12. 584-97.
Mohr and Mossige, 1943, Skrifter Norske phenotype: A weak allele; penetrance 10—20 per-
Videnskaps-Akad. Oslo, I: Mat.-Naturv. KL, No. cent. RK3.
7. 51 pp. (fig.). *kD: kidney-Dominant
phenotype: All bristles and hairs cylindrical instead origin: Spontaneous in chromosome containing k.
of tapered, with small enlargement before tip. discoverer: Puro, 60cll.
cytology: Placed between 64C12 and 65E1 on the phenotype: Eyes of heterozygote reduced at anterior
basis of its inclusion in Df(3L)Vn = edges. Expression variable; in extreme cases, eye
D[(3L)64C12-D1;65D2-E1. size about one-third normal. kD/k more extreme.
jvl: javelinlike Eyes of homozygote reduced about as much as
location: 3-56.7. kP/k, but occasionally one or both eyes missing;
origin: Spontaneous. antennae usually slightly deformed with thickened
discoverer: Ives, 4012. aristae. RK1.
references: 1942, DIS 16: 48. K-pn: Killer of prune
phenotype: Resembles ;v, bristles sometimes more location: 3-102.9 (0.2 to right of bv; recalculated
crooked. Viability and productivity somewhat from Sturtevant).
lower than normal. RK2. origin: Spontaneous.
*jyx: jaunty x discoverer: Sturtevant, 54a.
location: 1-24. references: 1955, DIS 29: 75.
origin: Spontaneous, 1956, Genetics 41: 118—23.
discoverer: Bridges, 14112. phenotype: No phenotypic effects, either when
phenotype: Wings curved up at tips. Viability about homozygous or when heterozygous, except that all
60 percent wild type. RK3. pn; K-pn flies die at end of second larval instar.
Kills all seven alleles of pn that have been tested.
K-pn eye disks transplanted to pn hosts develop
autonomously, as do the reciprocal transplants
(Grell, 1958, DIS 32: 123-24). RK3.
kar: karmoisin
location: 3-51.7.
discoverer: Pariser.
references: Gottschewski, 1935, DIS 4: 15.
phenotype: Eye color like st but less bright. Ocelli
ic: kidney white. Eyes contain 29 percent wild-type brown
From Bridges and Morgan, 1923, Carnegie Inst. Wash. pigment (Nolte, 1954, J. Genet. 52: 111-26).
Publ. No. 327: 77. Larval Malpighian tubes considerably lighter than
wild type but difficult to classify in living larvae
k: kidney (Brehme and Demerec, 1942, Growth 6: 351-56).
cytology: Placed in region 87D-F, on basis of its in- Viability somewhat reduced but fertility near
clusion inDf(3R)ry = Df(3R)87D-E;87E-F. normal. RK1 as heterozygote.
kar2 kidney: see k
origin: Spontaneous. kidney eye: see ke
discoverer: Bridges, 38blO. Killer of prune: see K-pn
phenotype: Like kar except that larval Malpighian Kinked: see Ki
tubes are bright yellow, similar to wild type kinked femur: see kf
(Brehtne and Demerec, 1942, Growth 6: 351—56),
RK1. *kk: kinky
kar31: karmotsin-3 lethal location: 1-42.
discoverer: Schalet. discoverer: Philip.
phenotype: Heterozygote of kar3I/kar resembles kar. phenotype: Bristles slightly bent or forked. RK3.
Homozygote lethal. RK2. other information: May be an allele of fw.
other information: kar-ry crossing over normal.
*ke: kidney eye KL: Male fertility complex in the long arm of the
location: 1-28.6. Y chromosome
origin: Induced by 2-chloroethyl methanesulfonate The male fertility complex of the long arm of the Y
(CB. 1506). chromosome, originally called KI by Stern (1929,
discoverer: Fahmy, 1956. Z. Induktive Abstammungs- Vererbungslehre 51:
references: 1959, DIS 33: 87. 253—353) and subsequently called KL by Brosseau
phenotype: Eyes small and extremely rough; anterior (1960, Genetics 45: 257—74), is subject to muta-
border indented, giving a kidney shape. Wings tions or deficiencies leading to male sterility,
small, abnormal, outspread, or upheld. Veins which are symbolized ms(Y)L. By complementation
thick, and often interrupted or fail to reach wing analysis, Brosseau divided KL into five different
margin, which is usually incised. Deformed complementing units designated kl-1 through kl-5.
antennae. Bristles straggly; occasionally one is By studying recombinants between the X and the
missing. Flies short lived; 50 percent die less y, i.e., detachments of attached X's, he ordered
than 24 hr after eclosion. Sterile, probably be- the complementation groups with respect to the
cause they are too weak to mate. RK3. centromere; he assigned the symbol kl-1 to the
*kf: kinked femur proximalmost and kl-5 to the distalmost.
location: 1-20.2. kn: knot
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- location: 2-72.3.
phenylalanine (CB. 3007). discoverer: Nichols-Skoog, 31hl.
discoverer: Fahmy, 1954. phenotype: Veins L3 and L4 shifted closer together
references: 1959, DIS 33: 87. in region of anterior crossvein, which is either
phenotype: Small fly with slightly dark, dull red eye extremely thick or eliminated by regional fusion of
color. Wings seldom fully expanded: when they do L3 and L4. Frequently extra crossvein between L3
expand, they are opaque and abnormal in shape. and L4 near end of wing. Wing narrowed. Head
Femurs kinked. Flies seem unable to move narrowed and flattened, so that long axis of eye is
normally and die on the food soon after eclosion. at oblique angle. May overlap wild type at high
RK3. temperatures and in late counts. Best at 19°C.
*Kg: Kugel RK2.
location: 3-48,2. *kno: knobbyhead
discoverer: B©nx, 1953. origin: Induced by triethylenemelamine (CB. 1246).
references: 1956, Rev. Suisse Zool. 63: 208-16. discoverer. Fahmy, 1951.
phenotype: Larva, pupa, and adult shorter and references: 1958, DIS 32: 70.
thicker than normal. Most striking in pup®. Homo- phenotype: Abnormal head; one or both eyes irreg-
xygote more estrero« than heterozygote. Homozy- ularly shaped, often drastically reduced in size.
gote viability 68 percent of wild type and fertility Occipital region frequently has hairy tufts, often
somewhat reduced. RK2. carried on protuberances. Males highly infertile;
Ki: Kicked viability about 10 percent wild type. RK2.
location: 3-47.6 (to the left ©f p). other information: One altele induced by CB. 2511.
origin: Spontaneous. knot: see Jen
discoverer: R. F. Grell, 571. Kr: KrUppel
references: 1958, DIS 32: 80. location: 2-108 (published value of 113 must be in-
pJwftotyfHK All bristles and hairs of beterozygote correct because the chromosome is only 108 units
shortened and twisted. Resembles am. Viability long)'
aKd fertility excellent; classification easy. Hooso- origin: Spontaneous,
has more-extretne bristle and limit effects. discoverer: Graber.
MUTATIONS
117
references: Gloor, 1950, Arch. Julius Klaus-Stift. phenotype: Eyes of L2/+ as small as or smaller than
Vererbungsforsch. Sozialanthropol. Rassenhyg. L/L. L2 homozygotes have tiny eyes and are
25: 38-44 (fig.). poorly viable or completely lethal, depending on
1954, Arch. Julius Klaus-Stift. Vererbungsforsch. background. Best used as heterozygote. Eyes
Sozialanthropol. Rassenhyg. 29: 277—87. further reduced in combination with M(3)w,
phenotype: Kr/+ adult sometimes has thoracic mal- M(3)h33i, and M(2)l2 (Dunn and Coyne, 1935, Biol.
formation; a leg or wing may be absent. Pene- Zentr. 55: 385—89). Classifiable in single dose in
trance low. Kr/Kr lethal before hatching. Primary triploids (Schultz, 1934, DIS 1: 55). Reduced
body segments of embryo abnormal, particularly number of cells enter into formation of eye disks
median segments. Ventral chain of ganglia discon- (Steinberg, 1944, Proc. Natl. Acad. Sci. U.S. 30:
nected. Tracheal system defective. Malpighian 5—13). RK1 as heterozygote.
tubules missing. Salivary glands normal. RK2.
KS: Male fertility complex in the short arm of the
Y chromosome
The male fertility complex of the short arm of the Y
chromosome, originally called K2 by Stern (1929,
Z. Induktive Abstammungs- Vererbungslehre 51:
253—353) and subsequently called KS by Brosseau
(1960, Genetics 45: 257—74), is subject to muta-
tions and deficiencies leading to male sterility,
which are symbolized ms(Y)S. By complementation
analysis, Brosseau divided KS into two comple-
menting units, ks-1 and ks-2. He believes that the
most probable order of factors on Ys from the tip L2: Lobe-2
toward the centromere is ks-2, ks-1, bb. Edith M. Wallace, unpublished.
Kugel: see Kg
kz: kurz
location: 1-0.9 (to the right of pn). origin: Spontaneous.
origin: Spontaneous. discoverer: Bridges, 24dlO.
discoverer: Stern, 26a23. references: Morgan, Bridges, and Sturtevant, 1925,
references: 1930, Z. Induktive Abstammungs- Bibliog. Genet. 2: 230.
Vererbungslehre 53: 279-86. phenotype: Eye reduction intermediate between L
1934, DIS 1: 35. and L2, but variability high. RK2.
phenotype: Bristles shorter and finer, like a slight
Minute. Postcutellars often absent. Hatches some-
what late. Viability fair; both sexes fertile. RK2. origin: Spontaneous.
cytology: Salivary chromosome location in region discoverer: Sturtevant, 23f.
2E1 through 2F6 (Demerec, Kaufmann, Fano, synonym: Lc.
Sutton, and Sansome, 1942, Carnegie Inst. Wash. phenotype: Heterozygote has fairly constant reduc-
Year Book 41: 191). tion in eye size, but not so great as to exclude its
use in combination with most eye colors. Expres-
L: Lobe sion more extreme at 25° than at 19°C. Hoaoxygot©
location: 2-72.0. has smaller eyes but lowered viability. Six® of
origin: Spontaneous. eyes of L4/+ reduced in combination with M(3)w,
discoverer: Bridges, 18i24. M(3)h33i, and M(2)!2 (Dunn and Coyne, 1935, Biol.
references: Morgan, Bridges, and Sturtevant, 1925, Zentr. 55: 385—89). Development similar to L3
Bibliog. Genet. 2: 230 (fig.). (Steinberg, 1944, Proc. Natl. Acad. Sci. U.S. 30:
phenotype: Heterozygous L eyes slightly smaller, 5—13). Reduced size of cephalic complex detect-
with nick in anterior edge, and lower half of eye able in 24-hr larva, but subsequent growth rate
reduced more than upper; overlaps wild type. similar to wild type (Medvedev, 1935, Z. Induktive
Homozygous L, eyes much smaller and less vari- Abstammungs- Vererbungslehre 70: 55—72, Tr.
able. Size of L/+ eyes reduced in combination with Inst. Genet. Akad. Nauk. SSSR 10: 119-51). RK2
M(3)w, M(3)h33), and M(2)l2 (Dunn and Coyne, as heterozygote.
1935, Biol. Zentr. 55: 385-89). Best used as a 5
recessive. RK2. origin: Spontaneous,
L.2 discoverer: Mohr, 3!k26.
origin: Spontaneous. references: Dunn, 1935, DIS 4: 14.
discoverer: Mohr, 20b2. pKenotyp©: Heterozygote ha* small nick in eyes,
references: 1924, Z. Induktive Abstammungs- overlap* wild type. Hotnoajygote has small ef®s
Vererbungslehre 32: 216. and tendency to antenna reduplication. May h«
Morgan, Bridges, and Sturtevant, 1925, Bibliog. used as a recessive but not as a dominant. Mor#
Genet. 2: 230. extreme than Lr but less so than L*. Development
similar to that of L? (Steinberg, 1944, Proc. Natl. phenotype: Eye reduction strong, with little varia-
Acad. Sci. U.S. 30: 5-13). RK3. tion in heterozygote. Homozygote more extreme;
L34 viability and fertility high. RK2.
origin: Spontaneous. cytology: Salivary chromosomes normal.
discoverer: Glass, 1934. Lr: Lobe-recessive
phenotype: Between L and L4. Dominance varies in discoverer: L. V. Morgan, 29h23.
different crosses. RK2. phenotype: Homozygote has small kidney-shaped
L52c eyes. Overlaps wild type at 19°; generally good at
origin: Spontaneous. 25°C. Heterozygote rarely shows seam or nick.
discoverer: Nakayama, 52c. RK2 as homozygote.
references: 1953, DIS 27: 59. Lrt>: Lobe-rough
phenotype: Like L. RK2. origin: Ultraviolet induced.
discoverer: Edmondson, 49k.
LB: Lobe of Becker references: Meyer, Edmondson, Byers, and Erickson,
origin: Spontaneous in In('2L)Cy +In(2R)Cy. 1950, DIS 24: 60.
discoverer: Becker. phenotype: Eye reduction similar to L4, but eye
references: 1957, Z. Induktive Abstammungs- surface slightly rough.
Vererbungslehre 88: 333—73 (fig.). Homozygote has extreme reduction of eyes; few or
phenotype: Not separable from Cy; homozygote no facets. L4/LTO has similar reduction in size of
therefore cannot be tested. Lower half of eyes eyes. Viability and fertility excellent. RK1 as
reduced or absent in heterozygote; more extreme at homozygote; RK2 as heterozygote.
25° than at 18°C. Sectors of ommatidia replaced by Lsi: Lobe-sinuate
chitin and bristles. Lower half of eyes apparently origin: Spontaneous.
produced from fewer than the normal 9 or 10 pre- discoverer: Morgan, 1932.
sumptive ommatidia-producing cells. Lower half of phenotype: Eyes of heterozygote flat, smooth, nearly
head also reduced at 25° but not 18°C. Tempera- full size, with sinuate margin; overlaps wild type
ture-sensitive period for ommatidia formation first only slightly. Eyes of homozygote smaller, with
and second ins tars; third instar as well for head flat or concave contour, smooth surface, and
reduction. RK2A. sinuate lower margin. RK3.
other information: Allelism to L inferred from pheno-
type and linkage to Cy alone. /( ): lethalf )
Lc: see L4 General term used to describe recessive mutations
*Ld: Lobe-duplicating that lead to death of most or all homozygous car-
origin: Spontaneous. riers. The symbol / is followed parenthetically by
discoverer: Kodani. the chromosome and then by the designation of the
references: Zimm, 1951, J. Exptl. Zool. 116: 289— particular mutant. Unfortunately, it is not practi-
319 (fig.). cable, except in special cases, to test allelism of
phenotype: Partially dominant. Characterized by sex-linked lethals, and it has not been common to
incomplete penetrance and variable expression: retain and test allelism of autosomal lethals. Con-
reduced single or bipartite eyes, kidney-shaped sequently, little information on allelism of lethals
eyes with bristles, large eyes with palps, or dupli- with similar genetic location is included.
cated antennae. Modifiers present on chromosomes 1-mah see l(l)m
I and 3. Penetrance affected by temperature during *l(l)h lethcl(l) J
development. RK3. location: 1-1.1.
*Ld*: Lobe-diminished discoverer: Rawls, 12b.
origin: Spontaneous. references: 1913, Biol. Bull. 24: 115-24.
discoverer: Kadel and Jenkins, 55g. Morgan and Bridges, 1916, Carnegie Inst. Wash.
synonym: dq. Publ. No. 237: 31.
references: Kadel, 1956, DIS 30: 73-74. other Information: First recessive lethal found in
1957, DIS 31: 83. D. melanogaster.
phenotype: LdQ/+ normal. Eyes of Ld<3/Ldti irregu- KD3C3
larly deformed; occasionally divided into two or location: 1-1.6 (between w and rst).
root* lobes. Expression variable; some overlap of origin: Synthetic.
wild type, especially in old cultures. Aristae discoverer: Lefevre and Wilkins.
reduced and deformed. RK2. references: 1964, Genetics 50: 264.
phenotype: Male lethal. l(l)3C3/w is normal. RK2.
LK: Lobe of Krivsh&nko cytology: Associated with the deficiency for band
origin: Spontaneous. 3-C3 obtained as a single recornbinant carrying the
discover or: Krivshenko, 1957. left ©nd of T(l;4)w**J « T(t;4)3C2-3;2O;lQ2C and
references: 1958, DB 32: 81. the right end of InfDnt3 * In(l)3C3-4;20B.
MUTATIONS 119
cytology: Associated with T(l;3)l-184 -

location: 1- (0.4 to the left of y). T(1;3)18A;81.
origin: Spontaneous. *I(J)272-J3
discoverer Bridges, 14d9. location: 1- (rearrangement).
references: 1916, Genetics 1: 149. origin: X ray induced.
1(1)7: see discoverer: Demerec, 1940.
references: Sutton, 1943, Genetics 28: 210—217.
location: 1-0. phenotype: Lethal. 1(1)272-13/'sc is scute. RK2A.
origin: Spontaneous in dor1. cytology: Associated with In(l)l-272-13 =
discoverer: Bridges, 1928. In(l)lA6-Bl;llA7-8;llF2-12Al;18A4-Bl.
phenotype: Probably a specific modifier of
1(1)7'e dor1 dies earlier than dor1. RK3. origin: Spontaneous.
cytology: Included in 2RDXP element of T(l;2)Bld discoverer: Demerec, 1936, 1937.
T(1;2)1C3-4;6OB12-13 whereas dor1 is not. references: Slizynski, 1938, Genetics 23: 283—90.
cytology: Salivary analysis (Slizynski, 1938) shows
location: 1-21.3 (19.0 to 23.6). some normal, some deficient, and some inverted for
discoverer: Sobels. single-lettered subdivisions of the X chromosome.
references: Gloor, 1962, Rev. Suisse Zool. 69: other information: A series of 13 independently
409-63 (fig.). occurring and genetically located lethals.
phenotype: Larvae lethal in third instar; survive up *l(l)294-
to 10 days. Testes and lymph glands degenerate. origin: X ray induced.
Imaginal disks develop normally after transplanta- discoverer: Demerec, 36d.
tion. Protein metabolism disturbed; free amino references: Slizynski, 1938, Genetics 23: 283—90.
acids and peptides abnormally high. RK2. cytology: Three normal, one deficient for 10B, and
one deficient for IOC.
l(l)48j: see l(l)mys other information: A series of five independently
*I(J)52 induced and genetically located lethals.
location: 1- (to the right of B). *l(l)296-
discoverer: Sobels. origin: Spontaneous.
references: Gloor, 1962, Rev. Suisse Zool. 69: discoverer: Demerec, 1936.
409-63 (fig.). references: Slizynski, 1938, Genetics 23: 283-90.
phenotype: Larvae die in second instar. Growth cytology: Salivary chromosomes normal.
retarded. Histology of nervous system, testes, and other information: A series of six independently
imaginal disks abnormal. Number of nuclei in occurring and genetically located lethals.
salivary glands increased. Amino acids and pep- *l(l)302-
tides increased. Transplanted testes and imaginal origin: Neutron induced,
disks autonomously lethal. RK2. discoverer: Demerec, 36k.
references: Slizynski, 1938, Genetics 23: 283-9Q.
location: 1- (claimed to be about 0.6 unit to the left cytology: Three normal; one deficient for 3F.
of y, making it the leftmost known locus on the other information: A series of four independently
first chromosome). induced and genetically located lethals.
discoverer: Burdick, 55a. */(7)304-
1957, DIS 31: 86. discoverer: Demerec, 37d.
phenotype: Presumed to be a lethal. Heterozygote references: Slizynski, 1938, Genetics 23: 283-90.
claimed to have viability about 1.5 times normal. cytology: Three normal; one deficient for 4C.
Not allelic to 1(1 )J1. RK2. other information: A series of four independently
other information: Map location of this mutation induced and genetically located lethals.
must be near y; however, the published data (1957, l(l)bt: see brl-
DIS 31: 86) permit the following alternative to a KDC
location to the left of y: l(l)55a is not completely location: 1-6 (between ec and bi).
lethal, and the 1.29 percent scored as crossovers origin: Spontaneous in sc t2 v si B chromosome.
to the left of y are actually surviving noncross- discoverer: Muller, 20j.
overs. references: 1928, Genetics 13: 279-357.
1(1)76: see dor13 phenotype: Dies as late embryo or, more commonly,
as first-instar larva (Brehme, 1937, Am. Naturalist
location: 1- (rearrangement), 71: 567). RK2A.
origin: X ray induced. cytology: Associated with the left breakpoint of
discoverer: Lindsley, Edington, and Von Halle, In(l)Cl «= In(l)4A5-Bl;17A6-Bl.
references: 1960, Genetics 45: 1649—70. *I(1)DM: lethol(l) of D. Mor/wofcj
phenotype: Almost completely lethal. The few sur- origin: X ray induced.
vivors have dark rough eyes. RK2A. discoverer: Moriwaki, 1932.
references: 1934, Japan J. Zool. 5: 585-602. I(1)EN7

1940, DIS 13: 50. location: 1- (rearrangement).
other information: Five independently induced and origin: X ray induced.
roughly located lethals. discoverer: Novitski.
I(1)ENT: lethal(l) from Eugene nonautonomous phenotype: Usually dies as early pupa; survives as
location: 1-46. patches of hemizygous tissue in gynandromorph.
origin: X ray induced. Fat bodies beaded instead of ribbonlike in third
discoverer: Novitski. larval instar. More free glutamine but less free
references: 1963, DIS 37: 51-52. tyrosine than normal. RK2A.
phenotype: Dies between first instar and prepupa; other information: Crossing over in X greatly re-
survives as patches of hemizygous tissue in gynan- duced.
dromorph. Salivary glands and gastric ceca small, I(1)EN8
and fat bodies usually absent in third-ins tar larva; location: 1- (close to the left of cv).
excess of free alanine or closely related sub- origin: X ray induced.
stance, and less free tyrosine than normal. RK2. discoverer: Novitski.
I(1)EN2 references: 1963, DIS 37: 52.
location: 1-0.3. phenotype: Usually dies in second larval instar; 25
origin: X ray induced. percent survive to third instar and a few to pupa-
discoverer Novitski. tion. Survives as patches of hemizygous tissue in
references: 1963, DIS 37: 52. gynandromorph. Fat bodies, Malpighian tubes, and
phenotype: Dies at end of third instar or in early salivary glands reduced. Rare adult survivors have
pupa; survives as patches of hemizygous tissue in soft exoskeleton and die within a few days. More
gynandromorph. More free glutamine but less free free glutamine but less free tyrosine than normal.
glutamic and as part ic acids than normal. RK2. RK2.
location: 1- (near car). I(1)EN9

origin: X ray induced. location: 1-10.
discoverer: Novitski. origin: X ray induced.
references: 1963, DIS 37: 52. discoverer: Novitski.
phenotype: Dies shortly after pupation; survives as references: 1963, DIS 37: 52.
patches of hemizygous tissue in gynandromorph. phenotype: Usually dies as third-instar larva; sur-
Larva usually has red-black pigmented areas in or vives as patches of hemizygous tissue in gynan-
on the cuticle. More free glutamine than normal. dromorph. Larva becomes transparent. Salivary
RK2. glands, Malpighian tubes, and fat bodies much re-
I(1)EH4 duced. Unknown fluorescent substance accumu-
location: 1-52. lates in larval cuticle. More free glutamine but
origin: X ray induced. less free tyrosine than normal. RK2(A).
discoverer Novitski. other information: Crossing over suppressed at tip
references: 1963, DIS 37: 52. of-X.
phenotype: Roughly 40 percent eclose but die imme- !(1)ENJO
diately; survives as patches of hemizygous tissue location: 1-59.
in gynandromorph. No morphological abnormalities origin: X ray induced.
observed in larva or pupa. More free glutamine discoverer: Novitski.
than normal. RK3. references: 1963, DIS 37: 52.
I(1)EN5 phenotype: About half die as pupae and half as few-
location: 1-47. day-old adults. Survives as patches of hemizygous
origin: X ray induced. tissue in gynandromorph. More free glutamine than
discoverer: Novitski. normal; free tyrosine nearly absent; low in free
references: 1963, DIS 37: 52. proline. RK3.
phenotype: Usually dies before third instar; sur- KDENJOa
vive® as patches of hemizygous tissue in gynan- location: 1-50.
droaaorpfa. More free glutamine than normal. RK2. origin: X ray induced.
1(1)EN6 discoverer: Novitski.
location: 1- 63 [between cmr and au(f)]. references: 1963, DIS 37: 52.
origin: X ray induced. phenotype: Dies as pupa. Third-instar larva shows
discoverer. Novitski. large excess of free glycine. RK2.
r«f©rtmc«s: 1963, DIS 37: 52. KDEHll
pbenotyp*: Dies at any stage; survives as patches location: 1-43.
of bemizygous tissue in gynandromorph. Larval fat origin: X ray induced.
bodies and Malpighian tubes reduced. More free discoverer: Novitski.
glutamine than normal. RK2. references: 1963, DIS 37: 52.
MUTATIONS 121
phenotype: Dies from second instar through pupa. movements but is unable to break through vitelline
Survives as patches of hemizygous tissue in membrane. Muscular activity persists several
gynandromorph. Melanotic spots on some larvae hours, but hatching does not occur and cell degen-
and inside pupae. Culture with dying larvae has eration begins at about 25 hr. Differentiation
distinct urinous odor. More free phenylalanine and abnormal in several ways: pharyngeal apparatus
less free tyrosine than normal. RK2. reduced and distorted; brain forms irregular mass;
I(1)ENU constriction forms behind head; segmentation dis-
location: 1-3. torted; and body wall usually incomplete dorsally.
origin: X ray induced. RK2.
discoverer: Novitski. *I(1)GSB: lethal(l) of Gershenson, Shapiro, and
references: 1963, DIS 37: 52. Borissenko
phenotype: Usually dies in third larval instar or origin: X ray induced in In(l)scs.
pupa. Rare survivors reach eclosion. Survives as discoverer: Gershenson, Shapiro, and Borissenko,
patches of hemizygous tissue in gynandromorph. 1931.
Flies that reach eclosion have soft exoskeleton references: Gershenson, 1934, DIS 1: 54.
with little pigmentation; appear almost translucent. other information: A series of 51 independently
Low in free tyrosine. RK2. induced and genetically located mutants.
KDEN13 *l(l)l: lethal(i) of Ives
location: 1-13.4. origin: Recovered from heat-treated lines.
origin: X ray induced. discoverer: Ives.
discoverer: Novitski. references: Plough and Ives, 1934, DIS 1: 32.
references: 1963, DIS 37: 52. 1935, Genetics 20: 42-69.
phenotype: Some survivors. No gross larval or pupal other information: A series of 13 independently
abnormalities. Low in free tyrosine. RK3. occurring mutants.
KVENU i(l)Jh lethal(i) of Jacobs-Duller
location: 1- (rearrangement). location: 1-0.0 (to the left of y).
origin: X ray induced. origin: X ray induced simultaneously with scJ*.
discoverer: Novitski. discoverer: Jacobs-Muller.
references: 1963, DIS 37: 52-53. references: Muller, 1932, Proc. Intern. Congr.
phenotype: Dies mostly in second, but also in third- Genet., 6th. Vol. 1: 225.
instar larva; survives as patches of hemizygous Muller, 1935, Genetica 17: 237-52.
tissue in gynandromorph. Accumulates propanol- phenotype: Lethal. Not cell lethal (Ephrussi, 1934,
ammonia-insoluble fluorescent substance in larval Proc. Natl. Acad. Sci. U.S. 20: 420-22). One
cuticle. Less free tyrosine and proline than recorded surviving male had rough eyes and was
normal. RK2A. sterile. RK2A.
other information: Crossing over in X greatly re- cytology: Probably in 1A6. Associated with
duced. In(l)scJ1 =In(l)lA4-5;lB4-5 (Muller, Prokofyeva,
KDEN15 and Raffel, 1935, Nature 135: 253-55).
location: 1- (near car). I(1)J12S9
origin: X ray induced. origin: X ray induced in y-bearing chromosome.
discoverer Novitski. discoverer Lindsley, Edington, and Von Halle,
references: 1963, DIS 37: 53. references: Frye, 1959, Genetics 44: 511.
phenotype: Dies as third-instar larva or early pupa; *l(l)jl: lethal(l) /aw/ess
survives as patches of hemizygous tissue in location: 1-14.
gynandromorph. Less free tyrosine and proline origin: Ultraviolet induced,
than normal. RK2. discoverer: McQuate, 1951.
t(l)ENU references: Oster, 1952, Heredity 6: 403-7.
location: 1-24. phenotype: Dies during first larval instar. Mouth
origin: X ray induced. parts poorly formed and sometimes absent. RK2.
discoverer Novitski. cytology: Salivary chromosomes normal (Valencia
references: 1963, DIS 37: 53. and McQuate, 1951, Genetics 36: 580).
phenotype: Dies between first-instar larva and pupa. *I(1)K: lethalO) of King
Less free tyrosine and proline than normal. RK2. origin: Recovered among progeny of males fed P32.
*!(l){fn: lethal(l) formalin food discoverer: R. C. King, 1948 and 1949.
location: 1- (not located). references: 1950, DIS 24: 58.
origin: Induced by formaldehyde. other informotion: Four independently induced and
discoverer: Auerbach. genetically located lethals.
synonyms: Ltlll. *IO)LB: lethal(l) of Luers and Belitz
references: Ede, 1956, Arch. Entwicklungsmech. discoverer: Luers and Belitz, 1951-1956.
Organ. 148: 416-36 (fig.). references: Belitz, 1954, Z. Induktive Abstarnrnungs-
phenotype: Develop® to late embryonic stage, at 22 Vererbungslehre 86: 173-84.
hr (normal hatching time) shows vigorous muscular 1956, DIS 30: 104.
other information: A series of nearly 500 mutations phenotype: 20-hr embryos (25°C) show mid-dorsal
recovered from untreated males or from males herniation of brain or mid-gut, or both, abnormal
treated with Miracil [l-(2-diethylaminoethylamino)- somatic, visceral, and pharyngeal muscles, and
4-methylthioxanthineJ, Mirasan, triethylenemela- incomplete morphogenesis of yolk-filled mid-gut.
mine, aminopterin, hesperidine, or 2,5-bisethylene- Development of embryo normal up to 13 hr.
imine-l,4-benzoquinone. All lethals were located Between 13 and 14.5 hr, first muscular contrac-
genetically. tions occur, while basement membrane is incom-
*l(l)m: lethal(l) malignant plete. This results in dorsal rupture of hypoderm,
location: 1- (not located). retraction of myogenic elements of somatic and
origin: Induced by mustard gas. pharyngeal muscles into spheroidal masses. Con-
synonym: l-m&l. tinuation of myogenesis produces spheroidal
references: El Shatoury, 1955, Arch. Entwicklungs- muscles with a cortex of disoriented fibrillae sur-
mech. Organ. 147: 496-522 (fig.). rounded by a medulla of nucleated sarcoplasm.
El Shatoury and Waddington, 1957, J. Embryol. RK2.
Exptl. Morphol. 5: 143-52 (fig.). cytology: Salivary chromosomes normal.
phenotype: Cells originating from lymph glands in
late third instar first spread to, and cause, *l(l)nd: lethal(l) no differentiation
destruction of imaginal buds and later may move location: 1- (not located).
along ventral nerve cord to attack posterior fat origin: Induced by mustard gas.
bodies and testes. The tumor cells eventually references: El Shatoury, 1955, Arch. Entwicklungs-
become melanotic after destruction of various mech. Organ. 147: 523-38 (fig.).
healthy tissues. Death occurs in late larval or phenotype: Some or all imaginal buds fail to differ-
early pupal stages. Claimed to be the only true entiate during larval third instar, apparently as a
malignancy in Drosophila melanogaster, RK2. result of abnormal proliferation of imaginal disk
mesoderm. Death in pupal or prepupal stage. RK2.
*I(1)MA: lethal(l) of Mailer and Altenburg
origin: Spontaneous. *l(l)ne: lethal(l) nonevaginated
discoverer: Muller and Altenburg. location: 1-0.1.
references: 1919, Proc. Soc. Exptl. Biol. Med. 17: origin: Induced by urethane.
10-14. discoverer: Vogt, 1949.
other information: A series of about 50 mutants of references: 1951, DIS 25: 76.
which only a few were located. Florschutz-de Waard and Faber, 1952, DIS 26: 99.
1(1 )ml: lethal(l) melanoma!ike Faber, Sobels, Florschiitz-de Waard, and
location: 1-10. Oppenoorth, 1954, Z. Induktive Abstammungs-
origin: Ultraviolet induced. Vererbungslehre 86: 293—321 (fig.).
discoverer: McQuate, 1951. phenotype: Lacks imaginal thoracic hypoderm.
references: Oster, 1952, Heredity 6: 403-7. Cephalic complex and thoracic imaginal disks fail
Oster and Sobels, 1956, Am. Naturalist 90: 55-60. to evaginate. The unaffected abdominal hypoderm
phenotype: Larvae die in third instar. At death, develops but ends anteriorly in a free edge that
they have internal melanotic masses (usually one folds back on itself and forms a darkly pigmented
or two, sometimes as many as ten). RK2. ring around the pupa. Genital disk capable of
cytology: Salivary chromosomes normal (Valencia normal evagination but vasa deferentia do not
and McQuate, 1951, Genetics 36: 580). connect to testes, which do not spiralize. Death
*l0)mt: lethal(l) midget occurs 3—5H days after prepuparium formation.
location: 1-2.5. Pupae darker than normal, with sticky, irregular
origin: Ultraviolet induced. surface and distinctly meandering tracheal trunks.
discoverer: McQuate, 1951. RK2.
references: Oster, 1952, Heredity 6: 403-7. *l(l)nib; lethal(l) no imaginal buds
phenotype: Dies as undersized third instar larva. location: 1- (not located).
RK2. references: El Shatoury and Waddington, 1957, J.
cytology. Salivary chromosomes normal (Valencia Embryol. Exptl. Morphol. 5: 143—52 (fig.).
and McQuate, 1951, Genetics 36: 580). phenotype: Dies in third larval instar. Imaginal
l(l)mys: lethal(l) myospheroid buds small or absent. Excessive proliferation of
location: 1-21.7. stomach epithelium leads to occlusion of gut. Pro-
origin: Induced by P 3 2 . liferations degenerate into melanotic masses.
discoverer: Poulson, 48j. RK2.
synonyms: 1(1)48). 1(1 )Q: lethal(l) Quinacrine mustard induced
references: Rizki, 1956, J. Exptl. Zool. 131: origin: Induced by 2-methoxy-6-[3-(ethyl-2-chloro-
203-22 (fig.). ethyl)aminopropylaminojacridine (ICR 100).
Wright, 1958,, Proc. Intern. Congr. Genet., 10th. discoverer: Carlson.
Vol. 2: 323. references: Carlson, Sederoff, and Cogan, 1967,
1960, J. Exptl. Zool. 143: 77-99 (fig.). Genetics 55: 295-313.
MUTATIONS 123
other information: A series of 64 independently Number Location

induced and genetically located lethals. Their * 67 20.3
numbers and locations are tabulated below. * 68 40.4
Number Location * 69 54.1
* 1 13.1 * 70 51.9
* 2 40.9 * 71b 54.4
* 3 26.5 * 72 10.6
* 4 52.0 * 73 53.9
* 5 0.0 * 74 18.2
* 6 30.2 75 29.3
* 7 28.8 * 76 49.1
* 8 65.5 * 77 0.0
* 9 1.4 * 78 0.0
* 10 28.6 * 79 48.0
* 11 16.0 * 80 54.2
* 12 42.0 * 81 6.8
* 13 49.7 * 82 18.7
* 14 64.2 * 83 12.1
* 15 19.6 * 85 0.0
* 16 52.6 86 56.7
* 17 23.6 87 0.0
* 18 56.7 * 89 51.5
* 19 62.5 *201 13.4
* 20 0.0 202 40.1
21 20.5 203 62.5
22 39.1 *2O4 22.6
* 23 57.8 *205 21.1
* 24 58.3 206 13.0
* 25 23.0 208 17.0
* 26 33.0 *209 0.0
* 27 9.3 •210 64.5
* 28 44.6 •211 33.0
* 30 22.7 212 0
* 31 56.7 *214 56.7
* 33 29.9 *215 10.4
* 34 56.7 216 8.6
* 36 0.0 217 0.0
* 39 1.7 218 2.8
* 40 1.5 *219 1.3
* 41 2.0 220 0.0
* 42 64.4 221 0.0
* 43 37.6 222 57.9
* 44 31.7 223 29.2
* 45 38.3 224 47.8
* 48 21.3 225 36.0
* 49 12.7 226 47.9
* 50 62.5 227 27.4
* 51 65.2 *228 42.2
52 36.9 *231 13.5
* 53 32.0 *232 16.5
* 54 33.0 233 15.2
* 55 60.6 234 38.9
* 56 11.2 235 28.1
* 57 16.8 236 21.0
58 38.2 237 21.7
* 59 44.5 *238 54.1
61 6.5 *240 33.0
* 62 62.5 • 244 24.2
* 63® 53.0 248 12.2
64 41.4 *l(l)R: tetbalO) of Rohrborn
65 20.5 discoverer: Rohrborn, 1955, 1956.
* 66 33.0 reference*: 1959, Z. Vererbungslehrts 90: 116-31.
other information: A series of 71 lethals including 3 discoverer: McQuate, 1951.

spontaneous ones, 55 from males treated with 1:4- references: Oster, 1952, Heredity 6: 403—7.
dimethanesulfonoxybutane (CB. 2041), and 13 from phenotype: Dies during third larval instar. Main
males treated with l:4-dimethanesulfonoxy-l:4- tracheal tubes greatly enlarged, sometimes lack
dimethylbutane (CB. 2348). All were located functional posterior spiracles. RK2.
genetically. cytology: Salivary chromosomes normal (Valencia
*l(l)rr: lethal(l) ring gland rudimentary and McQuate, 1951, Genetics 36: 580).
location: 1-0.3. *l(l)th lethal(l) tracheae lacking
origin: Ultraviolet induced. location: 1-59.
discoverer: McQuate, 1951. origin: Ultraviolet induced.
references: Oster, 1952, Heredity 6: 403-7. discoverer: McQuate, 1951.
phenotype: Dies during third larval instar. Larvae references: Oster, 1952, Heredity 6: 403—7.
live 15—30 days but do not become giant. Ring phenotype: Dies during first larval instar. Main
gland abnormally small, probably causing failure tracheal tubes absent, although small side
to undergo third molt. RK2. branches present. RK2.
cytology: Salivary chromosomes normal (Valencia cytology: Salivary chromosomes normal (Valencia
and McQuate, 1951, Genetics 36: 580). and McQuate, 1951, Genetics 36: 580).
*I(1)S: leihal(l) of Stark
origin: Spontaneous. *l(l)tr: lethal(l) tracheae ramified
discoverer: Stark, 1913, 1914. location: 1-56.
references: 1915, J. Exptl. Zool. 19: 531-58. origin: Ultraviolet induced.
Morgan and Bridges, 1916, Carnegie Inst. Wash. discoverer: McQuate, 1951.
Publ. No. 237: 64, 79. references: Oster, 1952, Heredity 6: 403-7.
other information: Four independently occurring phenotype: Dies during first larval instar. Main
lethals. tracheal tubes thick and have numerous side
*I(1)S9 branches. RK2.
location: 1- (to the right of car). cytology: Salivary chromosomes normal (Valencia
origin: Spontaneous. and McQuate, 1951, Genetics 36: 580).
discoverer: Auerbach.
references: Ede, 1956, Arch. Entwicklungsmech. *l(l)trs: lethal(l) tracheae stretched
Organ. 149: 256-66 (fig.). location: 1-8.0.
phenotype: Almost all embryos deformed at anterior origin: Ultraviolet induced.
end, where there is usually some undigested yolk. discoverer: McQuate, 1951.
Death occurs in embryonic, larval, and pupal synonym: l(l)ts (preoccupied).
stages. Primary abnormality is distribution of references: Oster, 1952, Heredity 6: 403—7.
cleavage nuclei, which causes blastoderm to be phenotype: Dies during first larval instar. Larvae
fragile at its anterior end. RK2. very large for this stage and all tracheal tubes very
l(l)sc: lethal(l) at scute thin, suggesting that they grow more slowly than
location: 1-0 (immediately to the right of sc). larvae and thus become stretched. RK2.
origin: Synthetic. cytology: Salivary gland chromosomes normal
discoverer: Muller. (Valencia and McQuate, 1951, Genetics 36: 580).
references: 1935, Genetica 17: 237—52. *l(l)ts: lethal(l) temperature sensitive
other information: Inferred from the inviability of location: 1-8.
In(l)nc*L&c9R = In(l)lB3-4;19F-20Cl^lB2- discoverer: Falbo and Re'.
3;18B8-9R [left break of In(l)sc9 in doubt] except references: 1945, DIS 19: 45, 57.
in the presence of Dpfi;2)sc* *\ phenotype: Inviable in cultures grown at 23°C but
shows more than 50 percent survival in cultures
*l(1)sd: lethal(l) schaiben defekt grown at 26.5°. RK3.
location: 1-17.9. *l(l)TS-45: lethal(l) no. 45 of T. Shiomi
origin: Induced by triethylenemelamine (CB. 1246). location: 1-5.8.
discoverer: M. J. Fahmy. origin: X ray induced.
references: Schnitter, 1961, Rev. Suisse Zool. 68: discoverer: Shiomi, 52f.
345-418 (fig.). references: 1954, DIS 28: 78.
phenotype: Dies during transition from larva to Imaizumi and Shiomi, 1955, Arch. Biol. (Liege) 66:
prepupa. Some larvae form puparia but do not dif- 483-87.
ferentiate further. Pattern of damage complex, most phenotype: Dies before hatching. No visible morpho-
severe defects being found in certain imaginal logical abnormality. Heterozygote of l(l)TS-45/
disks. Several larval organs abnormal, especially Base has average of 612 eye facets compared to
the salivary glands. RK2. only 402 in +/Bssc. Accumulation of urea or car-
*l(l)te: lethal(l) tracheae enlarged barnides in larvae of heterozygote; these compounds
location; 1-0.3.
presumably tend to normalize the Bar phenotype.
origin: Ultraviolet induced. RK2.
MUTATIONS 125
*l(l)TS-56 references: 1960, Genetics 45: 1649—70.

location: 1-1.5. phenotype: Viability of X/Y males 63 percent
origin: X ray induced. normal; further reduced in presence of E(var)7 and
discoverer: Shiomi, 52f. M(2)S2l°. X/Y males fertile. X/0 males lethal.
references: 1954, DIS 28: 78. Homozygous females viable, with fewer and smaller
phenotype: Lethal in late embryonic stage. Develop- bristles. RK2A as X/O male.
ment of tracheae, other chitinized parts, and body cytology: Associated with In(l)l-v59 = In(l)3-
segments abnormal. RK2. 4; 19-20.
I(1)v75
I(l)v3: lethal(l) variegated location: 1- (rearrangement).
location: 1- (rearrangement). origin: X ray induced.
origin: X ray induced. discoverer: Lindsley, Edington, and Von Halle.
discoverer: Lindsley, Edington, and Von Halle. references: 1960, Genetics 45: 1649—70.
references: 1960, Genetics 45: 1649-70. phenotype: Viability of X/Y males 26 percent
phenotype: Viability of X/Y males 74 percent normal; further reduced in presence of E(var)7 but
normal; recovery of X/Y males reduced by M(2)S210 not M(2)S210. X/Y males sterile, owing to failure
but not E(var)7. X/Y males sterile, owing to failure of sperm head to elongate. Viability of X/0 males
of sperm head to elongate. X/0 males lethal. less than 1 percent normal. RK2A as X/0 male.
RK2A as X/0 males. cytology: Associated with T(l;2)l-v75 - T(l;2)19-
cytology: Associated with T(l;3)l-v3 = T(1;3)4A;81, 20;41.
\(})vU9
location: 1- (rearrangement). location: 1- (rearrangement).
discoverer: Lindsley, Edington, and Von Halle. discoverer Lindsley, Edington, and Von Halle.
references: 1960, Genetics 45: 1649-70. references: 1960, Genetics 45: 1649—70.
phenotype: Viability of X/Y males 78 percent phenotype: Semilethal. Viability of X/Y males 91
normal; further reduced in presence of M(2)S2*° but percent normal, of X/0 males 26 percent normal,
not E(var)7. X/Y males fertile. Viability of X/0 X/Y males sterile, owing to failure of sperm head
males 4 percent normal. Homozygous females sur- to elongate. RK3A.
vive and have blistered wings and duplicate an- cytology: Associated with T(l;2)l-vl29 =
terior scute liar and postalar bristles; addition of T(1;2)18B;41.
y+Y eliminates wing effect. RK3A.
cytology: Associated with T(l;4)l-vll = M)vU2
I(l)v25 origin: X ray induced.
location: 1- (rearrangement). discoverer: Lindsley, Edington, and Von Halle.
origin: X ray induced. references: 1960, Genetics 45: 1649-70.
discoverer: Lindsley, Edington, and Von Halle. phenotype: Viability of X/Y males 83 percent
references: 1960, Genetics 45: 1649—70. normal. X/Y males fertile. Viability otX/0 males
phenotype: X/Y males of normal viability but less than 1 percent normal. Homozygous females
sterile, owing to failure of sperm head to elongate. viable. RK2A as X/0 male.
X/0 males lethal. RK2A as X/0 males. cytology: Associated with In(l)l-vl32 = ln(l)3-
cytology: Associated with T(l;2)l-v25 = T(l;2)19- 4,19-20.
20;40-41. I(l)vl35
*l(J)v47 origin: X ray induced,
location: 1- (between cv and v). discoverer: Lindsley, Edington, and Von Halle.
discoverer: Lindsley, Edington, and Von Halle. phenotype: Originally recovered as Y-suppressed
references: 1960, Genetics 45: 1649—70. lethal, but in subsequent tests both X/Y and X/0
phenotype: X/Y males have gg-like phenotype but males appeared to be lethal. Later tests by
with peripheral darkening of eye color; viability 41 Thompson show viability of X/Y males to be 40
percent normal; fertile. X/0 males lethal. RK2A. percent normal and X/0 males less than 1 percent
as X/0 male. normal. RK3A.
cytology: Associated with an insertion of an unspec- cytology: Associated with T(l;2)l-vl35 « T(l;2)18-
ified section of heterochromatin into 8F-9B. Link- 19;41. Induced simultaneously with T(2;3)135 =»
age tests suggest second chromosome origin of T(2;3)37;85A, from which it has since separated.
inserted material. T(l;2)l-v47 = T(1;2)8F-9B.
I(l)v59 I(l)vl39
location: 1- (rearrangement). location: 1-2 (between w and spj).
discoverer. Lindsley, Edington, and Von Halle. discoverer: Lindsley, Edington, and Von Halle.
references: 1960, Genetics 45: 1649-70. discoverer: Lindsley, Edington, and Von Halle.
phenotype: X/0 and X/Y males lethal. X/Y/Y references: 1960, Genetics 45: 1649—70.
males viable and fertile; show strong variegation phenotype: Viability of X/Y males normal, but
for w and rst. RK3A. reduced in combination with M(2)S210 but not
cytology: Associated with In(lLR)l-vl39 = E(var)7. X/Y males sterile, owing to failure of
In(lLR)3C6-7. sperm head to elongate. X/0 males lethal. RK2A
other information: Single recombinant carrying distal as X/0 male.
part of .X-centromere-bearing half of T(l;4)wmS = cytology: Associated with T(l;2)l-v219 =
T(l;4)3C3-4;101Fl-2 and proximal part of In(lLR)l- T(l;2)10A;40.
vl39 is variegated for w but not for rst and is *l(l)v223
viable. location: 1- (rearrangement).
*t(l)vl46 origin: X ray induced.
location: 1- (rearrangement). discoverer. Lind^sley, Edington, and Von Halle.
origin: X ray induced. references: I960, Genetics 45: 1649—70.
references: 1960, Genetics 45: 1649-70. normal. X/Y males sterile, owing to variegation for
phenotype: Viability oiX/Y males 41 percent absence of external genitalia, especially in com-
normal; further reduced by M(2)S21 ° and E(var)7. bination with E(var)7. X/0 males lethal. RK2A as
X/Y males fertile. X/0 males lethal. X/0 male.
I(l)vl46/I(l)vl46/Y more viable than cytology: Associated with T(l;2)l-v223 =
I(l)vl46/l(l)vl46 females. Frequently have fewer T(l;2)UF;41;50E.
dorsocentrals. RK2A as X/0 male. I(1)v227
cytology: Associated with In(l)l-vl46 = In(l)5- location: 1- (rearrangement).
6;19-20. origin: X ray induced.
l(l)vl50 discoverer: Lindsley, Edington, and Von Halle.
location: 1- (rearrangement). references: 1960, Genetics 45: 1649-70.
origin: X ray induced. phenotype: Viability of X/Y males 48 percent nor-
discoverer: Lindsley, Edington, and Von Halle. mal; further reduced in combination with M(2)S21 °
references: 1960, Genetics 45: 1649-70. and E(var)7. X/Y males fertile. X/0 males lethal.
phenotype: Viability of X/Y males 15 percent RK2A as X/0 male.
normal. X/Y males sterile, owing to failure of cytology: Associated with In(l)l-v227 = In(l)l-
sperm head to elongate. X/0 males lethal. RK2A 2,19-20.
as X/0 male. I(l)v231
cytology: Associated with T(l;2)l-vl50 = T(l;2)16- location: 1- (rearrangement).
17;40. origin: X ray induced.
discoverer: Lindsley, Edington, and Von Halle.
IO)v163 references: 1960, Genetics 45: 1649—70.
location: 1* (rearrangement). phenotype: X/Y males viable and fertile. Viability
origin: X ray induced. of X/0 males less than 1 percent normal; the few
discoverer: Lindsley, Edington, and Von Halle. survivors have reduced rough eyes. Homozygous
references: 1960, Genetics 45: 1649—70. females normal. RK2A in X/0 male.
phenotype: Viability of X/Y males 17 percent cytology: Associated with In(l)l-v231 =In(l)lC-
normal and of X/0 males less than 1 percent D-,19-20.
normal. X/Y males sterile, owing to failure of *IO)v252
sperm head to elongate. RK2A in X/0 male. location: 1- (rearrangement).
cytology: Associated with T(l;3)l-vl63 » T(1;3)17A- origin: X ray Induced.
B;8O-81. discoverer: Lindsley, Edington, and Von Halle.
*}(l)v216 references: 1960, Genetics 45: 1649-70.
location: 1- (rearrangement). phenotype: Viability of X/Y males normal, of X/0
origin: X ray induced. males 2 percent normal. X/Y males sterile, owing
discoverer: Lindsley, Edington, and Von, Halle. to failure of sperm head to elongate. RK2A as X/0
references: 1960, Genetics 45: 1649-70. male.
phenotype: Viability of X/Y males 15 percent cytology: Associated with T(l;3)l-v252; determined
normal; reduced further in presence of M(2)S2i ° but genetically; cytology not done.
not B(var)7. X/Y males sterile, owing to failure of *l(l)v306
sperm head to elongate. X/0 males lethal. RK2A location: 1-0.
as X/0 male. origin: X ray induced.
cytology: Associated with T(l;2;3)l-v216; deter- discoverer: Lindsley, Edington, and Von Halle.
mined genetically; cytology not done. references: 1960, Genetics 45: 1649—70.
phenotype: Viability of X/Y males 78 percent
location: 1- (rearrangement), normal. X/Y males fertile. X/0 males lethal.
origin: X ray induced. Homozygous females viable. RK2A in X/0 males.
MUTATIONS 127
cytology: Salivary chromosomes show insertion of I(l)v459

material of unknown origin into 1B-E. location: 1- (rearrangement).
t(1)v361 origin: X ray induced.
location: 1- (rearrangement). discoverer: Lindsley, Edington, and Von Halle.
origin: X ray induced. references: I960, Genetics 45: 1649—70.
discoverer: Lindsley, Edington, and Von Halle. phenotype: X/Y males have rough eyes and deformed
references: 1960, Genetics 45: 1649—70. wings and wing veins; 78 percent normal viability;
phenotype: X/Y males show normal viability but are fertile. X/0 males lethal. RK2A as X/0 male.
sterile, owing to failure of sperm head to elongate. cytology: Associated with T(l;2;3)l-v459 =
X/0 males lethal. RK2A as X/0 male. T(l;2;3)3D-F;XR;50;80-81.
cytology: Associated with T(l;3)l-v361 = T(l;3)19- l(J)v463
20;80-8t. location: 1- (rearrangement).
I(l)v451 origin: X ray induced.
location: 1-56.7 (inseparable from f). discoverer: Lindsley, Edington, and Von Halle.
references: 1960, Genetics 45: 1649-70. normal; further reduced in combination with E(var)7
phenotype: Viability of X/Y males 63 percent but not M(2)S2lO. x/Y males sterile, owing to
normal, of X/0 males 4 percent normal. X/Y males failure of sperm head to elongate. Viability of X/0
sterile; spermatogenesis appears normal but sperm males 18 percent normal. RK3A.
not motile. RK3. cytology: Associated with T(l;3)l-v463 = T(l;3)19-
cytology: Salivary chromosomes appear normal. 20-.81-82.
other information: No translocation detectable *l(l)w
genetically. Map distance between v and i reduced location: 1-66.
from standard 23.7 to 12. discoverer: Schubel, 1934.
I(l)v453 phenotype: Males survive; homozygous females die.
location: 1- (rearrangement). RK3.
origin: X ray induced. other information: Probably a lethal allele of bb.
references: 1960, Genetics 45: 1649—70. *I(1)X2: lethol(l) X ray induced
phenotype: Viability of X/Y males normal; reduced location: 1- (near forked).
in combination with M(2)S21 ° but not E(var)7. X/Y origin: X ray induced.
males sterile, owing to failure of sperm head to discoverer: Auerbach.
elongate. X/0 males lethal. RK2A as X/0 male. references: Ede, 1956, Arch. Entwicklungsmech.
cytology: Associated with T(l;3)l-v453 = Organ. 148: 437-51 (fig.).
T(1;3)12D; 80-81. phenotype: Embryos die in advanced stage of devel-
I(l)v454 opment. They live beyond normal hatching time,
location: 1- (rearrangement). move actively, but do not hatch. Embryo distorted;
origin: X ray induced. head material not involuted and pharyngeal material
discoverer: Lindsley, Edington, and Von Halle. external; body wall has disarranged segmentation
references: 1960, Genetics 45: 1649-70. in medial region. Mutant disrupts mechanism con-
phenotype: Viability of X/Y males 50 percent nor- trolling mitosis in early stages of gastrulation,
mal; further reduced in combination with M(2)S210 occasionally as early as blastoderm formation.
and E(var)7. X/Y males sterile, owing to failure of RK2.
sperm head to elongate. X/0 males lethal. RK2A cytology: Salivary chromosomes normal.
in X/0 males.
cytology: Associated with T(l;2;3;4)l-v454 = *IO)X10
T(1;2;3)12B;22-23;81 + T(2;4)44F;t01F. location: 1-0.0 (near <sc).
f(1)v455 discoverer: Auerbach.
location: 1-(rearrangement). references; Ede, 1956, Arch. Entwicklungsmech.
origin: X ray induced. Organ. 149: 247-58 (fig.).
discoverer: Lindsley, Edington, and Von Halle. phenotype: Variation in expression of factors dis-
references: I960, Genetics 45: 1649-70. continuous. There are three types of lethal em-
phenotype: Viability in X/Y males low; further bryos; some may survive into larval stage. Type 1
reduced in presence of both M(2)S210 and E(var)7. stops development after formation of a cap of undif-
X/Y males sterile, owing to failure of sperm head ferentiated cells. Type 2 has limited differentia-
to elongate. X/0 males invisible. Eye color varie- tion, often the nervous tissue exclusively, but no
gated in I(l)v455/w females. RK2A in X/0 males. organ formation. Type 3 survives beyond normal
cytology: Associated with T(l;3)l-v455 « hatching time, has no .gross abnormalities, but does
T(1;3)3C;81. not hatch. RK2.
*l(J)X20
location: 1- (near sc). origin: X ray induced.
origin: X ray induced. discoverer: Abrahamson, 64f5.
references: Ede, 1956, Arch. Entwicklungsraech. origin: X ray induced.
Organ. 149: 101-14 (fig.)- discoverer: Judd, 63g9.
phenotype: Four types of defective embryos pro-
duced. Types 1 and 2 reach late stage of develop- origin: X ray induced,
ment and are alive at time larvae normally hatch. discoverer: Judd, 62gl7.
Type 1 has a complete nervous system, but incom- cytology: Salivary chromosomes appear normal
plete hypoderm. Type 2 has hypoderm but a defi- (Judd).
cient nervous system. Types 3 and 4 stop devel-
oping at early stages. Type 3 has no development origin: X ray induced.
beyond gastrulation, and type 4 forms no blasto- discoverer: Judd, 63gl9.
derm. RK2.
*I(1)X27
origin: X ray induced. discoverer: Judd, 62g26.
references: Ede, 1956, Arch. Entwicklungsmech. origin: X ray induced.
Organ. 149: 88-100 (fig.). discoverer: Judd, 62g31.
phenotype: Embryos alive and in a late stage of de- cytology: Associated with In(l)Uzwl&31 =In(l)3A;6
velopment at normal hatching time but do not hatch. (Judd).
Degeneration begins at about 25 hr. Germ band
irregular at beginning of gastrulation, apparently origin: X ray induced.
the result of defective ventral furrow formation. discoverer: Judd, 62k5.
Consequently, hind-gut is open dorsally, nervous cytology: Salivary chromosomes normal (Judd).
system irregularly developed, and ventral nerve
cord interrupted in region of mid-gut. Other abnor-
malities from different causes are: (1) gut remains origin: X ray induced.
sac like; (2) ectoderm remains unsegmented; and (3) discoverer: Judd, 62k6.
musculature of body wall is underdeveloped. RK2. cytology: Salivary chromosomes normal (Judd).
l(1)zwl°h lethal(l) reste to white origin: X ray induced in z-bearing X chromosome.

location: 1-1.1. discoverer: Judd, 63k26.
origin: X ray induced. I(l)zw2a3
discoverer: Abrahams on, 62al. location: 1-1.2.
cytology: Salivary chromosomes appear normal; origin: X ray induced.
placed in region 3A5-7, on the basis of its inclu- discoverer: Abrahamson, 62a3.
sion in Df(l)64c4 = Dt(l)3A4~6;3C3-5 but not in phenotype: Lethal homozygous and when heterozy-
= Df(l)3A6-8;3Cl-3 (Judd), gous with all other alleles of I(l)zw2 except
\(l)zw2&6. RK2.
origin: X ray induced. cytology: Salivary chromosomes appear normal.
discoverer: Abrahams on, 62a2. Placed in region 3A7-B1, on the basis of its inclu-
sion in Df(l)64j4 = Df(l)3A6-8t*3Bl-2 (Judd).
discoverer: Judd, 62b22. I(l)zw2b26
cytology: Salivary chromosomes normal (Judd). origin: X ray induced.
discoverer: Judd, 62b26.
cytology: Salivary chromosomes appear normal
origin: X ray induced in z-bearing X chromosome. (Judd).
discoverer: Judd, 64d8. I(1)zw2c2i
origin: X ray induced in z-bearing X chromosome. discoverer: Judd, 62c21.
discoverer: Judd, 64dl3. cytology: Salivary chromosomes appear normal
CJudd).
origin: X ray induced, I(1)zw2c28
discoverer: Judd, 63e6. origin: X ray induced,
cytology: Salivary chromosomes normal (Judd). discoverer: Judd, 62c28.
origin: X ray induced, origin: X ray induced,

discoverer: A bra hams on, 64 f2. discoverer: Abrahamson, 64f3.
MUTATIONS 129
I(1)zw2g4 I(l)zw6»2
origin: X ray induced, origin: X ray induced.
discoverer: Lefevre, 62g4. discoverer: Judd, 6212.
cytology: Salivary chromosomes appear normal phenotype: Lethal homozygous and when heterozy-
(Judd). gous with other alleles of t(l)zw6\ only allele of
I(l)zw6 that survives when heterozygous with
origin: X ray induced. Dt(l)62dl8 =Df(l)3B2'Cl;3C3-5 (Judd). RK2.
discoverer: Lefevre, 62g6. cytology: Salivary chromosomes normal (Judd).
phenotype: Lethal homozygous and when heterozy- I(l)zw7*3
gous with all alleles of I(l)zw2 except I(l)zw2a3. location: 1-1.4.
RK2 origin: X ray induced.
discoverer Judd, 63e3.
location: 1-1.3 [based on position of I(l)zw3h22]. cytology: Salivary chromosomes normal. Placed in
origin: X ray induced- region 3B3-C2, on basis of its inclusion in both
discoverer: Judd, 62bl2. Df(l)62dl8 =Df(l)3B2-Cl;3C3-5 and Df(l)wrJ2 =
cytology: Associated with In(l)l-zw3bl2 = ln(l)3A8- Df(l)3A6'8;3Cl'3 (Judd).
Bl;13; placed in region 3A7-B1, on the basis of its
inclusion in Dt(l)64j4 = Df(l)3A6-8;3Bl-2 (Judd). origin: X ray induced.
discoverer: Judd, 63g20.
I(l)zw3h22
origin: X ray induced. location: 1-1.1 [between l(l)zwl and I(l)zw2\.
discoverer: Judd, 62h22. origin: X ray induced.
cytology: Salivary chromosomes normal Qudd). discoverer: Judd, 63glO.
I(l)zw4d28 cytology: Placed in salivary chromosome region
location: 1-1.1 [between l(l)zwl and I(l)zw2\. 3A5-7, on the basis of its inclusion in Df(l)6464 *=
origin: X ray induced. Di(l)3A4-6;3C3-5 but not in Dffijw*"/2 =Df(l)3A6-
discoverer: Judd, 62d28. 8;3Cl-3 (Judd).
cytology: Placed in salivary chromosome region I(l)zw9f4
3A5-7, on the basis of its inclusion in Df(l)64c4 = location: 1-1.4.
Df(l)3A4-6;3C3-5 but not in Df(l)wtJ2 = Df(l)3A6- origin: X ray induced.
8;3Cl-3 (Judd). discoverer: Abrahams on, 64f4.
cytology: Placed in salivary chromosome region
origin: X ray induced. 3B3-C2, on the basis of its inclusion in both
discoverer: Judd, 63e4. Dt(l)62dl8 = Df(l)3B2-Cl;3C3-5 and Df(l)w'J2 =
cytology: Salivary chromosomes normal (Judd). D/jfl^d-S^Cl-;? (Judd).
l(2)39a
I(1)zw4s24 location: 2-50 (right of Bl ?).
origin: X ray induced. origin: Spontaneous,
discoverer: Judd, 62g24. discoverer: Curry, 39a.
location: 1-1.4. l(2)55i
origin: X ray induced. location: 2-55.0 (probably to the left of the centro-
discoverer: Judd, 62jl. mere).
cytology: Placed in salivary chromosome region origin: Spontaneous.
3B3-C2, on the basis of its inclusion in both discoverer. Burdick, 55L
Dt(l)62dl8 =*Df(l)3B2-Cl;3C3-5 and references: 1956, DIS 30: 69.
Df(l)3A6-8;3C1.3 (Judd). Mukai and Burdick, 1959, Genetics 44: 211-32.
1960, Genetics 45: 1581-93.
) Schnick, Mukai, and Burdick, 1960, Genetics 45:
location: 1-1.3. 315-29.
origin: X ray induced, Mukai and Burdick, 1961, Japan. J. Genetics 36:
discoverer: Judd, 62b23. 97-104.
cytology: Salivary chromosomes appear normal phenotype: Larvae hatch but die before pupation.
(Judd). Females heterozygous for l(2)55i have higher
65 fecundity than homorygous wild-type females. The
origin: X ray induced, lethal is therefore not eliminated from laboratory
discoverer: Judd, 63e5. populations. RK3.
l(2)$6a
origin: X ray induced, tocotion: 2-90.
discoverer: Judd, 63el3. origin: Spontaneous.
cytology: Salivary chromosomes apparently normal discoverer: Burdick, 56a.
(Judd). references: 1956, D3S 30: 69.
phenotype: Homozygous lethal; heterozygote shows opening puparium. Occasionally, a fly sponta-
normal viability. RK3. neously escapes puparium without serious injury.
other information: Crossing over normal. Differences in content of free amino acids and pep-
*l(2)57 tides between l(2)bl and wild type can be distin-
origin: Spontaneous. guished in third instar larvae, prepupae, and early
discoverer: Paik. pupae. RK3,
references: I960, Evolution 14: 293-303. l(2)Bld: lethal(2) from Blond
other information: A series of 11 let ha Is selected location: 2-53.1.
from Korean wild populations. origin: Spontaneous.
discoverer: Bridges.
*l(2)1076 l(2)bw: lethald) with brown
location: 2-15 (a,bout 40 units from Bl). location: 2-104.
origin: Spontaneous. origin: Spontaneous in 6w 2b mr chromosome.
discoverer: Ives, 49h. discoverer: Curry, 36i.
references: 1951, DIS 25: 70. cytology: Salivary chromosomes seem to show slight
phenotype: Lethal homozygous and in combination deficiency or disturbance in 59C and D (Bridges).
with ln(2L)Cy. RK3. I(2)C: lethal(2) o/ Curry
*l(2)1323 location: 2-67.1.
location: 2-55 (0/162 crossovers with Bl). origin: Spontaneous.
origin: Spontaneous. discoverer: Curry, 34a21.
discoverer: Ives, 51 g. phenotype: Lethal before pupation. RK3.
references: 1951, DIS 25: 70. cytology: Placed in salivary region 49D4 through
phenotype: Lethal homozygous and in combination 49E5 on the basis of its inclusion in Df(2R)v£B =
with In(2L)Cy + In(2R)Cy. RK3. Df(2R)49D3-4;50A2-3 and in Di(2R)v^ =
K2)a Df(2R)49Cl-2;49E2-6 (Morgan, Bridges, and
location: 2-64.7. Schultz, 1938, Carnegie Inst. Wash. Year Book 37:
origin: Spontaneous. 306).
discoverer Bridges, 16a 15. l(2)cg: lethal(2) with comb gap
'references: Bridges and Morgan, 1919, Carnegie location: 2-15 (between dp and of).
Inst. Wash. Publ. No. 278: 286, 302. origin: Spontaneous.
phenotype: Almost completely lethal; body color of discoverer: Nichols-Skoog, 33dl9.
rare survivor pale. RK3. references: Curry, 1939, DIS 12: 46.
K2)ax 1(2) cn bwco-3a: s e e l(2)S3a
locotion: 2-106.9. 1(2) cn bwco-7; S ee 1(2)S7
origin: Spontaneous. l(2)crc: lethal(2) cryptocephal
discoverer Bridges, 19b28. location: 2-55.
references: 1937, Cytologia (Tokyo), Fujii Jub. Vol. origin: Spontaneous.
2: 745-55. discoverer: Hadorn, 1942.
phenotype: Lethal in very early larval stage. RK3. synonym: crc.
cytology: Located in 60B on salivary chromosome by references: Hadorn and Gloor, 1943, Rev. Suisse
Bridges, but not included in Df(2R)Px = Zool. 50: 256-61.
D£(2R)60B8-10;60Dl-2. Gloor, 1945, Arch. Julius Klaus-Stift.
Way Vererbungsforsch. Sozialanthropol. Rassenhyg.
location: 2-8.3. 20: 209-56.
origin: Spontaneous. Fristom, 1965, Genetics 52: 297-318.
discoverer: Bridges, 30d5. phenotype: Homozygotes undergo pupation but rarely
K2)B eclose from puparia. Imaginal head is not
location: 2- [in 2L of In(2L)t]. everted from thorax. Except for slightly reduced
discoverer Bridges, 1930. eyes and shortened legs, wings, and thoracic
l(2)bh tethaK2) bluter bristles, the head and thorax are fully differen-
location: 2-43.8. tiated. Head eversion is inhibited by integument
origin: X ray induced. being more rigid than normal. Mutant integument
discoverer: Kafer, 50b. contains more glucosamine than normal. Feeding
references: Benz, 1953, DIS 27: 55. glue osa mine to wild-type larvae produces a pheno-
1957, Z. Induktive Abstammungs- Vererbungslehre copy very similar to l(2)crc. Abdomen often shows
88: 78-114 (fig.). no differentiation and internal organ development
phenotype: Lethal at end of pupal stage. Homozy- arrested at pupal stage. RK3.
gotes make emerging movements, but puparia have l(2)gh lethal{2) giant /crvoe
abnormally thick protein layer so that iraaginal location: 2-0.0.
hypodermis is punctured in attempt to eclose. origin: Spontaneous.
Hemoljmph is lost and flies die. Apparently normal discoverer Bridges, 33e9.
hocnoxygotes may be obtained by artificially Synonym: lgl.
MUTATIONS 131
references: Hadorn, 1937, Proc. Soc. Exptl. Biol. origin: Spontaneous.

Med. 36: 632-34. discoverer: Redfield, 23b.
1937, Proc. Natl. Acad. Sci. U.S. 23: 478-84. references: 1924, Am. Naturalist 58: 566—69.
1938, Rev. Suisse Zool. 45: 425-29. 1926, Genetics 11: 482-502.
Vogt, 1947, Z. Naturforsch. 26: 292-94. phenotype: Homozygous females produce one
phenotype: Homozygotes undergo embryogenesis and daughter to 5.5 sons. Abnormal sex ratio caused
three larval ins tars. Larvae reach normal maximum by inviability of females. l(2)mat does not seem to
size but fail to pupate; they then become bloated to be allelic to da, which has a similar effect. RK3.
giant size. Pseudopupae are sometimes formed but l(2)me: lethal(2) meander
no morphogenesis occurs, and imaginal disks location: 2-72 (71-73).
degenerate. Ring gland small and appears immature origin: Spontaneous.
in third-instar larva (Scharrer and Hadorn, 1938, discoverer: Hadorn, 44g20.
Proc. Natl. Acad. Sci. U.S. 24: 236-42). A third- synonym: Itno.
instar l(2)gl host transplanted with a normal ring references: 1947, Exptl. Biol. Symp. Vol. 2:
gland pupates but does not metamorphose. Thus a 177—95, Cambridge Univ. Press.
deficiency of hormones from the ring gland is 1947, DIS 21: 68.
probably one result of l(2)gl but not the only one. Schmid, 1949, Z. Induktive Abstammungs-
Faulhaber (1959, Z. Vererbungslehre 90: 299-334) Vererbungslehre 83: 220-53 (fig.).
finds that the abnormal development affects the Chen and Hadorn, 1954, Rev. Suisse Zool. 61:
quantities of the different amino acids, peptides, 437-51.
and proteins. Welch (1957, Genetics 42: 544—59) 1955, Rev. Suisse Zool. 62: 338—47.
finds that DNA of nuclei in several tissues, espe- phenotype: Larvae do not grow normally; die while
cially salivary glands, is markedly reduced. RK3. small. Body length remains relatively shorter than
cytology: Locus lies between 21A1 and 21C1 tracheal stems, which become convoluted in a
(Lewis, 1945, Genetics 30: 137-66). meandering manner. Salivary glands reach 30 per-
other information: The order of 1(2)0 and net cent normal size; pharyngeal development normal.
unknown. Intestines lack proteolytic enzymes. RK3.
K2)gl2 I(2)mr2: lethal(2) with morula
origin: Ultraviolet induced. location: 2-70.
discoverer: Meyer, 51a. origin: Spontaneous,
references: Meyer and Edmonds on, 1951, DIS 25: 72. discoverer: Bridges, 25k24.
phenotype: Like l(2)gl. RK3. *I(2)NS: lethal(2) Nova Scotia
*I(2)9I3 location: 2-107.0 [to the right of l(2)ax and to the
origin: Spontaneous in In(2L)Cy + In(2R)Cy. left of sp].
discoverer: Meyer, 51a. discoverer: Bridges, 23j31.
references: Meyer and Edmondson, 1951, DIS 25: 73. references: 1937, Cytologia (Tokyo), Fujii Jub. Vol.
phenotype: Larvae heterozygous with l(2)gl2 are like 2: 745-55.
1(2)0. RK3A. phenotype: Lethal when larvae are about 2 mm long.
I(2)H: lethal(2) of Humphrey Development of tracheae and other chitinized parts
location: 2-50. abnormal. RK3A.
origin: Spontaneous. cytology: Exists only as In(2R)NS, px 1(2)NS sp.
discoverer: Humphrey, 32k. Salivary chromosome locus in 60B10-12 on the
references: Dunn, 1934, DIS 1: 30. basis of its inclusion in Df(2R)Px » D%2R)60B8'
1935, DIS 4: 9. 10;60Dl-2 but not in the 2RPXD element of
phenotype: Usually dies as pupa; 10—15 percent of T(l;2)Bld= T(1;2)1C3-4;6OB12-13 (Bridges).
flies survive, look normal but are weak. Homozy- l(2)pm: lethal(2) polymorph
gote usually sterile when inbred but fertile in out- location: 2-30,3.
crosses. RK3. origin: X ray induced,
l(2)hst: lethal(2) histolytic discoverer: Kafer, 50b.
location: 2-56. references: Benz, 1953, DIS 27: 55.
origin: X ray induced. 1957, Z. Induktive Abstammungs- Vererbungslehre
discoverer: Thompson, 59k. 88: 78-114 (fig.).
phenotype: Homozygote dies in early pupal stage. phenotype: Flies die throughout larval and pupal
Heterozygous viability good. RK3. stages. Larvae do not contract before pupation;
I(2)M: lethal(2) from Moftr hence, pupae are long and thin. Imagos often
location: 2-(b«tween dp and 6). cryptocephalic. Chief characteristic is a severe
origin: Spontaneous. muscular dystrophy. Protein metabolism extremely
discoverer: Bridges, 33118. disturbed. In larval stage, free amino acids and
l(2)Mad51"": see 1(2)S one peptidc are in abnormally high concentration.
l(2)Mass38*-: see 1(2)S Prepupae only slightly different from normal in this
t(2)maf: tethal(2) maternal respect. Occasional survivors viable and fertile.
location: 2- (near pr). RK3.
*l(2)pup: lethai(2) pupal *l(2)S3a

location: 2-47. location: 2- (between dp and Sp).
origin: Spontaneous. origin: Gamma ray induced.
discoverer: Ives, 38J25. discoverer: Seto, 1953.
references: 1945, Genetics 30: 175. synonym: l(2)cn bwc°-3a; Co-3A.
1945, DIS 19: 46. references: 1956, J. Heredity 47: 21-27.
phenotype: Dies in middle or late pupal stage. Ex- 1958, DIS 32: 157-58.
ternal anatomy appears normal except for heavy 1961, DIS 35: 94-95.
melanization of wings and legs. RK3. 1963, DIS 37: 128-29.
phenotype: Die predominantly in pupal stage; infre-
I(2)R: lethal(2) of RecHield quent weak sterile survivors with normal pheno-
location: 2- [in 2L with In(2L)t]. type. Death occurs earlier in crowded cultures
discoverer: Redfield, 1933. (Seto, 1957, DIS 31: 160-62). Pupation delayed
*I(2)S1: lethal(2) of Seto 1—2 days beyond normal. Respiration rate during
location: 2- (not located). pupation 50 percent normal (Seto, 1959, DIS 33:
origin: Spontaneous. 159-60). RK3.
discoverer: Seto, 1951. *I(2)S4
synonym: itfyMadSln-1; N-l. location: 2- (near pr).
references: 1954, J. Exptl. Zool. 126: 17-32. origin: Spontaneous.
1954, Am. Naturalist 88: 373-78. discoverer: Seto, 1951.
1956, J. Heredity, 47: 21-27 (fig.). synonym: l(2)Mad51"-4; N-4.
1961, DIS 35: 94-95. references: 1954, J. Exptl. Zool. 126: 17-32.
phenotype: Dies as larva or prepupa. Puparium 1954, Am. Naturalist 88: 373-78.
elongated, often with larval segmentation, en- 1956, J. Heredity 47: 21-27 (fig.).
larged, resembling 1(2)0.. Cephalic complex un- 1958, DIS 32: 157-58.
everted. Eye rudiment often pigmented. RK3. 1963, DIS 37: 128-29.
*I(2)S1A phenotype: Dies in prepupal and pupal stages.
location: 2- (between Sp and 6). Puparium small and slender with fragile exoskel-
origin: Spontaneous. eion. Appearance like l(2)crc, with uneverted
discoverer: Seto, 1953. cephalic complex. Development of hypodermal
synonym: l(2)Mad53n-tA', N-1A. derivatives and pigment variable. Pupation delayed
references: 1956, J. Heredity 47: 21-27 (fig.). 1—2 days. Crowding suppresses expression (Seto,
1958, DIS 32: 157-58. 1957, DIS 31: 160-62). RK3.
1961, DB 35: 94-95. *I(2)S7
1963, DIS 37: 128-29. location: 2- (between Sp and b).
phenotype: Dies in late pupa; differentiation of ex- origin: Gamma ray induced.
ternal structures almost complete. Color of pupa discoverer: Seto, 1953.
darker, changing to deep brown after a few days; synonym: l(2)cn bwc°-7; Co-7.
heavy melanization in thicker parts of exoskeleton; references: 1956, J. Heredity, 47: 21-27.
nephrocytes deeply pigmented. Respiration rate 80 1957, DIS 31: 160-62.
percent normal (Seto, 1959, DIS 33: 159—60). 1958, DIS 32: 157-58.
Slight delay in pupation. Development ceases 1961, DIS 35: 94-95.
earlier under crowded conditions (Seto, 1957, DIS phenotype: Die predominantly in pupal stage. Occa-
31: 160-62). RK3. sional sterile adults produced. Time of pupation
*l(2)$3 delayed. Respiration rate of prepupae and pupae 50
location: 2-(between c and px). percent normal (Seto, 1959, DIS 33: 159-60). RK3.
origin: X ray induced. *K2)sn
discoverer: Seto, 1951. location: 2- (not located).
synonym: l(2)MasB38x-3; X-3. origin: X ray induced.
references: 1954, J. Exptl. ZooL 126: 17-32. discoverer: Seto, 1951.
1956, J. Heredity 47: 21-27 (fig.). synonym: itfytiassSS*-11; X-ll.
1957, DIS 31: 160-62. references: 1954, J. Exptl. Zool. 126: 17-32.
1958, DIS 32: 157-58. 1956, J. Heredity 47: 21-27.
1961, DIS 35: 94-95.
1961, DIS 35: 94-95.
1963, DJS 37: 128-29.
phenotype: Dies as pupa. Puparium normal. Devel-
phenotype: Dies in prepupal stage. Puparium small opment may cease before or shortly after eversion
and dumpy. Deformed prepupa only partly fills
of frontal sac. Usually has melanized patches on
pupariswj; anterior structures rudimentary; rest of
dorsum. Wings and legs variably developed. RK3.
body saclike. Scattered small pigmented or
meiamijwd art as along tracrteal trunks; appendages location: 2- (not located).
and cephalic complex reduced or absent. Some origin: Spontaneous,
delay in pupation. RK3. discoverer. Seto, 1955.
MUTATIONS 133
synonym: l(2)Wau55n-l3; N-13. condition. Most flies complete but only rarely able
references: 1957, DIS 31: 160-62. to eclose; rare adults are weak and unproductive.
1961, DIS 35: 94-95. Time of pupation delayed about one-half day. RK3.
phenotype: Dies as late pupa or as adult. Puparium *l(2)S50
normal. RK3. location: 2- (rearrangement).
*I(2)S32 origin: Spontaneous.
location: 2- (between dp and Sp), discoverer: Seto, 1951.
origin: Spontaneous. synonym: l(2)Mad51n"S0; N-50.
discoverer: Seto, 1951. references: 1954, J. Exptl. Zool. 126: 17-32.
synonym: l(2)Mad51n-32; pj-32. 1954, Am. Naturalist 88: 373-78.
references: 1954, J. Exptl. Zool. 126: 17-32. 1956, J. Heredity 47: 21-27 (fig.).
1954, Am. Naturalist 88: 373-78. 1957, DIS 31: 160-62.
1956, J. Heredity 47: 21-27 (fig.). 1958, DIS 32: 157-58.
1958, DIS 32: 157-58. 1961, DIS 35: 94-95.
1961, DIS 35: 94-95. phenotype: Dies as pupa. Puparium normal. Devel-
1963, DIS 37: 128-29. opment ceases in early pupa; body generally unpig-
phenotype: Dies in early pupa. Usually no pigment mented; eyes rarely pigmented; hypodermal deriva-
or bristle formation; leg and wing sacs adhere to tives underdeveloped; melanotic degeneration of
pupa case, resulting in appendages developing in hypodermis in region of eye, external genitalia, and
cramped position; some melanization at extremi- appendage extremities. Pupation delayed. RK3A.
ties. Pupation delayed 1 day; pupa badly shrunken. cytology: Associated with In(2L) and In(2R) with
Crowding suppresses expression (Seto, 1957, DIS unknown break points.
31: 160—62). Respiration rate 50 percent normal *I(2)S51
(Seto, 1959, DIS 33: 159-60). RK3. location: 2- (near pr).
*I(2)S42 origin: Spontaneous.
location: 2- (between Bl and L). discoverer: Seto, 1951.
origin: Spontaneous. synonym: l(2)Mad51n-sl; N-Sl.
discoverer: Seto, 1951. references: 1954, J. Exptl. Zool. 126: 17-32.
synonym: l(2)Mad51n-42; N-42. 1954, Am. Naturalist 88: 373-78.
references: 1954, J. Exptl. Zool. 126: 17-32. 1956, J. Heredity 47: 21-27.
1956, J. Heredity 47: 21-27. 1957, DIS 31: 160-62.
1958, DIS 32: 157-58. 1958, DIS 32: 157-58.
1961, DIS 35: 94-95. 1961, DIS 35: 94-95.
phenotype: Dies in late pupa. Appears well differen- phenotype: Dies in prepupal stage. Puparium
tiated externally; eyes with little or no pigment; normal. Development similar to that of 1(2)S61, but
internal head structures poorly developed. RK3. with certain tissues further developed. Pupation
*I(2)S42* delayed 1—2 days. Respiration less than 50 per-
origin: Spontaneous derivative of 1(2)S42. cent normal (Seto, 1959, DIS 33: 159-60). RK3.
discoverer: Seto, 1954. *I(2)SSS
synonym: l(2)Mad51n-<2a; N-42A. location: 2- (between dp and Sp).
references: 1956, J. Heredity 47: 21-27 (fig.). origin: Spontaneous.
1957, DIS 31: 160-62. discoverer: Seto, 1955.
1961, DIS 35: 94-95. synonym: l(2)Wau55nS*; N-55.
phenotype: Ceases development in early pupa; references: 1958, DIS 32: 157-58.
puparium enlarged, elongated, and larva like; exo- 1961, DIS 35: 94-95.
skeleton thin and fragile; pupa adheres to anterior phenotype: Dies in late pupa; occasional weak adult
end of puparium, res-t of pupa contracted anteriorly. survivors. Phenotype similar to 1(2)S45. Crowding
Space between pupa and puparium filled with light, results in more and earlier mortality (Seto, 1957,
oily fluid. Time of pupation delayed 2—3 days. DIS 31: 160—62). Pupation delayed about one-half
Imaginal disks poorly developed. RK3. day. RK3.
*I(2)U5 *I(2)$S9
location: 2- (between Sp and o). location: 2- (not located).
discoverer: Seto, 1953. discoverer: Seto, 1951.
synonym: l(2)Mad53n-*3; N-45. synonym: l(2)Mad51n'^9; N-59.
references: 1956, J. Heredity 47: 21-27 (fig.). references: 1954, J. Exptl. Zool. 126: 17-32.
1957, DIS 31: 160-62. 1956, J, Heredity 47: 21-27.
1958, DBS 32: 157-58. 1961, D^ 35: 94-95.
1961, DIS 35: 94-95. phenotype: Dies in late larval or prepupal stage.
phenotype: Dies as late pupa just before eclosion. Puparium normal; frontal sacs uneverted; leg and
Slightly smaller than normal; leg sacs do not wing sacs incompletely developed; body sac like.
elongate, end legs develop in cramped and stunted RK3.
*I(2)S61 *l(2)SP9c
location: 2- (near pr). location: 2-55.1 (between rl and stw).
discoverer: Seto, 1951. discoverer: Spiess.
synonym: l(2)Mad51n-6l; N-61. references: Spiess, Helling, and Capenos, 1963,
references: 1954, J. Exptl. Zool. 126: 17-32. Genetics 48: 1377-88.
1954, Am. Naturalist 88: 373-78. phenotype: Lethal homozygous and in combination
1956, J. Heredity 47: 21-27 (fig.). with M(2)S2; viable in combination with l(2)Spll
1957, DIS 31: 160-62. and 1(2)SP15. RK3.
1958, DIS 32: 157-58. cytology: Placed in salivary region 41A based on
1961, DIS 35: 94-95. its inclusion in D{(2R)M'S21 ° = Df(2R)41A
1963, DIS 37: 128-29. (Burdick).
phenotype: Dies in prepupal stage. Puparium color *l(2)SP9d
darker than normal. Prepupa incompletely de- location: 2-55.1 (to the right of stw).
veloped; frontal sacs uneverted; wing and leg sacs origin: Spontaneous.
everted but development curtailed; free-floating fat discoverer: Spiess.
body fragments may fill fluid space between pupa- references: Spiess, Helling, and Capenos, 1963,
rium and prepupa. Pupation delayed 1—2 days. Genetics 48: 1377-88.
Respiration rate normal for first day and a half *l(2)SP10
then ceases (Seto,1959, DIS 33: 159-60). RK3. location: 2-37.5.
*l(2)Sph leihol(2) of Spiess origin: Spontaneous.
location: 2-35.0. discoverer: Spiess.
origin: Spontaneous. references: Spiess, Helling, and Capenos, 1963,
discoverer: Spiess. Genetics 48: 1377-88.
references: Spiess, Helling, and Capenos, 1963,
Genetics 48: 1377-88.
K2)sPn
location: 2-55.1 [between rl and stw; to the left of
*l(2)SP2b
1(2)SP15 (Burdick)].
location: 2-49.
discoverer: Spiess.
discoverer: Spiess.
references: Spiess, Helling, and Capenos, 1963,
references: Spiess, Helling, and Capenos, 1963, Genetics 48: 1377-88.
Genetics 48: 1377-88. phenotype: Lethal homozygous and in combination
*l(2)SP6b with M(2)S2; viable in combination with l(2)Sp9c
location: 2-50.0. and I(2)apl5. RK3.
origin: Spontaneous. cytology: Placed in salivary region 41A based on
discoverer. Spiess. its inclusion in Df(2RyM-S2^ 0 = Df(2R)41A
references: Spiess, Helling, and Capenos, 1963, (Burdick).
Genetics 48: 1377-88. *l(2)SpU
I(2)SP7 location: 2-61.5.
origin: Spontaneous, discoverer: Spiess.
discoverer: Spiess. references: Spiess, Helling, and Capenos, 1963,
Genetics 48: 1377-88. *l(2)SpU
*l(2)Sp8 location: 2-32.0.
origin: Spontaneous. discoverer: Spiess.
discoverer: Spiess. references: Spiess, Helling, and Capenos, 1963,
Genetics 48: 1377-88.
*!(2)Sp9a i(2)$P15
location: 2-1.9. location: 2-55.1 (between rl and stw).
discoverer Spiess. discoverer: Spiess.
references: Spiess, Helling, and Capenos, 1963, references: Spiess, Helling, and Capenos, 1963,
Genetics 48: 1377-88. Genetics 48: 1377-88.
l(2)Sp% phenotype: Lethal homozygous and in combination
location: 2-49. with M(2)S2; viable in combination with l(2)Sp9c
origin: Spontaneous. and l(2)Spll. RK3.
dl*cov»r#r. Spiess. cytology: Placed in salivary region 41A based on
r*f*r«nc«s: Spiess, Helling, and Capeno«, 1963, its inclusion in Df(2R.)M-S210 « Dt(2R)41A
Getseiics 48: 1377-8S. (Burdick).
MUTATIONS 135
*I(2)SP18 other information: Results of Bridges interpreted to

location: 2-65.3. show allelism to M(3)wB (3-79.7); may have been
origin: Spontaneous. related to maternal effect described by Schultz.
discoverer: Spiess. Position based on crosses by Muller (1918), in
references: Spiess, Helling, and Capenos, 1963, which he used l(3)a separated from In(3R)C.
Genetics 48: 1377-88. l(3)ac: lethal(3) accessory
l(2)Su(H): lethal(2) from Suppressor of Hairless location: 3- (midregion).
location: 2-99. discoverer: Schultz, 25g.
origin: Spontaneous. phenotype: Enhances maternal effect of In(3R)C,
discoverer: Bridges, 3717. l(3)a on recovery of M(3)w daughters from
cytology: Located in salivary region 58A1 through M(3)w/In(3R)C, l(3)a mothers (Schultz). RK3.
58F8 on basis of its inclusion in Df(2R)M-l = *l(3)blo-l: lethal(3) bloated larvae
Df(2R)57Fll-58Al;58F8-S9Al. location: 3- (to the left of p).
*I(2)T: lethal(2) of Thompson discoverer Bridges, 25k7.
origin: Spontaneous in normal chromosome of SM1/+ references: Chen, 1929, J. Morphol. 47: 135-99.
heterozygote. phenotype: Larvae become very large and trans-
discoverer: Thompson, 1956, 1957. parent; die in the prepupal stage. Growth of
synonym: I(2)56i24 through l(2)57hlO. imaginal disks irregular. RK2.
other information: A series of 13 independently 1(3)D1: see DP
occurring and genetically located lethals. *l(3)e: lethal(3) with ebony
l(2)Wau55n-: see 1(2)S location: 3- (not located).
1(3)1: see l(3)a origin: Spontaneous in In(3R)C, e.
1(3)26 discoverer: Schultz.
location: 3-52.2 [between l(3)S6 and 1(3)S7\. phenotype: Dies as fully developed normal-appearing
origin: X ray induced in a cu kat chromosome simul- imago unable to eclose. RK3A.
taneously with ry26.
discoverer: Schalet. *l(3)hd: lethal(3) head defect
synonym: 1(3)52.5.2. location: 3- (not located).
references: Schalet, Kernaghan, and Chovnick, discoverer: Bridges, 1924.
1964, Genetics 50: 1261-68. references: Morgan, Bridges, and Sturtevant, 1925,
other information: Placed between 1(3)S6 and 1(3)S7', Bibliog. Genet. 2: 230.
on the basis of its inclusion in Df(3R)ryK, phenotype: Dies in pupal stage with black tumorlike
Dt(3R)tvSi, and Df(3R)ry66 but not in Df(3R)ry2*', growth in head. RK3.
Df(3R)ry33, Df(3R)ry*2, or Df(3R)ry70f a l l of w hich I(3)PL
include 1(3)S6 but none of which include 1(3)S7. location: 3- (left arm).
l(3)36d!0 origin: Spontaneous in 3L carrying Itv(3L)P.
location: 3- (close to D, or rearrangement). I(3)PR
origin: Spontaneous, location: 3-90.2.
discoverer: Bridges, 36dlO. origin: Spontaneous inIn(3R)P.
*l(3)36d24
phenotype: Homozygous lethal; lethal in combination
location: 3- (near centromere),
with M(3)j. RK3.
K3)S1: lethal(3) of Schalet
location: 3-51 (to the left of kar),
origin: X ray induced in a kar3 chromosome-
Bridges and Bridges, 1938, Genetics 23: 111-14.
discoverer Schalet.
1(3)52.52: see 1(3)26
other information: Placed to the left of l(3)S2, on
*l(3)62g
the basis of its exclusion from Dt(3R)ry76, which
is deficient for 1(3)S2 and loci to the right.
discoverer: Paik.
l(3)Slo
origin: X ray induced in a kar2 chromosome.
Vol. 1: 163-64.
discoverer: Schalet.
other information: A series of 65 lethals recovered
other information: Allelism with 1(3)S1 tentative and
from Korean wild populations.
K3)a based on similarity in interaction with D£(3R)ry7*.
location: 3-81.6. I(3)S2
origin: Spontaneous in In(3R)C location: 3-51.5 [between l(3)Sl and kar].
discoverer: Morgan, 111. origin: X ray induced in a kar2 chromosome.
synonym: 1(3)1. discoverer: Schalet.
references: Muller, 1918, Genetics 3: 422-99. references: Schalet, Kernaghan, and Chovnick,
phenotype: Lethal homozygou®. Reduces recovery of 1964, Genetics 50: 1261-68.
M(3)w-be&ring daughters from In(3R}C, l(3)s/M(3)w other information: Placed between l(3)St and kmr, on
females (Schultx). RK3. the basis of its inclusion in Df(3R)ty76 but not
D%3R)ry29, D%3R)ry33, or Df(3R)ry36. None of
these deficiencies include 1(3)S1 and all include other information: Produces local reduction in
/car and genes to its right. crossing over.
I(3)S3 I(3)S7
location: 3-51.7 (between kar and mes). location: 3-53 [to the right of 1(3)26].
origin: X ray induced in a kar2 chromosome. origin: X ray induced in a kar2 chromosome.
discoverer: Schalet. discoverer: Schalet.
other information: Placed between /car and mes, on references: Schalet, Kernaghan, and Chovnick,
the basis of its exclusion from Df(3R)ry78 and its 1964, Genetics 50: 1261-68.
inclusion in Df(3R)ry27, Df(3R)ryS2, Df(3R)ry7S, other information: Placed to the right of 1(3)26* on
and Df(3R)ry77. None of these deficiencies in- the basis of its exclusion from Df(3R)ry66, which
clude /car and all include mes and loci to its right. includes 1(3)26 and loci to its left.
I(3)S4 l(3)S7o
location: 3-52.1 (to the right of pic). origin: X ray induced in a kar2 chromosome.
origin: X ray induced in a kar2 chromosome. discoverer: Schalet.
discoverer: Schalet. references: Schalet, Kernaghan, and Chovnick,
references: Schalet, Kernaghan, and Chovnick, 1964, Genetics 50: 1261-68.
1964, Genetics 50: 1261-68. other information: Extends farther to the right than
phenotype: Homozygous lethal, but there a few rela- l(3)S7 since it is lethal in combination with an
tively normal-appearing survivors that are mostly undescribed deficiency for red with which l(3)S7
females. RK3. survives.
other information: Placed to the right of ry by re- l(3)S7b
combination and to the right of pic on basis of its origin: X ray induced in a kar2 chromosome.
survival in combination with ry3$ which behaves discoverer: Schalet.
as though it were deficient for ry and pic. Placed references: Schalet, Kernaghan, and Chovnick,
to the left of 1(3)S5 by recombination. 1964, Genetics 50: 1261-68.
I(3)S5 other information: Extends farther to the right than
location: 3-52.1 [between 1(3)S4 and 1(3)S6]. 1(3)S7, by same criterion as l(3)S7a.
origin: X ray induced in a kar2 chromosome. *l(3)S7c
discoverer: Schalet. origin: X ray induced in a kar2 chromosome.
references: Schalet, Kernaghan, and Chovnick, discoverer: Schalet.
1964, Genetics 50: 1261-68. references: Schalet, Kernaghan, arid Chovnick,
other information: Placed to the right of 1(3)S4 by 1964, Genetics 50: 1261-68.
recombination analysis and to the left of l(3)5>6, on *l(3)S7d
the basis of its inclusion in Df(3R)ry75 and origin: X ray induced in kar2 chromosome.
Dt(3R)ry76, which do not include 1(3)S6. discoverer: Schalet.
l(3)S5o references: Schalet, Kernaghan, and Chovnick,
origin: X ray induced in a kar2 chromosome. 1964, Genetics 50: 1261-68.
references: Schalet, Kernaghan, and Chovnick, *l(3)Sph !ethal(3) of Spiess
1964, Genetics 50: 1261-68. location: 3-33.8.
I(3)S6 origin: Spontaneous.
location: 3-52.2 [between 1(3)S5 and 1(3)26]. discoverer: Spiess.
origin: X ray induced in a kar3 chromosome. references: Spiess, Helling, and Capenos, 1963,
discoverer: Schalet. Genetics 48: 1377-88,
references: Schalet, Kernaghan, and Chovnick, *I(3)SP2
1964, Genetics 50: 1261-68. location: 3-79.3.
other information: Placed to the right of 1(3)S5, on origin: Spontaneous.
the basis of its exclusion from Df(3R)ry7S and discoverer: Spiess.
D%3R)ry76, both of which include I(3)S5 and loci references: Spiess, Helling, and Capenos, 1963,
to the left. Genetics 48: 1377-88.
I(3)S6° *K3)$p5
origin: X ray induced in a kar2 chromosome. location: 3-41.0.
discoverer: Schalet. origin: Spontaneous,
references: Schalet, Kernaghan, and Chovnick, discoverer: Spiess.
1964, Genetics 50: 1261-68. references: Spiess, Helling, and Capenos, 1963,
other information: Causes local reduction in Genetics 48: 1377-88.
crossing over. *I(3)SP6
location: 3-40.4.
origin: X ray induced in a kar2 chromosome, origin: Spontaneous.
discoverer: Schalet. discoverer: Spiess.
references: Schalet, Kernaghan, and Chovnick, references: Spiess, Helling, and Capenos, 1963,
1964, Genetics 50: 1261-68. Genetics 48: 1377-88.
MUTATIONS 137
*I(3)SP9 phenotype: Lethal homozygous and in combination

location: 3-101.1. with Df(4)M. No interaction with other genes in
origin: Spontaneous. Df(4)M. Dies as larva. RK3.
discoverer: Spiess. cytology: Placed in region 101E through 102B16, on
references: Spiess, Helling, and Capenos, 1963, basis of its inclusion in Df(4)M =
Genetics 48: 1377-88. Df(4)101E-F;102B6-17.
*l(3)SP10 I (4) Jo
origin: Spontaneous, discoverer: Gloor and Green, 1957.
discoverer: Spiess. synonym: 1(4)27.
references: Spiess, Helling, and Capenos, 1963, references: Hochman, Gloor, and Green, 1964,
Genetics 48: 1377-88. Genetica 35: 109—26.
*I(3)SP17 phenotype: Like 1(4)1. RK3.
location: 3-38.4. i(4)n
discoverer: Spiess. discoverer: Gloor and Green, 1957.
references: Spiess, Helling, and Capenos, 1963, synonym: 1(4)28.
Genetics 48: 1377-88. references: Hochman, Gloor, and Green, 1964,
*I(3)SP19 Genetica 35: 109-26.
location: 3-100.9. phenotype: Like 1(4)1. RK3.
origin: Spontaneous. 1(4)1 c
discoverer. Spiess. origin: X ray induced.
references: Spiess, Helling, and Capenos, 1963, discoverer: Gloor and Green, 1957.
Genetics 48: 1377-88. synonym: 1(4)32.
l(3)tr: lethal(3) translucida references: Hochman, Gloor, and Green, 1964,
location: 3-20 (18.1 to 22.0). Genetica 35: 109-26.
origin: Spontaneous. phenotype: Like 1(4)1. RK3.
discoverer: Hadorn, 40116. 1(4)1 d
references: 1947, Exptl. Biol. Symp. Vol. 2: origin: X ray induced.
177-95, Cambridge Univ. Press. discoverer: Gloor and Green, 1957.
1947, DIS 21: 68. synonym: 1(4)30.
1956, Cold Spring Harbor Symp. Quant. Biol. 21: references: Hochman, Gloor, and Green, 1964,
363-73 (fig.). Genetica 35: 109-26.
phenotype: Larvae become bloated and transparent phenotype: Like 1(4)1. RK3.
from accumulation of abnormal amount of hemo- other information: Incorrectly reported as an allele
lymph. Concentration of amino acids in hemolymph of 1(4)25 by Hochman, Gloor, and Green (1964).
higher than normal; concentration of proteins 1(4)1 -JFC: see 1(4)9°
reduced. Pupation delayed one day (25°C); dwarfed 1(4)2
pupae formed in inflated puparia; death follows location: 4-(not located),
pupation or completion of imaginal differentiation origin: X ray induced.
of head and thorax; abdomen never metamorphoses. discoverer: Gloor and Green, 1957.
After transplantation into normal hosts, imaginal references: Hochman, Gloor, and Green, 1964,
disks develop normally; ovaries also develop Genetica 35: 109—26.
normally and are fully capable of producing viable phenotype: Lethal in embryo. Lethal in combination
eggs [Sobels, 1950, Experientia 6: 139-40 (fig.)]. with the so-called btD fourth chromosome. RK3.
In pure oxygen, frequency and extent of imaginal 1(4)2"
differentiation strongly increased [Sobels and origin: X ray induced.
Nijenhuis, 1953, Z. Induktive Abstammungs- discoverer: Gloor and Green, 1957.
Vererbungslehre 85: 579-92 (fig.)]. RK3. synonym: 1(4)21.
cytology: Salivary chromosomes normal (Rosin). references: Hochman, Gloor, and Green, 1964,
Genetica 35: 109-26.
location: 3- (right.arm). phenotype: Like 1(4)2. RK3.
origin: Spontaneous in 3R carrying In(3R)P. l(4)2b
l(3)XaR origin: X ray induced.
location: 3-91.8. discoverer: Gloor and Green, 1957.
other information: Used to balance T(2;3)®px*. synonym: 1(4)23.
1(4)1 references: Hochman, Gloor, and Green, 1964,
location: 4- [within Df(4)Ml Genetica 35: 109-26.
origin: X ray induced. phenotype: Unlike other alleles, dies as larva.
discoverer: Gloor and Green, 1957. l(4)2b/I(4)2h heterozygotes survive and have
references: Hochman, Gloor, and Green, 1964, narrow bodies both as pupae and imagos. Viability
Genetica 35: 109-26. reduced; fertile. RK3.
GENETIC VARIATIONS OF OROSOPHILA MELANOGASTEfi
138
I(4)2'>*D: leihal(4) 2 in bent Dominant 1(4)4"

origin: Spontaneous. Associated with supposedly origin: X ray induced.
btD chromosomes. discoverer. Gloor and Green, 1957.
references: Fox, 1947, DIS 21: 85. synonym: 1(4)18.
Hochman, Gloor, and Green, 1964, Genetics 35: references: Hochman, Gloor, and Green, 1964,
109-26. Genetica 35: 109-26.
phenotype: Lethal homozygous and in combination phenotype: Like 1(4)4. RK3.
with other 1(4)2 alleles. RK3. l(4)4b
1(4)2* origin: X ray induced.
origin: X ray induced. discoverer: Gloor and Green, 1957.
discoverer: Gloor and Green, 1957. synonym: 1(4)20.
synonym: 1(4)29. references: Hochman, Gloor, and Green, 1964,
references: Hochman, Gloor, and Green, 1964, Genetica 35: 109-26.
Genetica 35: 109-26. phenotype: Like 1(4)4. RK3.
phenotype: Like 1(4)2. RK3. l(4)4c
l(4)2d origin: Spontaneous.
origin: X ray induced. discoverer: Hochman, 61a.
discoverer. Gloor and Green, 1957. synonym: l(4)PT-3: lethal(4) Powell, Tennessee-3.
synonym: 1(4)37. references: 1961, Am. Naturalist 95: 375-82.
references: Hochman, Gloor, and Green, 1964, 1963, DIS 37: 45.
Genetica 35: 109-26. Hochman, Gloor, and Green, 1964, Genetica 35:
phenotype: Like 1(4)2. RK3. 109-26.
1(4)2* phenotype: Like 1(4)4. RK3.
origin: X ray induced. l(4)4d
discoverer. Gloor and Green, 1957. origin: Spontaneous.
synonym: 1(4)38. discoverer Lipe, 62k.
references: Hochman, Gloor, and Green, 1964, synonym: l(4)AM-2: lethal(4) Amherst, Massachu-
Genetica 35: 109-26. setts-2.
phenotype: Like 1(4)2. RK3. references: Hochman, Gloor, and Green, 1964,
1(4)2* Genetica 35: 109-26.
origin: Spontaneous. phenotype: Like 1(4)4. RK3.
discoverer Hochman, 61a.
synonym: l(4)PT-2: IeOial(4) Powell, Tennessee-2. 1(4)5
references: 1961, Am. Naturalist 95: 375-82. location: 4- (not located).
1963, DIS 37: 48. origin: X ray induced.
Hochman, Gloor, and Green, 1964, Genetica 35: discoverer: Gloor and Green, 1957.
109-26. references: Hochman, Gloor, and Green, 1964,
phenotype: Like 1(4)2. RK3. Genetica 35: 109-26.
1(4)2* phenotype: Homozygotes die as embryos. RK3.
origin: Spontaneous. 1(4)6
discoverer; Hochman, 63kl9. location: 4- (not located).
pti«rtofyp«: Like 1(4)2. RK3. origin: X ray induced.
discoverer: Gloor and Green, 1957.
<w!§i«s Spontaneous. references: Hochman, Gloor, and Green, 1964,
discoverer: Hochman, 63122. Genetica 35: 109-26.
•Imnetyp'e: Lethal &a embryo. In combination with phenotype: Homozygotes die as larvae. RK3.
l(4)2b, produce* a few survivors that have narrow 1(4)6°
bodies both as pupae ami imagos and are fertile. origin: X ray induced.
synonym: 1(4)36.
oriigin: 1 ray induced. references: Hochman, Gloor, and Green, 1964,
l i e o v m r : Gloor an-d Gr®en, 1947.
Genetica 35: 109—26.
sytKMtym: t{4)34,
phenotype: Homozygotes die as pupae. RK3.
references: Hochm&n, Gloor, aad Green, 1964,
l(4}6t>
Ge»@tfca 35: 109-26.
toe«fi«n: 4- (sot located). discoverer: Hochman, 61a.
•riffn: X ny induced, synonym: 1(4)PT-1; lethal(4) Powell, Tennessee-1.
disc»v««*r: Gloor tad Green, 1957. references: 1961, Am. Naturalist 95: 375—82.
1963, DIS 37: 48.
<wf«nmc«s: Hodman, Gloor, aed Green, 1964,
Hochman, Gloor, and Green, 1964, Genetica 35:
GHMKIC* 35: 109-26.
109-26.
it#« di® as larvae. RK3.
phenotype: Homozygotes die as pupae. RK3.
MUTATIONS 139
*1(4)6 c phenotype. Thus, whereas the at locus is within

origin: X ray induced, Df(4)M, 1(4)9 b is not.
discoverer: Gloor and Green, 1957. 1(4)9-
synonym: 1(4)15. origin: Spontaneous.
references: Hochman, Gloor, and Green, 1964, discoverer Hochman, 64a4.
1(4)7 1(4)10
location: 4- (not located). location: 4- (not located).
discoverer: Gloor and Green, 1957. discoverer: Gloor and Green, 1957.
references: Hochman, Gloor, and Green, 1964, synonym: 1(4)33.
Genetica 35: 109-26. references: Hochman, Gloor, and Green, 1964,
phenotype: Homozygotes die as embryos. RK3. Genetica 35: 109—26.
1(4)8 1(4)10: see 1(4)8*
location: 4- (not located). 1(4)11: see Di(4)ll
origin: X ray induced. 1(4)12: see Df(4)12
discoverer: Gloor and Green, 1957. 1(4)13
references: Hochman, Gloor, and Green, 1964, location: 4- [within Df(4)M\.
Genetica 35: 109—26* origin: X ray induced.
phenotype: Homozygotes die as pupae. RK3. discoverer: Gloor and Green, 1957.
*l(4)8° references: Hochman, Gloor, and Green, 1964,
origin: X ray induced. Genetica 35: 109-26.
discoverer: Gloor and Green, 1957. phenotype: Homozygous lethal. Lethal in combina-
synonym: 1(4)10. tion with Df(4)M and ciD but not with Df(4)17,
references: Hochman, Gloor, and Green, 1964, Df(4)34, or 1(4)18. 1(4)13/+ is normal. RK3.
Genetica 35: 109—26. cytology: Placed in region 101E through 102B16, on
phenotype: Homozygous lethal. RK3. basis of its inclusion in Dt(4)M =
Df(4)101E-F;102B6-17.
origin: X ray induced. 1(4)14
discoverer: Gloor and Green, 1957. location: 4- (not located).
synonym: 1(4)19. origin: X ray induced.
references: Hochman, Gloor, and Green, 1964, discoverer: Gloor and Green, 1957.
Genetica 35: 109-26. references: Hochman, Gloor, and Green, 1964,
phenotype: Like 1(4)8. RK3. Genetica 35: 109-26.
1(4)9 phenotype: Homozygotes die as larvae. RK3.
location: 4- [within Df(4)ti\. 1(4)14"
discoverer: Stowell, 62k. discoverer. Gloor and Green, 1957.
synonym: 1(4)BU-1: lethal(4) Bountiful, Utah-1. synonym: 1(4)26.
references: Hochman, Gloor, and Green, 1964, references: Hochman, Gloor, and Green, 1964,
Genetica 35: 109-26. Genetica 35: 109-26.
phenotype: Lethal homozygous and when heterozy- phenotype: Like 1(4)14. RK3.
gous with D£(4)3, Di(4)ll, Df(4)12, Dt(4)24, 1(4)14*
Df(4)34, Df(4)G, and spa c «<. RK3. origin: Spontaneous.
* 1(4)9° discoverer: Wrathall, 611.
origin: Spontaneous. synonym: l(4)ST-2: lethal(4) Solway, Tennessee-2.
discoverer: Crow. references: Hochman, 1963, DIS 37: 48.
synonym: 1(4)1-JFC: lethal(4) of /. F. Crow. Hochman, Gloor, and Green, 1964, Genetica 35:
references: Hochman, Gloor, and Green, 1964, 109-26.
phenotype: Like 1(4)9. RK3. 1(4)15
1(4)9* location: 4- (not located).
discoverer: Hochman, 61 e. discoverer: Grandmann, 62b.
synonym: l(4)ar: lethal(4) in chromosome containing synonym: l(4)ST-4: Mhal(4) Solway, Tmn@&Mee-4.
abdomen rota Cum. references: Hochman, 1963, DIS 37: 49.
references: Hochman, Gloor, and Green, 1964, Hochman, Gloor, and Green, 1964, Genetica 35:
Genetica 35: 109-26. 109-26.
ph*notype: Like 1(4)9. RK3. phenotype: Most homozygotes die as pupae. A few
other information: Proof that the lethal is not at the survive, especially in uncrowded cultures. Sur-
ar locus comes from the observation that vivors have spread wings and minor vein abnor-
D%4)M/ar 1(4)9*> survives and exhibits the ar malities such as crossveins between L2 and L3;
males lack external genitalia but produce motile synonym: l(4)AM-3: lethal(4) Amherst, Massachu-
sperm; both sexes sterile. RK3. setts-3.
1(4)15: see 1(4)6^ references: Hochman, Gloor, and Green, 1964,
1(4)15" Genetica 35: 109—26.
origin: Spontaneous. phenotype: Homozygotes die as embryos. RK3.
discoverer: Wrathall, 62a. 1(4)23: see 1(4)21>
synonym: 1(4)SLC-1: lethal(4) Salt Lake City-1. 1(4)24: see Df(4)24
references: Hochman, 1963, DIS 37: 49. 1(4)25
Hochman,, Gloor, and Green, 1964, Genetica 35: location: 4- [within Df(4)M\.
109-26. origin: Spontaneous.
phenotype: Like 1(4)15. RK3. discoverer: Hochman, 62a.
synonym: l(4)ST-3: lethal(4) Solway, Tennessee-3.
l(4)15t> references: 1963, DIS 37: 48-49.
origin: Spontaneous. Hochman, Gloor, and Green, 1964, Genetica 35:
discoverer: Lipe, 621. 109-26.
synonym: 1(4)MW-1: lethal(4) Madison, Wisconsin-1. phenotype: Homozygotes die as larvae. RK3.
references: Hochman, Gloor, and Green, 1964, cytology: Placed in salivary chromosome region
Genetica 35: 109-26. 101E through 102B16, on basis of its inclusion in
phenotype: Like 1(4)15. RK3. Df(4)M = Df(4)101E-F;102B6-17.
1(4)17: see Df(4)17 other information: Incorrectly reported as an allele
1(4)18 of 1(4)1* by Hochman, Gloor, and Green (1964).
location: 4- [within Df(4)M63*]. 1(4)26: see 1(4)14*
origin: X ray induced. 1(4)27: see 1(4)1*
discoverer: Gloor and Green, 1957. 1(4)28: see 1(4)1 *>
synonym: 1(4)35. 1(4)29
references: Hochman, Gloor, and Green, 1964, location: 4- Lwithin Dt(4)G\.
Genetica 35: 109-26. origin: Spontaneous.
phenotype: Homozygous lethal. 1(4)18/ ci is cf; discoverer: Hochman, 62k.
l(4)18/M(4)63a is lethal; l(4)l8/ciD is viable. synonym: l(4)BU-2: lethal(4) Bountiful, Utah-2.
About half the 1(4)18/ci+3 flies raised at 25°C references: Hochman, Gloor, and Green, 1964,
show L4 interruption. RK3A. Genetica 35: 109-26.
cytology: Placed in salivary chromosome region phenotype: Homozygotes die as pupae. Lethal when
101F2-102A5, on the basis of its inclusion in heterozygous with Di(4)3, Dt(4)ll, Df(4)12,
Df(4)M63" =Df(4)101F2-102Al;102A2-5. Associ- Df(4)24, Df(4)34, and Df(4)G. Wild type when het-
ated with T(3;4)l-18; breakpoints unknown. erozygous with alleles of 1(4)9, spa, or sv. RK3.
1(4)18: see 1(4)4° 1(4)29: see 1(4)2°
1(4)19: see l(4)8b 1(4)29°
1(4)20: seel(4)4b origin: Spontaneous.
1(4)21 discoverer: Kidwell, 621.
location: 4- (not located). synonym: l(4)OC'l; lethal(4) Ottawa, Canada-1.
origin: Spontaneous. references: Hochman, Gloor, and Green, 1964,
discoverer: Wrathall, 611. Genetica 35: 109—26.
synonym: 1(4)ST-1: lethal(4) Solway, Tennessee-1. phenotype: Like 1(4)29. RK3.
references: Hochman, 1963, DIS 37: 48. 1(4)29^
Hochman, Gloor, and Green, 1964, Genetica 35: origin: Spontaneous.
109-26. discoverer: Hochman, 6319.
phenotype: Homozygotes die as pupae. RK3. phenotype: Like 1(4)29. RK3.
1(4)21: see 1(4)2* 1(4)30: see 1(4)1d
1(4)22 1(4)31: see Di(4)31
location: 4- (not located). 1(4)32: see 1(4)1c
origin: Spontaneous. 1(4)33: see 1(4)10
discoverer: Wrathall, 62a. 1(4)34: see Df(4)34
synonym: 1(4)AM-1: lethal(4) Amherst, Massachu- 1(4)34: see 1(4)2*
setts-1, 1(4)33: see 1(4)18
references: Hochman, 1963, DIS 37: 49. 1(4)36: see l(4)6a
Hochman, Gloor, and Green, 1964, Genetica 35: 1(4)37: see 1(4)2*
109-26. 1(4)38: see 1(4)2*
phenotype: Homozygotes die as pupae. RK3. l(4)AM-t: see 1(4)22
1(4)23 l(4)AM-2: see l(4)4d
location: 4- (not located). l(4)AM-3: see 1(4)23
origin: Spontaneous. l(4)at: see 1(4)9*
discoverer: Lipe, 62k. 1(4)BU-1: see 1(4)9
MUTATIONS 141
l(4)BV-2: see 1(4)29 electrophoretic zone of mobility intermediate be-

1(4)MW-1: see 1(4)15* tween that of the slow and fast LAP A types. RK3.
l(4)OC-t: see 1(4)29* Lap-DF: Leucine aminopeptidase D-Fast
1(4)PT-1: see 1(4)6* location: 3-98.3 (Falke and Mactntyre).
l(4)PT-2: see 1(4)2* origin: Naturally occurring allele.
l(4)PT-3: see l(4)4c discoverer: Beckman and Johnson.
1(4)SLC-1: see 1(4)15' references: 1964, Hereditas 51: 221-30 (fig.).
1(4)ST-1: see 1(4)21 Falke and Maclntyre, 1966, DIS 41: 165-66.
l(4)ST-2: see /f4J24*> phenotype: Leucine aminopeptidase D is one of six
l(4)ST-3: see /(4J25 such enzymes detected by the method used to dem-
l(4)ST-4: see /(4Ji5 onstrate LAP A. LAP D is found in pupae and to
/ac: lacquered some extent in old larvae, but not in young larvae
location: 1-7.3. or adults. Lap-DF'/Lap-D? produces LAP D of high
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- mobility. RK3.
phenylalanine (CB. 3026). Lap-Ds: Leucine aminopeptidase D-S/ow
discoverer: Fahmy, 1953. origin: Naturally occurring allele.
references: 1959, DIS 33: 87. discoverer: Beckman and Johnson.
phenotype: Pale fly with chitin glistening as though references: 1964, Hereditas 51: 221-30 (fig.).
polished. Bristles long and straggly, frequently Falke and Maclntyre, 1966, DIS 41: 165-66.
duplicated. Eyes smaller and slightly bright. phenotype: Homozygotes produce slowly migrating
Wings often longitudinally pleated. Slightly LAP D. Lap-DF/Lap-Ds produce equal amounts of
delayed eclosion; viability and fertility reduced in slowly and rapidly migrating LAP D and no enzyme
both sexes. RK2. of intermediate mobility. RK3.
other information: Two alleles each induced by X Large: see Lg
rays, CB. 3034, CB. 1540. One allele each induced late hatching: see Ih
by CB. 3025, CB. 1506, CB. 2511. Six alleles Id: loboid
induced by CB. 1528. location: 3-102 [between ca and bv (Lewis, 1956,
lame: see I me DIS 30: 130)].
lanc&: see nw2 origin: Spontaneous.
lance-b: see // discoverer: Curry, 39a.
lanceolate: see // references: 1939, DIS 12: 45.
Lap-A°: Leucine aminopeptidase A-less phenotype: Eyes resemble L/+. Malformation of
location: 3- (near L,ap-D; no recombination yet ob- eyes ranges from slight dorsoventral seam across
served). middle of eyes to a more extreme effect in which
origin: Naturally occurring allele. growth of anterior part is completely inhibited in
discoverer: Beckman and Johnson. most-extreme cases. Antennalike outgrowth fre-
references: 1964, Hereditas 51: 221-30. quent where growth of eyes is suppressed. Tends
phenotype: Leucine aminopeptidase A is one of six to overlap wild type. RK3.
such enzymes that may be demonstrated in
Drosophila by starch gel electrophoresis when a origin: Spontaneous.
discontinuous Tris borate buffer is used at room discoverer: Edmonds on, 52a.
temperature at 6—8 V/cm. The enzyme is stained references: 1952, DIS 26: 60.
with L-leucyl-^S-naphthylamide and Black K salt in phenotype: Like Id. RK3.
0.2 M Tris Maleate buffer pH 5.2. LAP A is found leg tumor: see Igt
in homogenates of both larvae and pupae but not of *lem: lemon
adults. Lap-A°/Lap-A° horaogenates have no LAP location: 1-17.5.
A electrophoretic band. RK3. origin: Spontaneous.
F
Lap-A : Leucine aminopeptidase A-Fast discoverer E. M. Wallace, 12h.
origin: Naturally occurring allele. references: Morgan and Bridges, 1916, Carnegie
discoverer: Beckman and Johnson. Inst. Wash. Publ. No. 237: 48 (fig.).
references: 1964, Hereditas 51: 221-30 (fig.), phenotype: Body color pale yellow, with dark trident
phenotype: L&p-AF/Lap-AF and Lap-A^/Lap-A0 and black bristles. Wings and veins pale yellow.
produce LAP A, which migrates faster in starch gel Easily distinguished from wild type, but viability
electrophoresis than LAP A produced by Lap-As.
about 70 percent wild type, and most flies sterile.
RK3.
RK3.
Lap-As: Leucine aminopeptidase A-Slow lethal(): see l( )
origin: Naturally occurring allele. Leucine aminopeptidase A: see Lap-A
discoverer: Beckman and Johnson. Leucine aminopeptidase D: see Lap-D
references: 1964, Hereditas 51: 221-30 (fig.), If: little fly
phenotype: Lmp-As/Lmp-As and Lmp~As/L®p-A® location: 1-68.1.
produce slowly migrating LAP A. L,Mp~Ap'/L&p~A5 origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
produce enzyme characterized by a rather wide phenylalanine (CB. 3007).
discoverer: Fahmy, 1954. causes striking changes in compounds that

references: 1959, DIS 33: 87. fluoresce in ultraviolet light on paper chromato-
phenotype: Small fly with markedly narrow abdomen, grams of testes. Isoxanthopterin content of testis
frequently with small tumors. Low viability and sheath greatly reduced. A blue fluorescent com-
fertility in both sexes, but especially females. pound not otherwise detected in D. melanogaster
RK3. (the lix substance) is present. Drosopterins
other information: One allele each induced by CB. present in the testis sheath, and quantities of
3025, CB. 3051, CB. 1592, CB. 1414, CB. 1506, sepiapteridine, biopterin, "Compound A," and
and X rays. Two alleles induced by CB. 1528. "riboflavinlike" are elevated. The colored
Lffll: see l(l)ffll pteridine gives testis sheath its darker color.
*Lg: Large Pteridine accumulation in testis sheath alone is
location: 1-27. affected. RK3.
origin: Induced by P 3 2 . //; lanceolate
discoverer: Bateman, 1950. location: 2-106.7.
phenotype: Heterozygote large, late eclosing, with discoverer: Bridges, 23d3.
visibly smaller hairs; viability excellent. Tend- synonym: lance-b.
ency toward shortening of L4 and L5, missing post- references: Morgan, Bridges, and Sturtevant, 1925,
vertical bristles, and islands of vein tissue on Bibliog. Genet. 2: 227.
either side of L2. Homozygous lethal. RK2. Bridges, 1937, Cytologia (Tokyo), Fujii Jub. Vol.
*Igh: long haired 2: 745-55.
location: 1-20.7. phenotype: Wings narrowed at tips and slightly
origin: Induced by 2-chloroethyl methanesulfonate divergent. Eyes slightly smaller than normal and
(CB. 1506). bulging; head narrow. Waddington finds wing effect
discoverer: Fahmy, 1956. detectable in middle pupal stage. RK3.
references: 1959, DIS 33: 87. cytology: Placed in region between 59E2 and 60B10
phenotype: Small fly; size reduction most noticeable on the basis of its being to the right of
in head and thorax. Wings short and slightly In&RybwW*1 =ln(2R)41B2-Cl;59E2-4 and to the
altered in shape. Anterior thorax frequently dented left of Df(2R)Px = Df(2R)60B8-10;60Dl-2.
in the mid-dorsal line. Hairs deranged; bristles
long and straggly. Abdomen nearly always abnor-
mally pigmented, ranging from no melanization of
tergites 5—7 to small, irregular underpigmented
patches on these tergites. Male viability about 25
percent wild type. Males sterile. RK3.
lgl: see l(2)gl
*lgt: leg tumor
origin: Spontaneous, II2: lanceolate-2
discoverer: Spencer, 36c20. Edith M. Wallace, unpublished.
references: 1937, DIS 7: 14. 112
phenotype: Black tumor growths inside thorax ven- origin: Spontaneous.
trally at bases of posterior legs. Sterile in both discoverer: Bridges, 23d25.
sexes; poor viability. RK3. phenotype: Wings pointed and narrow. Eyes small
Ih: late hatching and bulging. Head narrow. Wing shape first seen
location: 1-57. in early contraction stage of wing development
origin: Spontaneous, (23-hr pupa at 25°C), according to Waddington
discoverer: Bridges, 31d6. (1939, Proc. Natl. Acad. Sci. U.S. 25: 303). More
phenotype: Slow-developing semigiant. RK3. extreme and more useful than //. RK2.
light: see It Im: limited
lightoid: see ltd location: 2-50.
limited: see Im origin: Spontaneous.
little fly: see If discoverer: Bridges, 29125.
lix: little isoxanthopterin phenotype: Sternites small, rounded, or irregular;
location: 1-23. bristles sparse. Females sterile. RK3.
origin: Spontaneous. cytology: Not Included in Dl(2L)64j = Df(2L)34E5-
discoverer: Hessler, 1959. F1;35C3-D1 (E. H. Grell).
references: 1960, DIS 34: 50. *lme: lame
Hubby, 1962, Genetics 47: 109-14. location: 1-47.8.
phenotype: Flies indistinguishable from wild type; origin: Induced by 2-chloroethyl methanesulfonate
dissected testis sheath dark yellow-orange, but (CB. 150§).
this character not dependable for classification; discoverer: Fahmy, 1956.
MUTATIONS 143
references: 1959, DIS 33: 87. phenotype: Like It3 in combination with other It
phenotype: Legs weak, frequently deformed and gen- alleles. Rare homozygotes obtained are short lived
erally shortened as a result of reduction in length and sterile. RK2.
of tarsal segments. Wings a typically shaped and *lts
abnormally held. Flies so crippled they cannot origin: Ultraviolet induced.
move; they die soon after eclosion. RK3. discoverer: Meyer, 51d.
lme: see l(2)me references: Meyer and Edmondson, 1951, DIS 25: 73.
Lobe: see L phenotype: Like It*. Homozygote lethal, as is
loboid: see Id It4'/It5. It3/It5 is viable and has mutant eye color.
long haired: see Igh RK2.
low xanthine dehydrogenase: see Ixd *lfS6e
lozenge: see Iz origin: Spontaneous (arose together with Atu56c).
lozenge-like: see rstl discoverer: Meyer, 56c.
lozengelike: see Izl references: 1956, DIS 30: 77.
It: light phenotype: Like It; has good viability. RK1.
location: 2-55.0 Oust to the left of spindle attach- */*•»*: light-mottled
ment). origin: X ray induced.
origin: Spontaneous, discoverer: Hessler, 1957.
discoverer: Bridges, 24dS. references: 1958, Genetics 43: 395-403.
references: 1931, Eos 7: 229-48. phenotype: Pale mottled; eyes a mixture of light and
de Zulueta, 1931, Eos 7: 249-53. wild type ommatidia. RK2A.
phenotype: Eye color yellowish pink — lighter at cytology: Associated with T(2;3)1P*1 = T(2;3)40B-
high temperatures, darker at low. Ocelli colorless. F;63B-F.
At 25°C, eyes have 12 percent wild-type red pig- *ltn,2
ment and 9 percent wild-type brown pigment (Nolte, origin: X ray induced.
1954, J. Genet. 52: 127—39); with st, color only discoverer: Hessler, 1957.
slightly lighter than with It alone; with bw, it is a references: 1958, Genetics 43: 395—403.
clear lemon yellow, pinkish in old flies (Schultz phenotype: Dark mottled; eyes a mixture of wild type
and Dobzhansky, 1934, Genetics 19: 344-64; and occasional darker ommatidia. RK2A.
Mainx, 1938, Z. Induktive Abstammungs- cytology: Associated with In(2L)lt™2 = In(2L)22F-
Vererbungslehre 75: 256—76). Eye color auton- 23A;4QB-F.
omous when larval optic disk is transplanted into Itm3
wild-type host (Beadle and Ephrussi, 1936, origin: X ray induced.
Genetics 21: 230). Larval Malpighian tubes color- discoverer: Hessler, 1957.
less in It offspring of lt/lt mothers; some color in references: 1958, Genetics 43: 395—403.
tubes if mother is lt/+. It stw/It stw is completely phenotype: Dark mottled like It*"2. RK2A.
in viable (Purdom); however, It stw3/ It stw3 has cytology: Associated with In(2LR)W»3 =
good viability. RK1. In(2LR)40B-F;60D.
cytology: Placed in 40B-F on basis of breakpoints *ltm4
common to rearrangements that produce mottling for origin: X ray induced.
It (Hessler, 1958, Genetics 43: 395-403). discoverer: Hessler, 1957.
*lf2 references: 1958, Genetics 43: 395—403.
origin: Spontaneous. phenotype: Dark mottled like ltm2. RK2A.
discoverer: Bridges, 30bl4. cytology: Associated with T(2;3)tta'4 ~ T(2;3)40B-
references: 1931, Eos 7: 229-48. F;67E.
phenotype: Eye color slightly maroon, differs little *ltmS
from wild type. Intensified by b&2 and more ex- origin: X ray induced.
treme in females. RK3. discoverer: Hessler, 1957.
It3 references: 1958, Genetics 43: 395—403.
origin: Spontaneous in In(2L)Cy + ln(2R)Cy, phenotype: Pale mottled like ltml, RK2A.
a/2 Cy L4 sp2. cytology: Associated with T(2;3)ltm5 = T(2;3)40B-
discoverer: Beadle, 36e23. F;98C.
phenotype: Eye color of It3/It darker than lt/lt. *lfn,6
Larval Malpighian tubes of U3/lt colorless when origin: X ray induced,
derived from 1/ mothers. Since it3 is in the rear- discoverer: Hessler, 1957.
ranged lethal-bearing chromosome the homozygote references: 1958, Genetics 43: 395-403.
has not been obtained. RK1A. phenotype: Pale mottled like ltm*. RK2A.
IH cytology: Associated with f]f2;3>/*«* « T(2;3)26E-
origin: Ultraviolet induced, F;4QB~F;96E.
discoverer: Meyer, 50d.
references: Meyer, Edmonds on, Byers, and Erickson, origin: X my induced.
19S0, DIS 24: 60. discoverer: Hessler, 1957.
references: 1958, Genetics 43: 395—403. discoverer: Hessler, 1957.

phenotype: Pale mottled like It0*1. RK2A. references: 1958, Genetics 43: 395-403.
cytology: Associated with T(2;3)1P*>7 = T(2;3)40B- phenotype: Pale mottled like ltml. RK2A.
F;l00F. cytology: Associated with T(l;2)ltm16 =
*ltm8 T(1;2)11A;12F;22D;4OB-F.
origin: X ray induced. *lfml7
discoverer: Hessler, 1957. origin: X ray induced.
phenotype: Dark mottled like ltm2. RK2A. references: 1958, Genetics 43: 395-403.
cytology: Associated with T(2;3)lt™8 = T(2;3)40B- phenotype: Pale mottled like ltml. RK2A.
F;92B. cytology: Associated with T(2;3)lt™17 = T(2;3)40B-
*lfm9 F;95C-D.
origin: X ray induced. * Ifm 18
phenotype: Dark mottled like ltm2. RK2A. references: 1958, Genetics 43: 395—403.
cytology: Associated with In(2LR)ltm? = phenotype: Dark mottled like ltm2. RK2A.
In(2LR)40B-F;56E. cytology: Associated with T(2;3)W«18 = T(2;3)40B-
*ffm 7 0 F;98A.
origin: X ray induced. *ltm19
phenotype: Dark mottled like ltm2. RK2A. references: 1958, Genetics 43: 395—403.
cytology: Associated with T(2;3)ltm10 =T(2;3)40B- phenotype: Dark mottled like It™2. RK2A.
F;64E. cytology: Associated with T(2;3)lt™19 = T(2;3)40B-
*ltmll F;94B.
origin: X ray induced. *ltm 20
references: 1958, Genetics 43: 395-403. discoverer: Hessler, 1957.
phenotype: Dark mottled like It1"2. RK2A. references: 1958, Genetics 43: 395—403.
cytology: Associated with T(2;3)ltm11 - T(2;3)40B- phenotype: Pale mottled like 1 tml. RK2A.
F;96F. cytology: Associated with In(2L)ltm20 =
Ifm 12 In(2L)32C;40B-F.
discoverer: Hessler, 1957. * Ifm 21
references: 1958, Genetics 43: 395—403. origin: X ray induced.
phenotype: Dark mottled like ltm2. RK2A. discoverer. Hessler, 1957.
cytology: Associated with ln(2LR)Hml2 = references: 1958, Genetics 43: 395—403.
In(2LR)40B-F;60D. phenotype: Dark mottled like ltm2. RK2A.
cytology: Associated with T(2;3)ltm21 = T(2;3)40B-
origin: X ray induced, F;93D,
discoverer: Hessler, 1957. *ltm22
phenotype: Dark mottled like ltm2. RK2A. discoverer: Hessler, 1957.
cytology: Associated with T(2;3)ltm13 = T(2;3)40B- references: 1958, Genetics 43: 395-403.
F;64F. phenotype: Dark mottled like im2. RK2A.
* ltm U cytology: Associated with In(2LR)ltm22 =
origin: X ray induced. In(2LR)4QB~F;59D.
discoverer: Hessler, 1957. *Ifm 23
phenotype: Dark mottled like ttm2, RK2A. discoverer: Hessier, 1957.
cytology: Associated with T(2;3)lt™1'* « T(2;3)40B- references: 1958, Genetics 43: 395—403.
F;9SF. phenotype: Pale mottled like ltm2. RK2A.
*Ifm IS cytology: Associated with T(2;3)lt**23 = T(2;3)40B-
origin: X my induced. F;62F.
discoverer: Hessler, 1957. *ltm24
phenotype: Pale mottled like ltml. RK2A. discoverer Hessler, 1957.
cytology: Associated with T(2;3)lt*** s = T(2;3)40B- references: 1958, Genetics 43: 395-403.
F;92B. phenotype: Pale mottled like ltmI. RK2A.
* ltm 16 cytology: Associated with T(2;3)ltm24 « T(2;3)4QB-
origin: X ray induced. F;59F;75C
MUTATIONS 145
*ltm2S
cytology: Associated with In(2LR)ltm33 =
origin: X ray induced. In(2LR)40B-F;58B.
discoverer: Hessler, 1957. *lfm34
phenotype: Pale mottled like !*»>•*. RK2A, discoverer: Hessler, 1957.
cytology: Associated with In(2LR)lt™25 = references: 1958, Genetics 43: 395—403.
In(2LR)40B'F; 57C-D. phenotype: Pale mottled like It™1. RK2A.
*lfm26 cytology: Associated with T(2;3)lf"34 - T(2;3)40B-
origin: X ray induced. F;61B.
discoverer: Hessler, 1957. *ltm3S
phenotype: Pale mottled like ltml. RK2A. discoverer: Hessler, 1957.
cytology: Associated with ln(2L)ltm26 = references: 1958, Genetics 43: 395-403.
In(2L)27C;40B-F. phenotype: Pale mottled like It1*1. RK2A.
*\fn>27
cytology: Associated with T(2;3)ltm3^ = T(2;3)40B-
origin: X ray induced. F;64C.
discoverer: Hessler, 1957. IfmlOO
phenotype: Pale mottled like ltml. RK2A. discoverer: Spieler, 60a25.
cytology: Associated with T(2;3)ltm27 - T(2;3)40B- references: Baker and Rein, 1962, Genetics 47:
F;88E-F.
1399-1407.
*lfn>28
phenotype: Homozygous lethal. Variegated for It.
origin: X ray induced. RK2A.
discoverer: Hessler, 1957. cytology: Associated with T(2;3)ltml°0 =
references: 1958, Genetics 43: 395—403. T(2;3)40;97F.
phenotype: Pale mottled like It1"1. RK2A.
cytology: Associated with T(2;3)ltm2S = T(2;3)40B- *lfpk: light-pinkoid
F,97E. origin: Spontaneous.
ltm29 discoverer: Lancefield, 18cl8.
origin: X ray induced. synonym: pinkoid; pink-wing.
discoverer: Hessler, 1957. references: 1918, Biol. Bull. 35: 207-10.
references: 1958, Genetics 43: 395-403. Bridges, 1931, Eos 7: 229-48.
phenotype: Pale mottled like ltml. RK2A. phenotype: Eye color like pink. Darker than It and
cytology: Associated with T(2;3)ltm29 = T(2;3)40B- lighter than It2. Wings short and crinkled. Via-
F;99F. bility 30 percent wild type. RK3.
*lfm30
ltd: lightoid
origin: X ray induced, location: 2-56 (cytology at variance with this loca-
discoverer: Hessler, 1957. tion).
references: 1958, Genetics 43: 395—403. origin: Spontaneous.
phenotype: Dark mottled like It™2. RK2A. discoverer Nichols-Skoog, 36d6.
cytology: Associated with T(2;3)Jf»30 _ T(2;3)40B- phenotype: Eye color clear light, translucent
F;99C. yellowish pink. Resembles // but is lighter;
darkens with age. Ocelli colorless; larval Mal-
origin: X ray induced, pighian tubes colorless. Eyes have 38 percent
discoverer: Hessler, 1957. wild-type red pigment and 7 percent wild-type
references: 1958, Genetics 43: 395—403. brown pigment (Nolte, 1954, J. Genet. 52: 127-39).
phenotype: Pale mottled like /*<"-*. RK2A. RK1.
cytology: Associated with TXltfllP*31 = cytology: Placed outside the region 41A-44C, on the
T(l ;2)8F; 28D;40B- F. basis of its not being included in Dp(2;3)P32 -
Dp(2;3)41A:42D-E;44C-Di89D7'El (E. B. Lewis).
*ltm32
discoverer: Hessler, 1957. discoverer: Poulson, 37b.
references: 1958, Genetics 43: 395—403. references: Poulson and King, 1948, DIS 22: 55.
phenotype: Pale mottled like It*"*, RK2A. phenotype: Eye color of newly hatched adult bright
cytology: Associated with T(2;3)lt™32 «* T(2;3)40B- red like v; darkens to a color like pr in old flies.
F;97A. Ocelli colorless; larval Malpighian tubes colorless.
*tfm33 Viability excellent. RK1.
origin: X ray induced. Ixd: low xanthine dehydrogenase
discoverer: Hessler, 1957. location: 3-33.
references: 1958, Genetics 43: 395—403. origin: Spontaneous (naturally occurring alleie ?).
phenotype: Pale mottled like im*. RK2A. discoverer: Keller and Glassman, 61jl2.
references: 1964, Genetics 49: 663—68. 49h

1964, DIS 39: 61. 50e
phenotype: Homozygous flies exhibit only 25 percent 37
normal xanthine dehydrogenase activity. No ob- 34
vious morphological expression. The ma!+ com- 49g
plementation factor of Glass man (1962, Proc. Natl. 3n
Acad. Sci. U.S. 48: 1491-97) inhibited in lxd 481
extracts and oxidase activity of Forrest, Hanly, 48 c 0.083 gm 0.057 3
and Lagowski (1961, Genetics 46: 1455-63) ab- 36 48f g4
sent. Purine analogs more toxic to lxd than to Bs 46 9
Oregon-R, mal, or ry* This might mean further de- s I
rangement of purine metabolism in lxd. RK3. I
Map of the Iz locus

Drawn from Green and Green, 1956, Z. Induktive
Iz: lozenge
location: 1-27.7.
discoverer: Bridges, 16bl2.
Ly: Lyra Bibliog. Genet. 2: 230.
From Bridges and Brehme, 1944, Carnegie Inst. Wash. Green and Green, 1956, Z. Induktive Abstammungs-
Publ. No. 552: 118. Vererbungslehre 87: 708-21.
phenotype: Eye narrower than wild type and ovoid.
Irregular facets in some areas cause rough patches;
Ly; Lyra areas of fused facets appear as smooth patches.
location: 3-40.5. Eye color appears normal, but in combination with
origin: X ray induced. st slight reduction in red pigment detectable.
discoverer: Dubinin, 1929. Tarsal claws reduced. Developmental study by
references: Coyne, 1935, DIS 4: 59. Waddington and Pilkington (1942, DIS 16: 70)
Morgan, Bridges, and Schultz, 1937, Carnegie Inst. shows failure of middle cell layer of optic disk to
Wash. Year Book 36: 301. penetrate between cells of outer layer; surface
phenotype: Lateral margins of wings excised, giving thus covered with primary pigment cells. Females
narrowed shape; angle between veins L2 and L5 sterile. Parovaria and spermathecae absent; some
reduced. Bristles shortened and stubby, post- lz/+ females have abnormal parovaria (Anderson,
scutellars frequently missing. Eyes somewhat de- 1945, Genetics 30: 280-96). RK1.
formed, with tufted vibrissae. Abdomen dark and cytology: Located in 8D (region 8D4 through 8E2)
narrow, with rear edge of tergites raised. Homo- by Green and Green (1956). Earlier Demerec,
zygous lethal. Ly/M(3yh33! is lethal. Modification Kaufmann, Fano, Sutton, and Sansome (1942,
of wings first visible as marginal scalloping of pre- Carnegie Inst. Wash. Year Book 41: 191) placed
pupal wing buds; wing fold narrower (Waddington, locus between 8C3 and 8C17, based on its inclu-
1939, Proc. Natl. Acad. ScL U.S. 25: 304; 1940, sion within Dt(l)t282-1 = Df(l)8C2-3;8C14-Dl;
J. Genet. 41: 75-139). RK1A. however, Green and Green suggest that
cytology: Placed in 70A3-5 on the basis of its asso- Df(l)ta*2-1 m a y extend into 8D, a region unfavor-
ciation with D%3L)Ly - Df(3L)7QA2-3;70A5-6 able for cytology.
(Bridges). other information: The Iz region has been sub-
divided into four recombinationally separable
lys: lysine groups (Green and Green, 1949, Proc. Natl. Acad.
location: 2-22.9. Sci. U.S. 35: 586-91; 1956; Green, 1961, Genetics
origin: Spontaneous. 46: 1169—76). First three groups called spe:
discoverer: E H. Grell, 1957. spectacle, Iz: lozenge, and gly: glossy; lzK is
references: I960, DIS 34: 50. sole member of fourth sub locus (see map). All
1961, Genetics 46: 925-33. double mutants show extreme phenotype resembling
phenotype: Larvae, pupae, and adults contain a / 2 s . Several comparative studies of Iz alleles have
higher concentration of lysine than wild type. Ac- been published [Gottschewski, 1936, Zool. Anz.
cumuiation of lysine is postulated to result from Suppi. 9: 104—12; Anderson, 1945, Genetics 30:
block in its degradation. Flies homozygous for lys 280-96; Oliver, 1947, Texas Univ. Publ. 4720:
occasionally have faintly reddish fat cells, espe- 167-84; Clayton, 1952, ibid. 5204: 227-51; 1954,
cially ia thorax. This effect enhanced by starva- iWd. 5422: 189-209, 210-43; Chovnick and
tion, by combining lya with re, rc*t or cho. RK3. Lefkowitz, 1956, Genetics 41: 79-92 (fig.);
MUTATIONS 147
Chovnick, Lefkowitz, and Fox, 1956, ibid. 41: secondary cause, lz34 ovaries, when transplanted
589-604; Clayton, 1957, ibid. 42: 28—41 (fig.); into normal females, however, are more productive
1958, ibid. 43: 261-73 (fig.); 1959, ibid. 44: than when in lz34 females (Clancy and Welborn,
1041-52 (fig.)]. 1948, Genetics 33: 606). RK1.
1x3 other information: Located in leftmost (spe) lz sub-
origin: Spontaneous. locus.
discoverer: Bridges, 22bl4. *IZ35
synonym: gly3: glossy-3. origin: Spontaneous,
references: Morgan, Bridges, and Sturtevant, 1925, discoverer: Gottschewski, 1935.
Bibliog. Genet. 2: 230. references: 1937, DIS 8: 12.
Green and Green, 1956, Z. Induktive Abstammungs- phenotype: Eyes reduced and diamond shaped; color
Veyerbungslehre 87: 708-21. opaque brown. Homozygous females sterile.
phenotype: Eye size sharply reduced; surface Iz3*/lz females fertile. RK1.
smooth. Optic disk of mature larva and prepupa 1x36
two-thirds normal size (Chen, 1929, J. Morphol. 47: origin: Spontaneous.
135—99). Red pigment greatly reduced; color discoverer: Spencer, 36c.
yellowish brown; cream colored in combination synonym: lz36c; lz36cS; spe36.
with v. Malpighian tubes of mature larvae lighter references: Green and Green, 1956, Z. Induktive
than wild type; variable (Brehme and Demerec, Abstammungs- Vererbungslehre 87: 708—21.
1942, Growth 6: 351—56). Tarsal claws vestigal. phenotype: Like lz3 in texture, color of eyes, and
Homozygous females lack parovaria and sperma- color of larval Malpighian tubes. Parovaria and
thecae, and are sterile; Iz3/+ females lack paro- spermathecae absent from homozygous females,
varia and many have abnormal spermathecae which are sterile, and tend to be abnormal in
[Anderson, 1945, Genetics 30: 280-96 (fig.)]. RK1. Iz36/+ females. [Anderson, 1945, Genetics 30:
other information: Located in the rightmost Q&ly) lz 280-96 (fig.)]- RK1.
sublocus. other information: Located in the leftmost (spe) lz
Iz3n sublocus.
origin: Spontaneous. *lz3 6cD: lozenge-36c of Dempster
discoverer Green. discoverer: Dempster, 36c.
synonym: spe3n: spectacled-3n. phenotype: Eyes small, narrow, oval, and glossy;
references: Green and Green, 1956, Z. Induktive color light brown with red rim and patches at
Abstammungs- Vererbungslehre 87: 708—21. center. Females fertile. RK1.
phenotype: Eyes sharply reduced in size; surface 1x3 7
smooth, red pigment sparse and confined primarily origin: Spontaneous.
to margin of eye. Tarsal claws vestigial. Females discoverer Curry, 37hl7.
sterile; spermathecae and parovaria absent. RK1. synonym: lz37h; spe37.
other information: Located at leftmost (spe) sub- references: Green and Green, 1956, Z. Induktive
locus of lz region. Abstammungs- Vererbungslehre 87; 708—21.
IZ34 phenotype: Eye size reduced. Areas of irregular
origin: Spontaneous. facets in posterior region of eye; eye color normal.
discoverer Beadle, 34k22. Larval Malpighian tubes somewhat lighter than
synonym: lz34k; spe34. normal; variable (Brehme and Demerec, 1942,
references: 1935, DIS 4: 9. Growth 6: 351—56). Tarsal claws reduced. Sper-
Green and Green, 1956, Z. Induktive Abstammungs- mathecae abnormal or absent from homozygous fe-
Vererbungslehre 87: 708—21. males, which are sterile; present in Iz37/+ females
phenotype: Eye phenotype intermediate between lz [Anderson, 1945, Genetics 30: 280—96 (fig.)]-
and lz3. Surface of eye has large areas of fused RK1.
facets with a few normal facets (Clayton, 1957, other information: Located in leftmost (spe) lz sub-
Genetics 42: 28—41); eye color dark red with small locus.
yellowish spots. Larval Malpighian tubes slightly
lighter than normal; variable (Brehme and Demerec, origin: Spontaneous,
1942, Growth 6: 351-56), Tarsal claws reduced. d I s coverer Green.
Spermathecae and parovaria absent from homozy- references: Green and Green, 1949, Proc. Natl.
gous females, which are sterile; some Iz34/+ Acad. ScL U.S. 35: 586—91.
females have abnormal parovaria (Anderson, 1945, 1956, Z. Induktive Abstammungs- Vererbungslehre
Genetics 30: 280—96). The female-fertile stock, 87: 708-21.
lz34; mt(lz34), described by Beader and Green phenotype: Eye size reduced; surface rough; color
(1960, Genetics 45: 1563—66) also lack sperma- near normal. Tarsal claws reduced. Spermathecae
thecae and parovaria. Bender and Green's observa- and parovaria absent from females; females sterile.
tions indicate that ovarian abnormalities are pri- RX1.
marily responsible for sterility of lx34 females and other information: Located in lz sublocus of lozenge
absence of spermathecae and parovaria are only a region.
*fz48c references: 1950, DIS 24: 58.

origin: X ray induced. Green and Green, 1956, Z. Induktive Abstammungs-
synonym: spe48c, Vererbungslehre 87: 708-21.
references: Green and Green, 1956, Z. Induktive phenotype: Like lz*?*1 in all respects. Eyes reduced
Abstammungs- Vererbungslehre 87: 708—21. in size, and almond shaped; no indication of
phenotype: Eye size sharply reduced; surface facets; covered with indentations, giving a pock-
smooth; red pigment reduced and largely confined marked appearance. Hairs on eye surface sparse or
to margin of eye. Tarsal claws vestigial. Females absent; eye surface glossy with many large black
sterile; lack spermathecae and parovaria. RK1. or brown flecks. Tarsal claws normal. Females
other information: Located in leftmost (spe) lz sub- fertile; spermathecae and parovaria present and
locus. normal. lzSOe/lz has normal eyes except for a few
lz48f flecks. Complements most other lz alleles except
origin: Induced by mustard gas. lz*9fl, lzS2c, and those associated with rearrange-
discoverer: Lindsley, 48f. ments or deficiencies. RK1.
references: Green and Green, 1956, Z. Induktive other information: Located in the leftmost (spe) lz
Abstammungs- Vererbungslehre 87: 708—21. sublocus.
phenotype: Like /z-*«c. RK1. *lzSld
other information: Located in lz sublocus. origin: Spontaneous.
discoverer: Mossige, 51dl0.
*IZ481 references: 1951, DIS 25: 69.
origin: X ray induced. phenotype: Eye size extremely reduced; surface
synonym: spe*81. smooth and glossy; color light yellow with
references: Green and Green, 1956, Z. Induktive brownish margin and spots. Females sterile. RK1.
Abstammungs- Vererbungslehre 87: 708—21. */ z 52c
phenotype: Like lz4S*. RK1. origin: Recovered among progeny of male fed H3BO3
other information: Located in the leftmost (spe) lz and exposed to thermal neutrons.
sublocus. discoverer: R. C. King, 52c28.
origin: Spontaneous. phenotype: Eyes mottled; yellowish brown, darker at
synonym: $pe*9<3. rim; facets fused. Males setnisterile, with missing
references: Green and Green, 1956, Z. Induktive tarsal claws, although pulvilli and endopodia
Abstammungs- Vererbungslehre 87: 708—21. normal. Third antennal segment slightly reduced.
phenotype: Like lz**<=. RK1. lzS2c/iz50e f em ales resemble /z^°*. RK1.
other information: Located in the leftmost (spe) lz
sublocus. *lz55d
*lz49h origin: X ray induced.
origin: X ray induced. discoverer Clark, 55d.
discoverer: W. K. Baker, 49h. references: 1956, DIS 30: 71.
synonym: ape49h. phenotype: Eyes smaller than wild type and oval in
references: Green and Green, 1956, Z. Induktive shape; all facets run together. Females sterile.
Abstammungs- Vererbungslehre 87: 708—21. RK1.
phenotype: Eye size sharply reduced; surface / Z 55/
smooth; red pigment distributed over entire eye. origin: Spontaneous.
Tarsal claws normal. Spermathecae and parovaria discoverer: Masters on, 551.
present and normal in females, which are fertile. references: Clancy, 1960, DIS 34: 48.
When heterozygous with all lozenge alleles except phenotype: Like lz8. Paper chromatography reveals
tz*9li, phenotype is wild type or very nearly so. trace of red eye pigments. RK1.
RK1.
other information: Located in the leftmost (spe) lz origin: X ray induced,
sublocus. discoverer Mayo, 57j.
IzSOd phenotype: Like lza. RK1.
origin: X ray induced. lz58d
discoverer: Ritterhoff, 50d. origin: Spontaneous.
references: Glass, 1951, DIS 25: 77. discoverer: Schreckengost, 58d.
phenotype: Like lzs. Females sterile. RK1. references: Clancy, 1960, DIS 34: 48.
phenotype: Like lza. Paper chromatography reveals
trace of red eye pigments. RK1.
origin: Recovered among progeny of male fed P32. */ z 59
containing medium. origin: X ray induced.
discoverer R. C. King, 50e30. discoverer: Polivanov, 1959.
synonym: 50e.
spe) references: 1963, DES 38: 30-31.
MUTATIONS 149
phenotype: Eyes reduced in size, and ovoid; facets parovaria absent from females, which are sterile.
fused; surface slightly rough and almost or com- RK1.
pletely hairless; color light brown with darker, other information: Located in the leftmost (spe) lz
slightly reddish rim; almost colorless in combina- sublocus.
tion with v. Tarsal claws practically absent as in */z c ': lozenge-clawless
lzcl. Males sterile; transmit no motile sperm to fe- origin: Appeared as a male from an ovary treated in
males; therefore homozygous females not observed. vitro with CuSO4.
lzS9/lz37 females intermediate between the two discoverer: Hadorn, 45b27.
mutants in eye phenotype, have reduced tarsal references: Hadorn and Anders, 1946, DIS 20: 65.
claws, and are weakly fertile. RK2. Anders, 1955, Z. Induktive Abstammungs-
lz61f Vererbungslehre 87: 113-86 (fig.).
origin: Spontaneous. phenotype: Eyes narrow and small, without facets;
discoverer: Moynehan, 61f. surface has rough spots; color amber; both
references: Burdick, 1963, DIS 37: 47. pteridines and ommochromes affected; darker at
phenotype: Facets completely fused; eye color dark, rim. Tarsal claws absent. Third antennal segment
but pigment unevenly distributed and concentrated reduced; sensillae on antennae abnormal. Pheno-
at margin. Females fertile. Iz6*l/lz females more type similar in both sexes. Females infertile and
normal than either mutant, with facets disrupted lack spermathecae and parovaria. Autonomous in
and fused only in posterior third of eye; also transplants. RK1.
fertile. RK1. lzD: lozenge-Dominant
lz62k origin: Spontaneous.
origin: X ray induced. discoverer: Novitski, 47i.
discoverer: Mickey, 62kll. references: 1949, DIS 23: 61.
references: 1963, DIS 38: 28. phenotype: Males and homozygous females resemble
phenotype: Like lza. RK1. lza. Heterozygous females sometimes have
IZ63 roughened eyes. Apparent dominance shown by
origin: X ray induced. H. Bender to be caused by the presence of
discoverer: Halfer, 1963. spae(Iz); heterozygous expression additionally en-
phenotype: Eye shape oval; color brown, darkest at hanced by presence of In(2LR)bwvl. RK1 as re-
margin; surface smooth and glossy. Viability and cessive; RK3 as dominant.
fertility of both sexes good. RK1. *lzf: lozenge-fertile
lz63f origin: Spontaneous.
origin: Spontaneous. discoverer: Muller.
discoverer Burdick, 63fl7. references: 1946, DIS 20: 67.
references: Seiger and Bender, 1963, DIS 38: 31. phenotype: Intermediate allele like lz. Female mod-
phenotype: Eye size moderately reduced; surface erately fertile. RK2.
smooth; color brownish with darker margin. Tarsal Izs: lozenge-glossy
claws and pulvilli strongly reduced. Spermathecae origin: X ray induced.
and parovaria absent; female reproductive capacity discoverer: Oliver, 31a7.
strongly reduced. lz63t complements lz50e but not synonym: ily1.
lz34, lzD, or lz61t (Klingele). Spermathecal number references: 1935, DIS 4: 15.
o£iz63i/izK 0-3. RK1. Green and Green, 1956, Z. Induktive Abstammungs-
*lz268-29 Vererbungslehre 87: 708—21.
origin: X ray induced. phenotype: Eyes smaller than wild type; surface
discoverer: Hoover, 38d. glossy from fused facets; a few normal facets also
phenotype: Lethal, but not shown that lethality is at present; color dark blood red, bright red in combi-
Iz locus. Fertile in combination with lz. RK2A. nation with st or v. Larval Malpighian tubes
cytology: Associated with T(l;3)lz26!*-29 - slightly lighter than normal (Brehme and Demerec,
T(1;3)8D8-9;81F. 1942, Growth 6: 351-56). Tarsal claws reduced.
other information: Induced simultaneously with Spermathecae and parovaria absent from homozy-
gous females, which have reduced fertility; Iz6/+
lzBS: lozenge from Bar-Stone females tend to have abnormal parovaria [Anderson,
origin: X ray induced, 1945, Genetics 30: 280-96 (fig.)]. RK1.
synonym: spe^^. other information: Located in rightmost (jffty) lz sub-
references: Oliver, 1947, Texas Univ. Publ. 4720: locus. lz6/lzB provided probably the first recorded
167-84. case of intra-allelic recombination (Oliver, 1940,
Green and Green, 1956, Z. Induktive Abstammungs- Proc. Natl. Acad. Sci. U.S. 26: 452-54; 1940, DIS
Vererbungslehre 87: 708—21. 13: 73).
phenotype: Eye size reduced; surface rough with *lz9>: lozenge-glued
scattered fused facets; color nearly normal; some origin: X ray induced.
reduction of red pigment detected in combination discoverer: M. A Bender, 53k.
with st. Tarsal claws reduced; spermathecae and references: 1955, DIS 29: 69.
phenotype: Eyes of male reduced and roughened like 1940, Proc. Natl. Acad. Sci. U.S. 26: 452-54;
Gl; color dark; female eyes somewhat less extreme. 1940, DIS 13: 73).
Iz6l/lz intermediate between lz&1 and Iz and *lz*B: lozenge-spectacled of Bishop
sterile. Homozygous females fertile. RK1. origin: X ray induced.
: lozenge-glossy of Muller references: Oliver, 1947, Texas Univ. Publ. 4720:
origin: Spontaneous. 167-84.
references: Green and Green, 1956, Z. Induktive phenotype: An extreme lozenge allele similar to lzs.
Abstammungs- Vererbungslehre 87: 708—21. Eye color yellowish brown. Homozygous females
phenotype: Eye size reduced; surface rough; red lack spermathecae and parovaria and are sterile;
pigment distributed over entire eye. Tarsal claws lzaB/+ females have reduced numbers of sperma-
reduced. Spermathecae and parovaria absent. Fe- thecae and parovaria (Anderson, 1945, Genetics 30:
males sterile. RK1. 280-96). RK1A.
other information: Located in the Iz sublocus of the cytology: Associated with In(l)lzsB = In(l)8;20
lozenge region. (Green).
lzK: lozenge of Krivshenko IZY*: lozenge in yellow-4
discoverer: Krivshenko, 55k9. synonym: £\yY*-
synonym: amxss: almondex-55; lzk. references: Oliver, 1947, Texas Univ. Publ. 4720:
references: 1956, DIS 30: 74. 167-84.
Green, 1961, Genetics 46: 1169-76 (fig.). Green and Green, 1956, Z. Induktive Abstammungs-
phenotype: Eyes narrow and moderately rough; Vererbungslehre 87: 708-21.
facets irregular; eyes of homozygous females more phenotype: Similar to lza but eye color redder. Ho-
nearly normal than those of males. Tarsal claws mozygous females lack spermathecae and parovaria
normal. Females fertile; spermathecae and paro- and are sterile; lzVt/+ females have abnormal par-
varia present. Interactions of lzK with other Iz ovaria and tend to lack spermathecae and parovaria
alleles described by Green (1961). RK1. (Anderson, 1945, Genetics 30: 280-96). RK1.
other information: Located between the spe and Iz other information: Located in the rightmost (gly) Iz
subloci. sublocus.
*lzK'; lozenge of Kill lz-1: see rstl
origin: Spontaneous. *lzl: lozengelike
discoverer: Kiil, 45kl4. location: 1-11.
references: 1946, DIS 20: 66. discoverer: Oliver, 29k24.
phenotype: A less extreme allele of Iz. Some references: 1935, DIS 3: 28.
females fertile. RK1. phenotype: Eyes rough. Both sexes fertile. RK3.
*/xM58« lozenge of Meyer other information: Possibly an allele of rg (1-11.0).
discoverer: Meyer, 58k.
phenotype: Eyes small and oval; surface glossy;
color brownish. Tarsal claws missing. Homozy-
gous females moderately fertile, although sperma-
thecae absent; lzM5*/lz* also fertile. RK1.
Iz*: /oxenge-spectacled
discoverer Patterson, 1928.
synonym: ape1.
references: Patterson and Muller, 1930, Genetics 15:
495-577.
Green and Green, 1956, Z. Induktive Abstammungs-
Vererbungslehre 87: 708—21,
phenotype Eye size reduced; narrower titan normal;
no true facets and whole eye has glossy surface; m: miniature
color yellow-brown with darker rim, creamy in com- From Morgan and Bridges, 1916, Carnegie Inst. Wash.
bination with v. Tarsal claws vestigial. Homozy- Publ. No. 237.
gous females lack spermathecae and parovaria and
are sterile. lza/+ females tend to have abnormal m: miniature
parovaria (Anderson, 1945, Genetics 30: 280—96), location: 1-36.1.
RK1. origin: Spontaneous,
other information: Located in the leftmost (spe) Iz discoverer Morgan, lOh.
•ubloctts. lz*/lz$ provided probably the first re- references: Morgan and Bridges, 1916, Carnegie
corded case of intz*-*llelic recombination (Oliver, Inst. Wash. Publ. No. 237: 26 (fig.).
MUTATIONS 151
phenotype: Wing size reduced; only slightly longer references: Burdick, 1961, DIS 35: 45.
than abdomen and with normal proportions. Angle phenotype: Like m, but females poorly fertile. RK2.
between L2 and L5 reduced. Wings dark gray and other information: Recombines with Df(l)m2S9'4 to
less transparent than normal. Wing cells smaller its left and with m, mD, and the dy alleles to its
than normal (Dobzhansky, 1929, Arch. Entwick- right (Dorn and Burdick, 1962, Genetics 47: SOS-
lungsmech. Organ. 115: 363—79). In poor cultures, IS).
wings may become divergent and stringy. Cell 60
expansion inhibited in prepupae and pupae origin: Gamma ray induced.
[Waddington, 1940, J. Genet. 41: 75-139 (fig.)]. discoverer: Ives, 601.
Different m mutants complement slightly; m/ dy is synonym: m601.
wild type. RK1. references: 1961, DIS 35: 46.
cytology: Locus probably lies in 10E1-2 and extends phenotype: Like m. RK1.
to the right for a short distance. Salivary chromo- other information: Recombines with Df(l)m2S9~4 to
some studies by Demerec and Sutton show the its left but has not been extensively tested for
locus to lie in region 10C3 to 10E2 (Demerec, recombination with other m alleles.
Kaufmann, Fano, Sutton, and Sansome, 1942,
Carnegie Inst. Wash. Year Book 41: 191); however, origin: Gamma ray induced.
Dorn and Burdick (1962, Genetics 47: 503-18) discoverer: Ives, 61e.
showed that Dt(l)m259-4 (the basis for the Demerec synonym: m61a.
and Sutton location) is deficient for only part of references: 1962, DIS 36: 38.
the m locus, and some alleles are located by re- phenotype: Like m. RK1.
combination to the right of the deficiency. m2S9-4
other information: The miniature-dusky region has origin: X ray induced.
been divided into four recombinationally separable discoverer: Demerec, 33i.
sites (Dorn and Burdick 1962); m occupies the synonym: Df(l)m259-4.
third site from the left. No reverse mutations phenotype: Heterozygote with m has miniature
among 2 X 10 s progeny of m/m females (Gagne). phenotype. Lethal and cell lethal. RK2A.
cytology: Associated with Df(l^S9-4 - Df(l)10C2-
3;10E2-3 (Demerec).
dy61
other information: This deficiency must be for only
dy58 part of the m region since it recombines with m,
,60 dy m59, and mD, all of which are to its right (Dorn
Dffljm 259-4 S9 and Burdick, 1962, Genetics 47: 503—18).
I mD: miniature-Dominant
origin: X ray induced (discovered as a mosaic).
,57.
discoverer: Slatis, 48kl7.
Map of the m-dy region Slatis and Willermet, 1954, Genetics 39: 45—58
Drawn from Dorn and Burdick, 1962, Genetics 47: 503 (figO-
18. phenotype: Wings of homozygote smaller than m/m.
m2 mD/+ wings intermediate between homozygote and
origin: X ray induced in In(l)dl-49. wild type. Viability 20—50 percent normal in males
discoverer. Glass, 1929. and 5 percent in homozygous females; most die in
references: 1935, DIS 4: 9. embryo. Fertility low in homozygous females. Wing
phenotype: Like m. RK1A. size of m D /m and mD/dy intermediate between
other information: Has not been separated from mD/+ and mD/mD. RK2.
In(l)dl-49. other information: Recombines with m alleles to its
left but not dy alleles (Dora and Burdick, 1962,
origin: X ray induced. Genetics 47: 503-18).
discoverer:. Mayo, 57i. mK: miniature of KriYshenko
synonym: m^?*. origin: X ray induced.
references: 1958, DIS 32: 82. discoverer: Krivshenko, 5513.
phenotype: Like m. RK1. references: 1956, DIS 30: 75.
other information: Re combines with dy, dySSkf (fy61 phenotype: Wings thin textured, smaller than normal,
and mD but not with m, m59, m60, or Df(l)maS9-4. sometimes crumpled, with tips bent slightly upward
May be a submicroscopic rearrangement. No or downward. Sometimes, fly has m phenotype.
reverse mutation among 2 X 105 progeny of m^/m female varies from m-like to nearly normal.
mS7/m57 females (Gagne). Viability and fertility high. RK2A.
*» cytology: Associated with In(I)mK În(t)10E;20B,
origin: Spontaneous, mps: miniature-Penn State
discoverer: Krawinkel, 59a. origin: Gamma ray induced.
synonym: m^9m. discoverer: Keller and Nash.
references: 1960, DIS 34: 51. Df(l)N264-73 = Df(l)3C3-4;4C6-7 versus slight M

1961, DIS 35: 47. phenotype \M(1)3E] of Df(l)N264-42 = Df(l)3C4-
phenotype: Like mD. Homozygous females produce 5;4B4-6.
very few progeny. RK2. *M(l)30
m-Hke: see dy3id location: 1-14 (to the left of cv).
M-prd: see T(l;4)M-pro origin: Spontaneous.
discoverer: Schultz.
M: Minute references: 1929, Genetics 14: 366-419.
A class of genes lethal in homozygous or hemizy- cytology: Associated with Di(l)M-30; breakpoints
gous condition, producing smaller (short, fine) unknown. Placed in region 5D3-7B2, on the basis
bristles and increasing developmental time in heter- of the Minute phenotype of Df(l)ct2*8-37 =
ozygotes. Heterozygotes often exhibit secondary Dt(l)5D2-3;7B2-3.
effects such as small body size, large and some- M(l)34i28: see M(1)OSP
what rough eyes, missing aristae, thin-textured M(l)36f: see M(l)n3 6
wings with tendency to plexus venation, missing M(l)Bld: Minuted) Blond
bristles (usually postverticals), and low fertility, location: 1-0.1.
especially in females. Certain Minutes increase origin: Synthetic.
somatic crossing over (Stern, 1936, Genetics 21: discoverer: Patterson.
625—730). Most Minutes enhance dominance of such synonym: Vi: Viability.
venation characters as px and net or of such bristle references: 1932, Z. Induktive Abstammungs-
characters as sc. Complementary dominant lethal Vererbungslehre 60: 125-36.
effects are frequent, in combination with Dl, J, and Stern, 1936, Genetics 21: 630.
occasionally D. Recessive to two wild-type alleles phenotype: Extreme Minute of low viability. In
in triploids; lethal when two doses are present. Patterson's work, the nonappearance of Minutes
K. C. Atwood has suggested that Minute loci are the led him to postulate a factor for viability (Vi).
sites of synthesis of soluble or transfer RNA. He Stern (1936, Genetics, 21: 625-730) found it in-
argues that the best estimate of the number of creased frequency of somatic crossing over.
Minute loci agrees with the probable number of dif- RK3A.
ferent soluble RNA types; furthermore, the slow cytology: Locus in region 1B11 to 1C2-3 (Demerec,
rate of development and the weakness of the M/+ Kaufmann, Fano, Sutton, and Sansome, 1942,
fly is a reasonable manifestation of the reduced Carnegie Inst. Wash. Year Book 41: 191). Associ-
rate of protein synthesis that might be expected to ated with the 2RDXp element of T(l;2)Bld =
result from decreased production of a particular T(1;2)1C3-4;6OB12-13 when the X&2RP element is
transfer RNA; and the lethality oiM/M is the ex- replaced by a normal second chromosome.
pected result from the absence of a transfer RNA. *M(J)k
location: 1-36.3.
M(1)3E: Minute(l) in region 3E discoverer: Bridges, 23d28.
location: 1-5. phenotype: A strong Minute. Male lethal. RK2.
discoverer Demerec, 1938.
references: Demerec, Kaufmann, Fano, Sutton, and
Sansome, 1942, Carnegie Inst. Wash. Year Book
41: 191.
phenotype: Slight Minute; barely distinguishable
from wild type. RIGA,
cytology: Found and located in salivary chromosome
bands 3E3-4, on basis of slight M phenotype of
females heterozygous for Dîy>]264-76 « Di(i)3B4-
Cl;3E4-5 and non-Af phenotype of females hetero-
zygous for DffiyV*6*-** = Df(l)2D3-4;3E2-3 and
7 m Dt(l)3A6~7;3E2-3.
: Minutc(l) in region 4BC

location: 1-6.8.
discoverer: Demerec, 1938.
references: Demerec, Kaufmann, Fano, Sutton, and
41: 191.
phenotype: Strong Minute; easily distinguishable
frota wild type. RK.2A.
cytology: Found and located in salivary gland chro-
mosome region 4B5 through 4C6, on basis of ex- M(l)n: Minuted) n
trero© M phettotype of females heterozygous for Edith M. Wallace, unpublished.
MUTATIONS 153
location: 1-62.7. location: 2-(not located).

origin: Spontaneous. discoverer: Schultz.
discoverer: Bridges, 1923. references: 1929, Genetics 14: 366-419.
references: Morgan, Sturtevant, and Bridges, 1924, phenotype: Moderate Minute. Survives in combina-
Carnegie Inst. Wash. Year Book 23: 231-36. tion with M(2)e, and M(2)l2. RK2.
Bridges, 1925, Proc. Natl. Acad. Sci. U.S. 11: cytology: Occurred in chromosome with In(2R)Cy.
701-5. M(2)33et: s e e M(2)c33*
phenotype: Heterozygous females have Minute *M(2)33d
bristles. Lethal in males. Viability and fertility location: 2- (not located).
low. Pupation delayed about 42 hr at 25°C origin: X ray induced in In(2L)Cy +In(2R)Cy.
(Brehme). Wing cells smaller than normal (Brehme, discoverer: Oliver, 33dl4.
1941, J. Efcptl. Zool. 88: 135-60). Increases so- references: 1939, DIS 12: 48.
matic crossing over in X chromosome (Stern, 1936, *M(2)34b
Genetics 21: 625-730). RK2. location: 2- (not located).
origin: X ray induced in In(2L)Cy +In(2R)Cy.
origin: Spontaneous in attached X. discoverer: Oliver, 34b3.
discoverer: Curry, 36flO. references: 1939, DIS 12: 48.
synonym: M(l)36f. *M(2)34d
phenotype: A slight Minute. RK3. origin: X ray induced in In(2L)Cy +In(2R)Cy.
other information: Allelism inferred from location of discoverer: Oliver, 34d25.
M(l)n36 at 62. references: 1939, DIS 12: 48.
phenotype: Associated with rough eye variegation.
location: 1-56.6. RK3.
origin: Spontaneous. *M(2)34k
discoverer Bridges, 24b4. location: 2- (not located).
phenotype: Heterozygous females have Minute origin: X ray induced in In(2L)Cy +In(2R)Cy.
bristles. Normal in combination with the duplica- discoverer: Oliver, 34k22.
tion for 15 through 16A7 formed by combining the references: 1939, DIS 12: 48.
distal portion of the X from TCl^B3 = T(1;4)15F9- *M(2)38b
16A1;16A7-B1;1O2F with the proximal portion of location: 2-57.
the X from T(l;4)l-vll = T(l;4)15;101 (Von Halle). origin: Spontaneous.
Increases somatic crossing over (Kaplan, 1953, discoverer: Curry, 38bl8.
Genetics 38: 630-51). Lethal in males. RK2. phenotype: Extreme Minute with small bristles and
cytology: Demerec and Sutton place locus between compact body. Viability varies with modifiers.
15B1.-2 and 15E7 (Demerec, Kaufmann, Fano, M(2)38b/stw is non-sttv; M(2)38b/M(2)p viable.
Sutton, and Sansome, 1942, Carnegie Inst. Wash. RK3.
Year Book 41: 191). *M(2)38k4
M(1)OSP: Minute(l) o of Spencer location: 2- (not located).
origin: Spontaneous as mosaic female. origin: Spontaneous.
discoverer. Spencer, 34i28. discoverer Mossige, 38k4.
synonym: M(l)34i28; M(l)Sp. phenotype: Medium Minute. RK2.
references: 1935, DIS 3: 28. *M(2)40c
1937, DIS 7, 14. location: 2-65.
phenotype: Moderate Minute. Normal in combination origin: Spontaneous.
with the duplication for 15 through 16A7 formed by discoverer. Ives, 40c.
combining the distal portion of the X from references: 1941, DIS 14: 39.
T(1;4)BS = T(1;4)15F9-16A1;16A7-B1;1O2F with phenotype: Medium Minute with probable eye effect.
the proximal portion of the X from T(l;4)l-vll = RK2.
T(l;4)15;101 (Von Halle). Lethal in males. RK2. other information: Crossing over normal.
M(l)Sp: see M(l)oSp M(2)50j: see M(2)S2soj
M(2)U5: seeM(2)c*3*
M(2)21Cl-2: Minufe(2) in 21C1-2 M(2)173
location: 2-0.0. location: 2-92.3.
phenotype: Extreme Minute. RK2A. discoverer: Csik.
cytology: Placed in 21C1-2, on basis of inclusion in references: 1930, Magy. Biol. Kut. Int. Munk.
Dt(2L)al = Dt(2L)21B8'Cl;2lC8-Dl but not in (Tihany) 3: 438-53.
Dt(2L)S5 »Dt(2L)21C2-3;22A3'4 (Lewis, 1945, Gottschewskl, 1935, DIS 4: 15.
Genetics 30: 137-66). phenotype: Moderate Minute. RK2.
other information: Exists only as Df(2L)al and var- cytology: Salivary chromosomes apparently normal
ious aneuploid deficiencies for the tip of 2L. (Bridges).
U(2)At s e e M(2)c other information: Useful as balancer for recessive

( male-sterile genes in the second chromosome.
location: 2-87.5. Crossing over probably reduced.
discoverer Bridges, 19k22. M(2hs: Minute(2) e of Schultz
references: Morgan, Bridges, and Sturtevant, 1925, origin: X ray induced.
Bibliog. Genet. 2: 231. discoverer: Schultz, 34k21.
phenotype: Bristles extremely small. Abnormal synonym: M(2)S11.
abdomen effects in 90 percent of females and 40 phenotype: Bristles almost normal. Not late
percent of males. RK2. hatching. RK3.
other information: First Minute found in chromosome other information: Allelism to M(2)e inferred from
2. location of M(2)e at 40±5 and M(2)e$ at 43.
M(2)B: s e e M(2)zB
location: 2-108 [based on location discoverer: Bridges, 24116.
discoverer: Sturtevant, 20a7. synonym: M(2)t.
synonym: M(2)a. phenotype: Medium Minute. RK2.
references: Morgan, Bridges, and Sturtevant, 1925, other information: Allelism to M(2)e inferred from
Bibliog. Genet. 2: 231. location of M(2)e* at 46±5.
Bridges, 1937, Cytologia (Tokyo), Fujii Jub. Vol. M(2)H: Minute(2) from T(Y;2)H
2: 745-55. location: 2-53.5 [based on location of M(2)H.S^; be-
phenotype: Extreme Minute. Very late hatching. tween M(2)m and It].
Low fertility. RK3. origin: Synthetic.
cytology: Placed in salivary gland chromosome discoverer: Schultz.
region 60E3-11, on basis of Dt(2R)M-c33a = references: Morgan, Bridges, and Schultz, 1931,
Df(2R)60E2'3;60El 1-12. Carnegie Inst. Wash. Year Book 30: 408—15.
Morgan, Bridges, and Schultz, 1935, Carnegie Inst.
discoverer: Schultz, 33a7. Wash. Year Book 34: 284-91.
synonym: M(2)115; M(2)33a. phenotype: Weak Minute. RK3A.
references: Bridges, 1937, Cytologia (Tokyo), Fujii cytology: Located in salivary region 37B2 through
Jub. Vol. 2: 745-55. 4OB2, on basis of its association with deficiency
phenotype: Fairly strong Minute. Late hatching, but from T(Y;2î = T(Y;2)37Bl-2;40B2-3.
with good fertility and viability. RK2A. M(2)H*S: Minuted) H of Schultz
cytology: Associated with Di(2R)M-c33a - origin: X ray induced.
Df(2R)60E2-3;60El 1-12. discoverer: Schultz, 33a9.
other information: Allelism inferred from location of synonym: M(2)S5.
M(2)c at 107 and M(2)c33* at 108, phenotype: Medium Minute. RK2.
M(2)C: s e e M(2)zC other information: Allelism with M(2)H. inferred from
location at 53.5 and its survival in combination
with M(2XnS6 (Schultz).
location: 2-72.
discoverer: Bridges; 20b25. origin: X ray induced.
references: Bridges and Morgan, 1923, Carnegie discoverer: Schultz, 33b7.
Inst. Wash. Publ. No. 327: 231-34. synonym: M(2)S12,
Morgan, Bridges, and Sturtevant, 1925, Bibliog. phenotype: Slight Minute. Bristles nearly normal.
Genet. 2: 231. RK3.
phenotype: Heterozygote has no effect except when other information: Allelism with M(2)H inferred from
also heterozygous for M(3)d. The double heterozy- slight phenotype and inseparability from pr.
gote has Minute bristles in about 95 percent of
flies. Probably lethal in homozygote. RK3. location: 2-101.2 [based on M(2)l2].
M(2)D: s e e M(2)S2*> origin: Spontaneous.
*M(2)m references: Morgan, Bridges, and Sturtevant, 1925,
location: 2-43 [based on location of M(2)es], Bibliog. Genet. 2: 231.
origin: Spontaneous. Bridges, 1937, Cytologia (Tokyo),Fujii Jub. Vol. 2:
discoverer: Bridges, 2Ob25. 745-55.
references: Morgan, Bridges, and Sturtevant, 1925, phenotype: Fairly strong Minute. Flies eclose about
BibUog. Genet. 2: 231. 2 days late (at 25°C) because of delay in puparium
phenotype: Medium Minute with delayed hatching. formation (Dunn and Mossige, 1937, Hereditas 23:
Fifty percent of females and 10 percent of males 70—90). Eyes somewhat rough; veins often show
show abnormal abdomen effect. Most females plexus. Abdominal sclerites often abnormal. Ocelli
sterile and remainder produce few progeny, often reduced. Viability 80—90 percent wild type
RK3(A>. and fertility low. Homozygote dies in egg stage;
MUTATIONS 155
eggs recognizable by a thin chorion (Li, 1927, *M(2)mS13

Genetics 12: 1-58). RK2A. origin: X ray induced.
cytology: Associated with Df(2R)M-l = Df(2R)57Fll- discoverer: Schultz, 33b3.
58AI;58F8-59A1. Location further restricted to synonym: M(2)S13.
58F on the basis of its inclusion in Dp(2;3)P from phenotype: Small-bristled Minute with chunky body.
T(2;3)P = T(2;3)58E3-F2;60D14-E2;96B5-Cl RK2.
(Bridges, 1937). other information: Allelism with M(2)m inferred from
phenotype and location at 50.
origin: Spontaneous. *M(2)p
discoverer: Schultz, 26a7. location: 2- (to the right of msf).
synonym: M(2)l . discoverer: Bridges, 24b6.
references: Bridges, 1937, Cytologia (Tokyo), Fujii references: Curry, 1939, DIS 12: 46.
Jub. Vol. 2: 745-55. Morgan, Schultz, Bridges, and Curry, 1939,
Morgan, Bridges, and Schultz, 1937, Carnegie Inst. Carnegie Inst. Wash. Year Book 38: 273—77.
Wash. Year Book 36: 298-305. phenotype: Bristles small. Survives in combination
phenotype: Medium Minute. Puparium formation de- with M(2)e, M(2)38b, or Df(2R)M-S2v6ll. RK3(A).
layed about 13 hr at 25°C (Brehme, 1939, Genetics other information: May also have a second Minute
24: 131—61); slight delay in time of second larval factor to left of pr. Crossing over possibly re-
molt. Viability, fertility, and classification excel- duced.
lent. Homozygote lethal in first larval instar. In- M(2)pt>: see M(2)§2»
creases somatic crossing over (Kaplan, 1953, : see M(2)m*
Genetics 38: 630-51). RK2.
cytology: Salivary chromosomes apparently normal *M(2)S1: Minute(2) of Schultz
(Bridges). location: 2-15.0 (between dp and tkv).
M(2)l': see M(2)P origin: X ray induced.
*M(2)m discoverer: Schultz, 33al2.
location: 2-54. references: Curry, 1939, DIS 12: 46.
discoverer: Bridges, 23gl2. phenotype: Small-bristled Minute with heavy body.
references: Morgan, Bridges, and Sturtevant, 1925, Classification good. Viability and fertility fairly
Bibliog. Genet. 2: 231. good. RK2.
phenotype: Rather strong Minute with viability about other information: Not deficient for neighboring loci.
60 percent wild type. RK3. M(2)S2
cytology: The deficiency from T(Y;2)G = location: 2-55.1.
T(Y;2)36B5-C1;4OF with the duplication from origin: X ray induced.
T(Y;2ya = T(Y;2)37Bl-2;40B2-3 (i.e., a deficiency discoverer: Schultz, 33al2.
for 36B6 through 37B2 and from 40B3 into 40F) in references: Morgan, Bridges, and Schultz, 1938,
combination with a normal second chromosome Carnegie Inst. Wash. Year Book 37: 304—9.
produces an extreme Minute (Morgan, Bridges, and Morgan, Schultz, Bridges, and Curry, 1939,
Schultz, 1935, Carnegie Inst. Wash. Year Book 34: Carnegie Inst. Wash. Year Book 38: 273-77.
284—91). Because of its genetic location and ex- Morgan, Schultz, and Curry, 1940, Carnegie Inst.
treme phenotype, M(2)m is assumed to be the type Wash. Year Book 39: 251-55.
mutant at the locus revealed by the above defi- phenotype: Moderate Minute with relatively good
ciency, although the appropriate genetic tests can- viability. Pale body color. RK2(A).
not be made. cytology; Placed in region 41 A, on basis of
*M(2)m' Df(2R)M~S2lO = Df(2R)41A. Salivary gland chromo-
origin: Spontaneous. somes of M(2)S2 apparently normal, but locus in
discoverer: Bridges, 24128. difficult chromocentric region.
synonym: M(2)s. other information: Gives mutant phenotype in combi-
references: Schultz, 1929, Genetics 14: 366—419. nation with stw but not with Jag, It, rl, ap, tk, std,
phenotype: Medium Minute. RK2. or mat; thus genetic evidence suggests deficiency.
other information: Allelism inferred from location at
54.4, but could equally well be an allele of M(2)H. origin: X ray induced.
*M(2)mS*: Minute(2) m of Schultz discoverer: Schultz, 33a.
origin: X ray induced, synonym: M(2)S3.
discoverer: Schultz, 33al2. phenotype: Medium Minute. RK2(A).
synonym: M(2)S6. other information: May be associated with an inver-
phenotype: Medium Minute. Survives in combination sion, since there is no crossing over between b
with M(2)HS5 (Schultz). RK2. and pr.
cytology: Included in duplication from T(Y;2)G =
T(Y;2)36B5-C1;4OF but not that from T(Y;2)H » origin: X ray induced.
T(Y;2)37Bl-2;4QB2-3; thus occurs in cytological discoverer: Schultz, 33a5.
region assumed to contain M(2)m, synonym: M(2)S4.
references: Morgan, Bridges, and Schultz, 1938, phenotype: Medium Minute. RK2(A).
Carnegie Inst. Wash. Year Book 37: 304—9. other information: Gives nonmutant phenotype in
Morgan, Schultz, Bridges, and Curry, 1939, combination with rl and stw. Recombination be-
Carnegie Lost. Wash. Year Book 38: 273-77. tween pr and en reduced to 1.5 map units.
Morgan, Schultz, and Curry, 1940, Carnegie Inst.
Wash. Year Book 39: 251-55. *M(2)S2D: Minuted) of Schultz 2 in T(Y;2;3)D
pbemorype: Medium Minute bristles. Large pale body origin: X ray induced with T(Y;2;3)D.
with heavy, malformed, bloated abdomen. Female discoverer: Schultz, 1934.
fertility low. RK2A. synonym: M(2)pD; M(2)D.
cytology: Salivary chromosomes apparently normal, references: 1937, DIS 7: 14.
but the region is in chromocentric part of 2R. phenotype: Bristles almost normal. Bristle and body
ether information: Gives mutant phenotype in com- color pale. Presence of M(2)S2D enhances varie-
bination with l(2)Sp9c, l(2)Spll, l(2)Spl5, atw, and gated position effects to same extent removal of Y
up btit not with/a||, It, tl, tk, std, or mat. Thus from male. RK3A.
geaetic evidence suggests deficiency. cytology: Presumably associated with deficiency of
salivary chromosome regions 41A-C found by
origin: X ray induced. Whittinghill in T(Y;2;3p (1937, DIS 8: 82).
dlseeverer: Schulte, 33a3.
synonym: M(2)S8. ": Minuted) of Schultz 2 from
references: Morgan, Bridges, and Schultz, 1938, vestige!-11
Cft«*gi@ fast. Wasa. Year Book 37: 304-9. origin: X ray Induced; arose simultaneously with
Morgan, Schultz, Bridges, and Curry, 1939,
Carnegie lust, W«»h. Yea* Bo@k 3& 273—77. discoverer: Ruch, 1931.
phmtmtypmi Long-bristled Minute; readily classifi- synonym: M(2)v^1.
•Me. P®l© body color. Eyes often deformed; post- references: Morgan, Bridges, and Schultz, 1938,
scutellar bristle* may be erect or abs«st. RO(A), Cana«gie last. Wash. Year Book 37: 304-9.
cytology: No detectable change in salivary chromo- Morgan, Scimitar, Bridges, and Curry, 1939,
sosses, bat region, is in cbremioceotrk: part of 2M. Carnegie fast. Wash. Year Book 38: 273-77.
other information: Give® mutant pfcenotyp® in c&mtl- phenorype: Slight Minute of good viability. Pale
wdttlm with I(2pp9c, l(2)SplI, l(2)Spl5, sad ntw body color. RK2A.
bat not rl or ap. Then ges*tic evWenc* »»gf e*t» cytology: Associated with D f 2 R ) M - S 2 ^ " «
dWicie-nc?. Di(2R)40F-41Al;42Al 9-Bt.
other infamcrtt©* Originally considered to be part
®rl§ft*: X mf iodoced. ©f FJJ** pt*@»otyp@ but Bridges and Curry showed it
itmtsmmmr. Sefento, 32U31. to be separable,
M(2)S3: »ee
Leag-b«s!ied Minnie, RKJ. M(2)S4i me
ee M(2)HSs
rS^«; X ray
Setato* 32k22.
l©csrt®n: 2-77.5.
®#ij|!»s X ray iaiaced (©ce«rr©d as a «©»aic).
, and Cwtf,
, C«ittrgi« test. l%sk, T « » Botk 38- 273-
77, ftristle* very tsiall, aristae often re-
Srfcutlc, ; veaation plexasltfee. Hatches late. Via-
mbctxt 70 percent wild type, btut fariafel«.
, C«*a»gMt last, Wa»k Y««r »»r* 4§; Fertility good, toeffasrs somatic er©»«i»g over
(K»p!«Bp 1953, Ge»#tic» 3S; 630-51), Rati© of
aad total »ac!#te acid coatenl to total nitrogen content
less than normal (Altmttt, 1953, Experieotia fft
to sa-*e
Ms wlf% D f<"<•>*,far* <f»f , cyteAeffy: SaSi*ary cfere?r2%ea'*'s apparently ttenm!,

organs, v « * » g «§#»*, t® t»e right of ?2D, ce the basis ©f its te-
ia ttw duplicated s««t*^fl of chr /jros'jes* 2
* fc ««it stir cst»«ei ia nkfomrn'me 3 in T'i. Ijlo.f 7*'*, 1'??-
MQ)SJ3t: me Mtj)**
MUTATIONS 157
M(2)t: see M(2)e* M(3)33d: see M(3)h33d

M(2)vg11>: see M(3)33j: see M(3)h33j
M(2)z M(3)364: see M(3)be36e
location: 2-12.9. *M(3)39b
origin: Spontaneous. location: 3- (not located but probably in 3R).
discoverer: Schultz. discoverer: Curry, 39bl7.
references: 1929, Genetics 14: 366—419. references: 1939, DIS 12: 45.
phenotype: Medium Minute with good character- phenotype: Short-bristled Minute of low viability.
istics. About 2 days delay in puparium formation Females infertile except in mass culture. RK3.
(Dunn and Mossige, 1937, Hereditas 23: 70—90). M(3)54c
Increases somatic crossing over (Kaplan, 1953, location: 3- (rearrangement),
Genetics 38: 630-51). RK2. origin: Neutron induced.
cytology: Located between 24E2 and 25A2, based discoverer: Mickey, 54clO.
on its inclusion in Df(2L)M-zB = Df(2L)24E2- references: 1963, DIS 38: 29.
Fl;25Al-2 (Morgan, Schultz, Bridges, and Curry, cytology: Associated with In(3LR)M-54c =
1939, Carnegie List. Wash. Year Book 38: 273— In(3L)73A9-10;75D7-El + In(3LR)61C2-3;80C4-
77). 5;93B4-5;100B8-9.
other information: Carries dp + and tkv+. M(3)124: seeM(3)wi24
M(3)B: see M(3)wB
: Minute(2) z of Bridges M(3)B2; see M(3)wB2
origin: Spontaneous. *M(3)bb
discoverer: Bridges, 38dl2. location: 3- (not located).
synonym: M(2)B. origin: Spontaneous.
references: Morgan, Schultz, Bridges, and Curry, discoverer: Mossige.
1939, Carnegie Inst. Wash. Year Book 38: 273 references: 1946, DIS 20: 68.
77. phenotype: Medium Minute. RK2.
Curry, 1939, DIS 12: 46.
Curry, 1941, DIS 14: SO. *M(3)be: Minute(3) beta
phenotype: Medium Minute. RK2A. location: 3-84.5 [based on location of M(3)be36e]m
cytology: Associated with Df(2L)M-zB = origin: Spontaneous.
Df(2L)24E2-Fl; 25A1-2. discoverer: Stern, 26a20.
references: 1927, Naturwissenschaften 15: 740—46.
*M(2)zC: Minute(2) z of Curry 1934, DIS 1: 35-36.
origin: Spontaneous, phenorype: Medium Minute of excellent viability.
discoverer: Curry, 37g27. Increases somatic crossing over (Stern, 1936,
synonym: M(2)C. Genetics 21: 625-730). RK2.
references: Morgan, Bridges, and Schultz, 1938,
Carnegie Inst. Wash. Year Book 37: 304-9. origin: Spontaneous.
Morgan, Schultz, Bridges, and Curry, 1939, discoverer Bridges, 36e22.
Carnegie Inst. Wash. Year Book 38: 273—77. synonym: M(3)36e.
phenotype: Fairly strong Minute. Late hatching. phenotype: Medium Minute. Good viability and fer-
Eyes rough. Viability and fertility low. RK2A. tility. Wing shows plexus effect along vein L2 and
cytology: Associated with Df(2L)M-zc = at posterior cross vein. RK2.
Dt(2L)24D2-5;25A2-3 (Bridges). other information: Allelistn to M(3)be based on the
location of M(3)be at 87± and M(3)be36* at 84.5.
location: 3-101.0. *M(3)d
discoverer. Bridges, 19b8. discoverer: Bridges, 20b25.
references: Bridges and Morgan, 1923, Carnegie references: Bridges and Morgan, 1923, Carnegie
Inst. Wash. Publ. No. 327: 206—7 (fig.). Inst. Wash. Publ. No. 327: 231.
phenotype: Bristles slender and shorter than wild phenotype: Part of digenic Minute. Produces no
type. Somewhat late hatching. M(3)l/M(3)g sur- effect except when M(2)d is also heterozygous.
vives. RK2. Homozygote probably lethal. RK3.
other information: First Minute found. *M(3)f
M(3)6: see location: 3-105.
discoverer: Bridges, 20i9.
location: 3- (not located), references: Bridges and Morgan, 1923, Carnegie
origin: X ray induced, Inst. Wash. Publ. No. 327: 236.
discoverer: Oliver, 32122. phenotype: Small bristles. Poorly viable; females
references: 1939, DIS 12: 48. infertile. RK3.
other information: Permits no crossing over with M(3)f: see M(3)S35{
3-p!@ except In spindle-fiber region. M(3)Fla: see
other information: Allelismbased on lethal interac-

location: 3-106.2. tion with M(3$iy (Von Halle).
origin: Spontaneous in M(3)d strain. *Ul(3)hS38
discoverer: Bridges, 20i27. origin: X ray induced.
references: Bridges and Morgan, 1923, Carnegie discoverer: Schultz, 33a 12.
Inst. Wash. Publ. No. 327: 236. synonym: M(3)S38.
phenotype: Very slight Minute. Stronger in presence phenotype: Rather extreme Minute. Wings flimsy,
of an enhancer located near p. Survives in combi- with plexus effect along vein L2 and at posterior
nation with M(3)l. RK3. crossvein. Low viability and fertility. RK3.
cytology: Schultz found it to be in the deficiency for other information: Allelism inferred from location
the tip of 3R from T(3;4)d (cytology not recorded); 3.6 units to the left of st (Schultz).
Dobzhansky (1930, Genetics IS: 347-99) claimed
otherwise. origin: Spontaneous.
other information." May be same as M(3)d. discoverer Bridges, 25dl8.
synonym: M(3)v.
location: 3-40.2 (to the left of D). phenotype: Medium Minute. RK2.
origin: Spontaneous as a mosaic male. other information: Allelism based on lethal interac-
discoverer: P. R. Sturtevant. tion with M(3)h.
Inst. Wash. Publ. No. 327: 244. origin: Spontaneous.
Coyne, 1935, DIS 4: 59. discoverer: Sturtevant, 25gl9.
Mossige, 1938, Hereditas 24: 110-16. synonym: M(3)y.
phenotype: Medium Minute. Delayed about 2 days in references: Stern, 1927, Naturwissenschaften 15:
puparium formation (Dunn and Mossige, 1937, 740-46.
Hereditas 23: 70-90). M(3)h/Ly survives. RK2. Mossige, 1938, Hereditas 24: 110-16.
phenotype: Medium Minute. Good viability and fer-
tility. Increases somatic crossing over in X (Stern,
origin: Recovered among progeny of heat-treated 1936, Genetics 21: 625-730). RK2.
flies. other information: Allelism based on lethal interac-
discoverer: Ives, 33d30. tion with
synonym: M(3)33d.
references: Plough and Ives, 1934, DIS 1: 33. location: 3-28.9.
1935, DIS 2: 35. discoverer: Bridges, 23d23.
phenotype: Bristles thin and short. Eyes small and phenotype: Medium Minute of good viability. RK2.
rough. Wings broad and waxy. Viability poor. *M(3)ii
RK3. discoverer: Bridges, 24b28.
other information: Allelism inferred from location at synonym: M(3)q.
41. phenotype: Extreme Minute. Very late hatching.
Poor viability; females infertile. RK3.
origin: X ray induced. other information: Allelism to M(3)i inferred from its
discoverer: Demerec, 33j25. published position of 30±10.
synonym: M(3)6; M(3)33j.
Coyne, 1935, DIS 4: 59. discoverer: Schultz, 33a6.
Mossige, 1938, Hereditas 24: 110-16. synonym: M(3)S33.
phenotype: Medium Minute. Prolongs larval life 2 references: 1940, DIS 13: 51.
days at 25°C (Dunn and Mossige, 1937, Hereditas phenotype: Extreme Minute. Females usually
23: 70-90). Good viability and fertility. sterile. RK3.
M(3)h33}/Ly and M(3)h33i/M(3)ti are lethal. In- other information: Allelism with M(3)i inferred from
creases somatic crossing over in X (Stern, 1936, its published location near ft.
Genetics 21: 625-730). RK2A.
cytology: Genetic data on lethal interaction with location: 3-90.2.
both M(3î and Dt(3h%,y = Df(3L)70A2-3;70A5-6 discoverer: Bridges, 23dl2.
suggest that M(3]h33i is a deficiency that includes phenotype: Extreme Minute. Late hatching. Fe-
bands in 70A. males sterile or of low fertility. RK3(A).
other information: Lethal in combination with
Mf3)ri«7; Mhute(3) h of Schultz 1(3)PR; possibly a deficiency.
origin: X ray induced. *M(3)J*P: Minute(3) j of Spencer
discoverer- Schultz, 33al2. origin: Spontaneous.
synonym: M(3)$37. discoverer: Spencer, 36c21.
phenotype: Extreme Minute with fine bristles and synonym: M(3)Sp.
small body. RK3. references: 1937, DIS 7: 14.
MUTATIONS 159
phenotype: Extreme Minute with very small bristles. M(3)v: see

Wings broad with plexus of veins. Abdominal
bands somewhat abnormal. Female almost never location: 3-79.7.
fertile and then only sparingly. Male has fair via- discoverer: Schultz, 1925.
bility and fertility. RK3. references: Stern, 1927, Naturwissenschaften 15:
other information: Allelism with M(3)j inferred from 745.
phenotype and published location of 90+10. Schultz, 1929, Genetics 14: 366-419.
: see M(3)ii phenotype: Strongly reduced bristles. Good viability
M(3)S31: Minute(3) of Schultz and fertility. Delays puparium formation about 42
location: 3-50.0. hr at 25°C; first and second instars also delayed
origin: X ray induced. (Brehme, 1941, Growth 5: 183-95). Homozygote
discoverer: Schultz, 33alO. dies in first instar. M(3)w enhances L, B (Dunn
references: 1940, DIS 13: 51. and Coyne, 1935, Biol. Zentr. 55: 385-89), Bx3,
phenotype: Fine-bristled Minute of medium viability. Co, fa, ap4, Jag, Ser, Ly, and apXt» (Bryson, 1940,
RK3(A). Genetics 25: 113). Treanor (1962, Ph.D. Thesis,
other information: Gives mutant interaction with cu Univ. Buffalo) suggests that the mitochondria 1
but not ma. May reduce crossing over in sf-sr membrane is defective and labile phosphate forma-
region. tion is disturbed. Recovery of M(3)w from
M(3)S32 M(3)w/In(3R)C, l(3)a mothers is reduced (Schultz).
location: 3- (not located). RK2.
discoverer: Schultz, 33a5. discoverer: Csik.
phenotype: Medium Minute. Most flies thickset. synonym: M(3)124.
RK3. references: 1930, Magy. Biol. Kut. Int. Munk.
M(3)S33: s e e M(3)is3 3 (Tihany)3: 438-53.
M(3)S34 Gottschewski, 1935, DIS 4: 15.
location: 3-44.3 [3.2 units to the left of Dfd phenotype: Bristles shorter than average Minute.
(Schalet, 1960)J. Hatching later. Viability good. RK2.
origin: X ray induced. other information: Allelism based on lethal interac-
discoverer: Schultz, 33a6. tion with M(3)w, M(3)wB, M(3)wB^, and M(3)wFl*.
references: Schalet, 1960, DIS 34: 55. : Minute(3) w of Burkarf
phenotype: Slight Minute. Overlaps wild type. In discoverer: Burkart.
existing lines bristles appear normal, but reces- synonym: M(3)B.
sive lethal effect at 44.3 remains. RK3. references: 1935, DIS 4: 15.
*M(3)S35 phenotype: Moderate Minute. Good viability and
location: 3-64. fertility. RK2(A).
origin: X ray induced. other information: Interacts lethally with l(3)a
discoverer: Schultz, 33all. (Bridges), whereas M(3)w does not (Schultz). Pos-
phenotype: Extreme Minute with small body. RK3. sibly indicates that M(2)wB is a deficiency. Alle-
*M(3)S35f lism based on lethal interaction with M(3)w,
discoverer: Moriwaki, 38f2. (3)wB2t a n d M(3)wF**.
synonym: M(3)L ; Minute(3) w of Bridges
references: 1939, DIS 12: 50. discoverer. Bridges, 38c6.
phenotype: Minute bristles. RK2. synonym: M(3)B^.
other information: Allelism to M(3)S35 based on its phenotype: Bristles quite small. Body size reduced.
location at 62.4. Medium late hatching. RK2.
M(3)$36 other information: Allelism based on lethal interac-
location: 3- (not located). tion with M(3)w, M(3)w124, M(3)wB, and M(3)wFl*.
origin: X ray induced. M(3)wFI°: Minute(3) w-F/orit/o
discoverer: Schultz, 32k26. discoverer. Mossige, 35d.
phenotype: Variable phenotypes appear in stock; synonym: M(3)Fta.
Minute and variegated for ss-like. Not studied. references: Bryson, 1937, DIS 7: 18.
RK3. 1939, DIS 12: 50.
M(3)S37: see M(3)h*3 7 phenotype: Strongly reduced bristles. Good viability
M(3)S38: s e e M(3)hsss and fertility. M(3)Fta/+ females form puparia at
*M(3)S39 129 hr after hatching; 41 hr after in wild type.
location: 3-47. Larval molts also delayed to a lesser extent
origin: X ray induced. (Brehme, 1940, Genetics 26: 141), RK2.
discoverer: Schultz, 33a3. other information: Allelism based on lethal interac-
phenotype: Extreme Minute with small body. Low tion with M(3)w, M(3)w*2*, M(3)wB, and M(3)wB*.
viability and fertility. RK3. M(3)x: Minuto(3) with C(3)x
M(3)Sp; s e e M(3)JSP location: 3- (on the left arm).
160 GENETIC VARIATIONS OF DRO5OPHILA MELANOGASTER
origin: Spontaneous in In(3L)P. divergent or upheld. Development severely re-

discoverer: Muller, 1929. tarded; viability low. RK3A.
phenotype: Rather extreme Minute; expression re- cytology: Associated with Df(4)M63e =
duced by H. RK3A. Df(4)10lE;102D13-El (Fahmy).
M(3)y: s e e M(3)hv
origin: Gamma ray induced.
mlocation: 4-0. discoverer: Fahmy, 62f.
origin: Spontaneous. phenotype: Medium Minute. Development slightly
discoverer: Bridges, 25128. retarded; viability good. RK2A.
references: 1935, Biol. Zh. (Moscow) 4: 401-20. cytology: Associated with Di(4)M62t —
1935, Tr. Dinam. Razvit. 10: 463-74. Dt(4)101E;102B10-17 (Fahmy); Df(4)101E-
phenotype: Medium Minute. Viability good; develop- F;102B2-5 (Hochman).
ment only slightly delayed. M(4)/+/+ triplo-fours M(4)63°
are non-Minute (Mohr, 1933, Hereditas 17: 317-32). origin: Recovered from progeny of male injected
Homozygotes die in embryonic stage (Farnsworth, with thymus extract from leukemic mice (Gross
1951, Genetics 36: 550). RK2A. Factor).
cytology: Placed in salivary gland chromosome sec- discoverer: Fahmy, 63a.
tion 1O1F2-102A5, on the basis of Df(4)M63B - phenotype: Medium Minute. Development slightly
Di(4)101F2-102Al;102A2-S (Fahmy and Hochman). retarded; viability good. RK2A.
Associated with Df(4)M = Df(4)101E-F;102B6-17. cytology: Associated with Df(4)M63a = Df(4)101F2-
*M(4)2 102Al;102A2-5 (combined from observations of
origin: X ray induced. Fahmy and Hochman).
discoverer: Schultz, 32k29. m(B): see su(B)
references: Bridges, 1935, Biol. Zh. (Moscow) 4: m(g): see e(g)
401-20. ma: maroon
phenotype: Like M(4) but more extreme; a slight location: 3-49.7.
grooveless phenotype. Viability and fertility lower origin: Spontaneous.
than M(4). RK2A. discoverer: Bridges, 12cl3.
other information: Gives mutant phenotype in combi- references: 1918, Proc. Natl. Acad. Sci. U.S. 4:
nation with at, d, and ciD; therefore probably a 316-18.
deficiency. Bridges and Morgan, 1923, Carnegie Inst. Wash.
Publ. No. 327: 53 (fig.).
origin: X ray induced. phenotype: Eye color dull ruby, approaching wild
discoverer Schultz, 33a8. type with age; classification slow. Larval Mal-
phenotype: Similar to M(4). RK2A. pighian tubes pale yellow (Beadle, 1937, Genetics
cytology: Associated with Dt(4)M3, which looks like 22: 587—611). Eye color autonomous in transplant
Df(4)M (Bridges, 1935, Tr. Dinam. Razvit. 10: into wild-type host (Beadle and Ephrussi, 1936,
463-74). Genetics 21: 230). Eyes contain 31 percent normal
red pigment and 59 percent normal brown pigment
origin: X ray induced. (Nolte, 1955, J. Genet. 53: 1-10). RK2.
discoverer: Glass, 42hl2. ma49d
phenotype: Like M(4). RK2A. discoverer: Oftedal, 49d.
other information: Gives mutant interaction with d references: 1951, DIS 25: 69.
and Ce and therefore probably associated with a phenotype: Eye color like bw, darkening with age.
deficiency. RK1.
*Ma: Mo dominigene
H(4)S79 location: 1- (not located).
discoverer. Gloor and Green, 1957. discoverer: Goldschmidt, 1935.
references: Hochman, Gloor, and Green, 1964, references: Gardner, 1942, Univ. Calif. (Berkeley)
Geoetica 35: 109-26. Publ. Zool. 49: 95.
phenotype: Like M(4). RK2. phenotype: In combination with Ma, vg/+ is strongly
cytology: Salivary chromosomes apparently normal scalloped. RK3.
(Hochman). macro fine: s e e mf
other information: No interaction with ci or ci°. mah: mahogany
location: 3-88.
origin: Recovered among progeny of male injected discoverer: Beadle, 36b26.
with homologous DNA. references: Beadle and Ephrussi, 1937, Am.
discoverer. Fahmy, 62e. Naturalist 71: 91-95.
phenotype: Small fly with extremely Minute bristles. phenotype: Eye color translucent brown in young
Eyes large and slightly rough. Wings frequently flies, changing toward wild type and becoming dark
MUTATIONS 161
brown with age. Eyes contain 77 percent normal references: 1961, DIS 35: 47.
red pigment and 102 percent normal brown pigment 1963, DIS 38: 82.
(Nolte, 1955, J. Genet. 53: 1-10). Larval Mal- phenotype: Male lethal. RK3A.
pighian tubes wild type in color (Beadle, 1937, other information: Shows mutant interaction with
Genetics 22: 587-611). RK3. sw, mal, su(f), at least one lethal locus left of sw,
mal: maroon I ike and at least five lethal loci between mal and su(f)
location: 1-64.8 (Schalet, 1963, DIS 38: 82). but not bb. Therefore, associated with a defi-
origin: X ray induced. ciency.
discoverer: Oliver, 3011. *mal60
references: 1935, DIS 3: 28. origin: Induced by DNA.
phenotype: Eye color brownish purple. Larval Mal- discoverer: Fahmy, 60j.
pighian tubes short, bloated, and irregularly synonym: malbz60L
formed; contain yellow to orange pteridine globules phenotype: Resembles mal. Noncomplementing with
(Schwinck, 1960, DIS 34: 105). Lacks detectable mal and malbz. RK3.
amounts of xanthine dehydrogenase and the malbx: maroon like-bronzy
products of its activity, uric acid and isoxanthop- origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
terin (Forrest, Glassman, and Mitchell, 1956, phenylalanine (CB. 3007).
Science 124: 725—26; Glassman and Mitchell, discoverer: Fahmy, 1954.
1959, Genetics 44: 153—62; Hubby and Forrest, synonym: bz: bronzy.
1960, Genetics 45: 211—24). Accumulates the references: 1958, DIS 32: 68.
enzyme's substrates (Mitchell, Glassman, and phenotype: Morphologically and biochemically like
Hadorn, 1959, Science 129: 268—69). mal progeny mal. Shows maternal effect like mal. mal/malbz
of mal mothers appear normal in both eye color heterozygote appears normal in eye color and Mal-
and Malpighian tube morphology, but not chromato- pighian tube morphology, but produces only 10 per-
graphically (Glassman and Mitchell, 1959, cent normal level of xanthine dehydrogenase ac-
Genetics 44: 547—54; Glassman and McLean, 1962, tivity (Glassman and Pinkerton, I960, Science 131:
Proc. Natl. Acad. Sci. U.S. 48: 1712-18; 1810—11; Ursprung, 1961, Z. Vererbungslehre 92:
Schwinck, 1960). mal/malbx heterozygotes appear 119-25; Schwinck, 1960, DIS 34: 105). Produces
normal in eye color and Malpighian tube morphology in vitro complementation with ry (Glassman, 1962,
but show only about 10 percent the normal amount Proc. Natl. Acad. Sci. U.S. 48: 1491-97). Be-
of xanthine dehydrogenase activity and accumulate haves nonautonomously in transplants (Ursprung,
enzyme's substrates (Glassman and Mitchell, 1959; 1959, DIS 33: 174-75). RK3.
Schwinck, 1960). In vitro complementation of mal malbzeoj; see malô
and malbz has not been demonstrated, mal and ry Mal: Malformed
extracts complement to produce xanthine dehydro- location: 2- (near right end of 2R) and 4- (multi-
genase activity (Glassman, 1962, Proc. Natl. genic, according to Bridges),
Acad. Sci. U.S. 48: 1491-97); they do not comple- origin: Spontaneous.
ment intercellularly in vivo, however, since recip- discoverer: Steinberg, 36kl3.
rocal eye-disk or Malpighian-tube transplants be- references: 1937, DIS 7: 15,20.
have autonomously with respect to drosopterin for- phenotype: Heterozygote has either malformed pit in
mation (Schwinck, 1960; 1963, DIS 38: 87). mal is middle of eye or, oftener, nick at front edge of eye,
nonautonomous in mosaics with wild-type tissue with bristle or antennalike outgrowth. Penetrance
(Glassman, 1957, DIS 31: 121—22) and in trans- low; enhanced by addition of extra brewer's yeast
plants of eyes into wild-type hosts (Ursprung, to medium. Homozygote shows larger nick and an-
1961, Z. Vererbungslehre 92: 119-25). Xanthine tennal outgrowth, with 100 percent expression in pr
dehydrogenase level the same in flies with 1—3 Mal stock. RK3.
doses of nja/ + (Grell, 1962, Z. Vererbungslehre 93: male and female sterile( ): see mfs( )
371—77; Glassman, Karam, and Keller, 1962, Z. male sterile( ): see ms( )
Vererbungslehre 93: 399-403. RK3. Malformed: see Mal
other information: One allele each induced by CB. maroon: see ma
1414, CB. 3007, CB. 3025, CB. 3051, and X rays maroonlike: see mal
(Fahmy, 1958, DIS 32: 68). Mas: Masculinizer
origin: X ray induced. discoverer. Mischaikow, 581.
discoverer: Schalet, 1961. references: 1959, DIS 33: 98.
references: 1961, DIS 35: 46-47. phenotype: Heterozygous female transformed into
phenotype: Brownish red eye color like mel; does sterile malelike fly. Last abdominal segments
not complement with mal, mat3, or mmlblt, RK3. show male-type pigmentation; external genitftlia
mal3 essentially mal©, sometimes completely absent.
origin: X ray induced. Sex combs may be present, but vary in size. In-
discoverer: Schalet, 1961. ternal sex organs degenerate; ovaries and uterus
rudimentary; spermathecae seldom present. Hetero- references: 1958, DIS 32: 71.
zygous male normal. Homozygous lethal. RK2. phenotype: Thoracic hairs irregularly distributed;
other information: May be allele of tra such as traD occasionally reduced in number. Bristles small;
of Gowen. sparse on scutellunu Eyes small and rough. Wings
matt brown: see mtb ovoid and short. Tergites in female sometimes dis*
*mb: minus bar arranged. Viability and fertility good in both
location: 3-43.4. sexes, me2/mch is wild type. RK2.
discoverer: Nordenskiold, 33a30. other information: Allelism inferred from location of
references: 1934, DIS 2: 7. me2 at 52.1 and from phenotype.
phenotype: Modifies Bar in such a way that B/B re- Me: Microcephalus
sembles B/+, and B/+ appears almost wild type; location: 3-59.0 (about 0.2 unit to the right of bx).
B male modified to resemble BK Homozygous fe- origin: Spontaneous.
male highly infertile. RK3. discoverer: Bateman.
*mbs: miniature blistered references: 1944, DIS 18: 40.
location: 2-56. 1945, DIS 19: 47.
origin: Spontaneous. phenotype: Eyes of heterozygote small or absent.
discoverer: Neel, 41cl3. Scutellars curve upward. Viability and fertility
references: 1942, DIS 16: 51. good. Homozygote usually more extreme than het-
phenotype: Wings small, curled, blistered, and erozygote, but not reliably distinguishable. Via-
plexate. Bristle positions irregular, and bristles bility of homozygote varies from 100 down to 40
often bent and twisted. Viability and fertility poor. percent. RK1A.
RK3. cytology: Probably associated with a minute rear-
rangement, perhaps a tandem repeat, of one or more
bands in 89E7-11 (E. B. Lewis).
mc-tik6: see me2
*mch: minute chaetae

location: 1-52.0.
origin: Induced by methyl methanesulfonate (CB.
1540).
phenotype: Extremely short, fine bristles. Hairs and
body also small; delayed eclosion. Male viable
and fertile, mch/mc2 is wild type. RK2.
other information: One allele each induced by CB.
1246, CB. 1356, and CB. 3026.
*md: melanotic lesions
location: 3-38.0.
origin: Found in experiments using benzopyrene.
discoverer: Gowen, 1933.
phenotype: Lesions occur in many places throughout
me: microchaete head, thorax, and abdomen. RK3.
Edith W. Wallace, unpublished. *mdg: midgoid
location: 1-64.7.
me: microchaete origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
location: 1-54.0. phenylalanine (CB. 3026).
origin: X ray induced. discoverer: Fahmy, 1955.
discoverer: Demerec, 28f20. references: 1958, DIS 32: 71.
synonym: tb-53. phenotype: Small in all dimensions; frequently
references: 1935, DIS 3: 13. underpigmented. Male infertile; viability about 20
phenotype: Hairs on thorax fewer than wild type, percent wild type. RK3.
more irregular, and frequently doubled. Bristles *me: focal melanosis
smaller, more sparse on scute Hum and occasion- location: 1-29.0.
ally on head. Eyes rough. Wings ovoid and short; origin: X ray induced.
marginal bristles disarranged. Abdominal sclerites discoverer: Gowen, 1928.
ridged. RK1. references: 1934, Arch. Pathol. 17: 638-47 (fig.).
1934, Cold Spring Harbor Symp. Quant. Biol. 2:
origin: Induced by D-p-NN-dH2-chloroethyl)amino- 128-36 (fig.).
phenylalanine (CB. 3026). phenotype: Melanotic degeneration occurs at junc-
discoverer: Fahmy, 1955. tion of tibia and femur. Lethal at end of pupal
synonym: molike. stage or shortly after eclosion. RK2.
MUTATIONS 163
Me: Moire Inner margin removed to various degrees and vena-

location: 3-19.2 (to the left of jv; based on location tion abnormal. Viability good, but both sexes in-
of Me<S5d). fertile. RK2.
origin: X ray induced. other information: One allele induced by CB. 3025.
discoverer: Muller, 1929. me/: melanized
synonym: Mo. location: 1-64.1.
references: 1930, J. Genet. 22: 299-334 (fig.). origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
Glass, 1933, J. Genet. 28: 69-112 (fig.). phenylalanine (CB. 3007).
1934, Am. Naturalist 68, 107-14. discoverer: Fahmy, 1953.
phenotype: Eye has watered-silk, shimmering, iri- references: 1958, DIS 32: 71.
descent pattern owing to a ring of six flecks phenotype: Body color darker than normal, espe-
around normal fleck. Eye color brownish and trans- cially in thorax; trident pronounced. Eye color
lucent; 79 percent normal red pigment and 85 per- dull red. Wing tips frequently curve upward. Clas-
cent normal brown pigment (Nolte, 1955, J. Genet. sification rather difficult; best in young fly. Via-
53: 1—10). Larval Malpighian tubes considerably bility and fertility good in both sexes. RK3.
lighter in color than normal but mutant classifiable other information: One allele induced by CB. 3025.
with difficulty (Brehme and Demerec, 1942, Growth
6: 351—56). Contains a modifier of dominance of melanoscutellum: see msc
dp such that dp/+; Me/+ has truncated wings. melanotic lesions: see md
Classifiable in single dose in triploids (Schultz, melanotic tumor-A: see tu-bw
1934, DIS 1: 55). Homozygous lethal. Me/In(3L)P mes: messy
is viable. RK1A. location: 3-51.9.
cytology: Placed in region 64C12-65E1, on the origin: X ray induced in a kar2 chromosome.
basis of its inclusion in Df(3L)Vn ~ Df(3L)64C12- discoverer: Schalet.
D1;65D2-E1 (Mohr, 1938, Avhandl. Norske Viden- references: Schalet, Kernaghan, and Chovnick,
skaps-Akad. Oslo, I. Mat.-Naturv. Kl. No. 4: 1-7). 1964, Genetics 50: 1261-68.
Associated with In(3L)P = In(3L)63C;72El-2 phenotype: Extra head and thoracic bristles. Wings
(Morgan, Bridges, and Schultz, 1937, Carnegie inflated, turned somewhat upward and outward, and
Inst. Wash. Year Book 36: 301). shorter and broader than normal. Posterior cross-
vein with a gap or missing. Setnilethal; male con-
origin: X ray induced. siderably less viable than female; sterile.
discoverer: Moore, 1929. mes/mes2 like mes/mes but mes/mes3, mes/mes4,
references: Glass, 1933, J. Genet. 28: 69—112. mes/messt, and mes/mes*' appear normal. RK3.
phenotype: Like Me. RK1A. other information: mes locus subdivisible into two
cytology: Arose with T(2;3)Me2 (2L broken near functional units by complementation analysis; mes
centromere). placed in the left unit, on the basis of its being
wild type when heterozygous with Df(3R)ry74.
origin: Induced by ethyl methanesulfonate. "mes*
discoverer: E. H. Grell, 65d. origin: X ray induced in a kar2 chromosome.
phenotype: Like Me. Eyes brownish with watered- discoverer: Schalet.
silk effect. Tips of large bristles slightly lighter references: Schalet, Kernaghan, and Chovnick,
than wild type. Homozygote and Me6Sd/Me lethal. 1964, Genetics 50: 1261-68.
dp°lv/+; Me65d/+ occasionally has truncated wing phenotype: Similar to mes. Mutant homozygous and
tips. RK1. in combination with mes but normal in combina-
other information: Crossing over normal in 3L. tion with mes 5 , mes4, mes 5 ', and mes61. RK3.
Me6Sd apparently not associated with a gross other information: In the left complementing unit.
chromosomal rearrangement like other Me alleles. mes3
*Me*°; Mo/re of Sytko origin: X ray induced in a kar2 chromosome.
discoverer: Sytko. discoverer: Schalet.
references: Agol, 1936, DIS 5: 7. references: Schalet, Kernaghan, and Chovnick,
phenotype: Like Me. RK1A. 1964, Genetics 50: 1261-68.
cytology: Arose with T(2;3)MeSo (breaks in 2R and phenotype: Similar to mes. mes3/mes and
3R). mes3/mes2 normal; mes3/mes4, and mes3/messt,
*meg: megaoculus mes3/mes61, and mes3/ry74 mutant. RK3.
location: 1-61.9. other information: Placed to the right of mes and
origin: Induced by DL-p-NN-di-(2-chlaroethyl)amino- mes2, on the basis of its mutant interaction with
phenylalanine (CB. 3007). Dt(3R)ry74.
discoverer: Fahmy, 1954. mes4
references: 1958, DIS 32: 71. origin: X ray induced in a kar2 chromosome.
phenotype: Eyes large, abnormally shaped, and discoverer. Schalet.
rough. Wings abnormally shaped and sometimes references: Schalet, Kernaghan, and Chovnick,
extremely small. Wing surface irregularly curved. 1964, Genetics 50: 1261-68.
phenotype: Similar to mes. mes4/mes and mgt: midget

mes4/tnes2 normal; mes4/mes3, mes4/messl, location: 1-48.7.
mes4/mes61, and mes4/ry74 mutant. RK3. origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
other information: Placed to the right of mes and phenylalanine (CB. 3007).
mes 2 , on the basis of its mutant interaction with discoverer: Fahmy, 1954.
Df(3R)ry74. references: 1958, DIS 32: 71.
mes51: messy-S lethal phenotype: Small fly with delayed eclosion. Not
origin: X ray induced in a /car2 chromosome. easily classified. Male fertile, but viability about
discoverer: Schalet. 20 percent wild type. Expression more extreme in
phenotype: Lethal homozygous and in combination female and viability further reduced. RK3.
with mes61 .and Df(3R)ry74. messl/mes3 mutant; other information: One allele each induced by CB.
messi/mes4 mutant but with low viability; 3025 and X rays; two alleles induced by CB. 1506.
messi/mes and mes51/mes2 normal. RK3. mi: minus
other information: In the right complementing unit. location: 2-104.7.
mes6' discoverer: Biddle, 281.
origin: X ray induced in a kar2 chromosome. references: Bridges, 1937, Cytologia (Tokyo), Fujii
discoverer: Schalet. Jub. Vol. 2: 745-55.
phenotype: Lethal homozygous and in combination phenotype: Bristles almost as small as hairs, and
with mes5i and Df(3R)ry74. mes61/mes3 and hairs reduced in number and size. Body size
mes6{/mes4 mutant; mes6I/mes and mes61'/mes2 small. Eclosion delayed. Viability low and er-
normal. RK3. ratic. Female entirely sterile; male fertile. RK2.
other information: In the right complementing unit. cytology: Locus is in 59E1-2 of salivary gland
*Mef: Metatarsi irregular chromosome (Schultz), on the basis of its being be-
location: 2- or 3- (rearrangement). tween the right breakpoints of In(2R)bwVD^1 =
origin: X ray induced. In(2R)41B2-Cl;59E2-4 and In(2R)bwVDe2 -
discoverer: Jonsson, 56alO. In(2R)41A-B;59D6-El.
references: Liining, 1956, DIS 30: 73. *mib: miniature bristles
phenotype: First and second tarsal joints fused and location: 1-8.7.
swollen, with extra hairs. Male sex combs en- origin: X ray induced.
larged. Fully penetrant when balanced with Cy; discoverer: Fahmy, 1956.
however, Met/ss is wild type or nearly so. RK2A. references: 1959, DIS 33: 88.
cytology: Associated with T(2;3)Met. phenotype: Short, thin bristles. Body slightly darker
*mf: macrofine than normal, particularly thorax and posterior
location: 1-5.5. border of tergites. Wings occasionally upheld and
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- inner margins frequently incised. Male viable but
phenylalanine (CB. 3025). sterile. RK3.
discoverer: Fahmy, 1955. micro-oculus: see mo
references: 1958, DIS 32: 71. M/crocep/io/us: see Me
phenotype: Fly slightly smaller than normal with microchaete: see me
short, thin, bristles. Male viable and fertile. Fe- microptera: see mp
male slightly delayed in eclosion and reduced in midget: see mgt
viability. RK3. midgoid: see mdg
miniature: see m
mfs(3)G: male and female sterile of Gill minus: see mi
location: 3-59. minus bar. see mb
origin: X ray induced, Minute-producer: see T(l;4)M-pro
discoverer: Gill, 59a. minute chaetae: see men
synonym: fs(3)4S9a. MinuteO; see M( )
references: 1960, Anat. Record 138: 351. minutelike: see ml
1961, Ph.D. Thesis, Yale Univ. Mio: see Dt*'io
1962, DIS 36: 37. *mis: misproportioned
1963, J. Exptl. Zool. 152: 251-77 (fig.). location: 1-1.3.
phenotype: Oogenesis incomplete; follicles usually origin: Induced by l:4-dimethanesulfonoxybut-2-yne
cease development early in vitellogenesis (at or (CB. 2058).
before stage 9); occasional breakthrough produces discoverer: Fahmy, 1951.
adult fly. Primary compound chambers in which references: 1938, DIS 32: 71.
two, occasionally three, incipient cysts are en- phenotype: Abdomen deformed: in male, large and
closed occur in about 10 percent of the cases. broad; in female, tergites abnormal and hairs dis-
Male sterile. Adult fat body hypertrophied; body arranged. Wings shortened in both sexes. Bristles
size reduced. Occasionally, metathcracic legs thin and body color rather pale. Eclosion slightly
with tibiae more curved than normal, and tarsi delayed. Male viability and fertility normal; female
crooked. Viability low. RK3. viability 50 percent wild type. RK3.
MUTATIONS 165
other information: One allele each induced by CB. phenotype: Body color black. Homozygous lethal in
1540 and CB. 3034. male; female viability about 10 percent normal.
mis formed: see msf RK3.
misheld wings: see mwi morula: see mr
misproportioned: see mis *moh28: mottled
missing: see msg location: 3-46.0.
mk: murky origin: Found among progeny of males given super-
location: 1-0.8. sonic treatment.
origin: Induced by triethylenenielamine (CB. 1246). discoverer: Hersh, 28il9.
discoverer: Fahmy, 1950. references: Hersh, Karrer, and Loomis, 1930, Am.
references: 1958, DIS 32: 71-72. Naturalist 64: 552-59.
phenotype: Small fly with dull red eyes and extra Hersh, 1934, DIS 1: 30.
body pigmentation; trident pattern especially Surrarrer, 1935, Genetics 20: 357—62 (fig.)-
marked. Delayed eclosion. Male fertile but via- 1938, Genetics 23: 631-46 (fig.).
bility 50 percent wild type; female sterile. RK3. 1940, DIS 13: 51.
other information: One allele each induced by CB. phenotype: Eyes mottled with patches of dark brown
1414, CB. 1506, CB. 1540, CB. 3007, and CB. or black on wild-type background. Sensitive to
3034; two alleles induced by CB. 3025. temperature. Always mottled at 18°; almost never
*ml: minutelike above 25°C. Temperature-effective period is 25—
location: 3-46. 35 hr after beginning of pupation. Mottling more
discoverer: Mohr, 24c3. easily seen in presence of v; also manifested in w
synonym: sb: short-bristle. homozygotes (Schultz). RK1 at 18°C, RK3 above
references: 1924, Brit- J. Exptl. Biol. 2: 189-98 25°.
(fig-). *mot-32l
phenotype: Bristles small, as in Minute. Late location: 1- (not located).
hatching and poorly fertile. RK3. origin: X ray induced.
*ml2 discoverer: Oliver, 32128.
discoverer: Nichols-Skoog, 36c. phenotype: Eye color mottled in female only. RK3.
phenotype: Like ml. RK3. *mot-36e
other information: Allelism inferred from phenotype location: 3- [left arm, with In(3L)p].
and location on third chromosome. discoverer: Bridges, 36ell.
mti: see references: 1937, DIS 7: 12.
phenotype: Eyes mottled with translucent spots and
mo; m / c r o - o c u / u s roughness. Bristles twisted and stubby; hairs ir-
location: 1-6.7. regular. Wing venation plexoid around posterior
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- crossvein. Female sterile. Enhances somatic
phenylalanine (CB. 3007). crossing over in first, second, and third chromo-
discoverer: Fahmy, 1954. somes. RK3.
references: 1958, DIS 3 2 : 72. Mot-K: Mottled of Krivshenko
phenotype: E y e s small. Wings narrow and frequently location: 2- or 3- (rearrangement).
pleated longitudinally, with irregular hairs, giving origin: X ray induced,
slight opacity. Body s i z e slightly reduced. Not discoverer: Krivshenko, 54c25.
easily c l a s s i f i e d . Viability and fertility good in synonym: MoK.
both s e x e s . RK3. references; 1954, DIS 28: 75.
other information: Two a l l e l e s each induced by C B . 1955, DIS 29: 76.
3007 and C B . 3026; four induced by C B . 1528; one phenotype: Eyes liberally mottled with dark color on
each induced by CB. 1506, CB. 1540, C B . 1592, wild-type background; character barely noticeable
and CB. 3025. in young flies but striking in older ones; number
mti: see moo and size of spots variable. Homozygous lethal.
Mti: see Me Viability and fertility of heterozygotes good. RK2A.
Mo«: see Mot-K cytology: Associated with T(2;3)Mot-K =
modifier ol Bat: see su(B) T(2;3)41;60D;80~81.
modifier of garnet: see e(g) mo tiler of white: see mw
Moire: see Me mp: microptera
location: 3-0.0.
*moo: moorish discoverer: Serebrovsky, 40g8.
origin: X ray induced, phenotype: Wings small and spoonlike; veins irreg-
discoverer: Thompson. ular. Tarsi four jointed (rarely 3 or 5); joints 3 and
synonym: too (preoccupied). 4 usually fused. Antennae shortened. Ecloses
references: 1959, DIS 33: 99. somewhat late. Viability and fertility low. RK2.

discoverer: Meyer, 48c.
references: Meyer, Edmondson, Byers, and
Erickson, 1950, DIS 24: 60.
phenotype: Male sterile; female fertile. Sperm
present but not motile. RK3.
ms(2)2
location: 2-44.0 (Meyer).
discoverer: Muller, 1951.
synonym: ms.
references: Meyer, 1959, DIS 33: 97.
phenotype: Homozygous male completely sterile;
female fairly fertile. RK3.
*ms(2)E3: male sterite(2) of Edmondson
location: 2-28.
discoverer: Edmondson, 1951.
synonym: ms2,3.
phenotype: Male sterile. Fertile in heterozygotes
with fs(2)E2 (2-22.0). RK3.
*ms(2)E4
mr: mow I a location: 2-47.9.
From Bridges and Brehme, 1944, Carnegie Inst. Wash. origin: Ultraviolet induced.
Publ, No. 552: 132. discoverer: Edmondson, 1951.
synonym: ms2.4.
mr: morula references: 1952, DIS 26: 61.
location: 2-106.7. phenotype: Male sterile. Fertile in heterozygotes
discoverer: Bridges, 13c8. with fs(2)E3 (2-47.5), ts(2)E4 (2-48.5), fs(2)E5
references: Bridges and Morgan, 1919, Carnegie (2-50.4), is(2)E6 (2-54.4), ms(2)E5 (2-54.8),
Inst. Wash. Publ. No. 278: 230 (fig.). ms(2)E6 (2-54.8), ms(2)B7 (2-54.8), ts(2)E7
Bridges, 1937, Cytologia (Tokyo), Fujii Tub. Vol. (2-55.2), and ms(2)E8 (2-55.6). RK3.
2: 745-55. *ms(2)E5
phenotype: Eyes rough. Bristles irregularly reduced location: 2.54.8.
in size and number. Abdominal sclerites often origin: Ultraviolet induced.
smaller. Developmental study by Lees and discoverer: Edmondson, 1951.
Waddington [l942, Proc. Roy. Soc. (London), Ser. synonym: ms2.5.
B 131: 87-100] shows that effect on bristles re- references: 1952, DIS 26: 61.
sults from general slowing of bristle growth. Fe- phenotype: Male sterile. Fertile in heterozygotes
male entirely sterile, with underdeveloped ovaries. with fs(2)E3 (2-47.5), ms(2)E4 (2-47.9), fs(2)E4
At 19°C, bristles nearly normal and eyes nearly (2-48.5), fs(2)E5 (2-50.4), fs(2)E6 (2-54.4),
wild type. RK2 at 25°C and above. ms(2)E6 (2-54.8), ms(2)E7 (2-54.8), fs(2)E7
cytology: In salivary chromosome region between (2-55.2), ms(2)E8 (2-55.6), and ms(2)E9 (2-57.0).
59E2 and 60B10 based on its being to the right of RK3.
ln(2R)bwyD«l - In(2R)41B2-Cl;59E2'4 and to the *ms(2)E6
left of Df(2R)Px = Df(2R)6QB8-10;6QDl-2 (Bridges, location: 2-54.8.
1937). origin: Ultraviolet induced.
mr2 discoverer: Edmondson, 1951.
origin: Spontaneous. synonym: ms2.6<
discoverer: Bridges, 25k24. references: 1952, DIS 26: 61.
phenotype: Less extreme than mr. Nearly wild type phenotype: Male sterile. Fertile in heterozygotes
at 19°C. Female entirely sterile. Oogenesis with fs(2)E3 (2-47.5), ms(2)E4 (2-47.9), is(2)E4
normal through stage 4; then compound nurse cell (2-48.5), fs(2)E5 (2-50.4), (s(2)E6 (2-54.4),
chromosomes fall apart and degenerate. Karyosome ms(2)E5 (2-54.8), m&(2^;7 (2-54.8), fs(2)E7
of oocyte also disappears. Oogenesis does not (2-55.2), ms(2)E8 (2-55.6), and ms(2)E9 (2-57.0).
proceed beyond sixth stage (King, 1964, Royal RK3.
Entomol. Soc. London Symposium 2, Insect Repro- *ms(2)E7
duction pp. 13-25). RK2 at 25°C or above. location: 2-54.8.
m&: see msc origin: Ultraviolet induced.
ms(2}h male stcri!e(2) discoverer. Edmondson, 1951.
location: 2-65.5. synonym: ms2.7.
MUTATIONS 167
references: 1952, DIS 26: 61. phenotype: Male sterile. Fertile in heterozygotes
phenotype: Male sterile. Fertile in heterozygotes with fs(2)E6 (2-54.5), ms(2)E5 (2-54.8), ms(2)E6
with fs(2)E3 (2-47.5), ms(2)E4 (2-47.9), fs(2)E4 (2-54.8), ms(2)E7 (2-54.8), fs(2)E7 (2-55.2),
(2-48.5), fs(2)E5 (2-50.4), fs(2)E6 (2-54.4), ms(2)E8 (2-55.6), ms(2)E9 (2-57.0), fs(2)E8
ms(2)E5 (2-54.8), ms(2)E6 (2-54.8), fs(2)E7 (2-62.6), ms(2)E10 (2-66.5), and ms(2)Ell (2-68.0).
(2-55.2), ms(2)E8 (2-55.6), and ms(2)E9 (2-57.0). RK3.
RK3. ms(Y)Ll: male sferile in long arm of Y
*ms(2)E8 location: F.
location: 2-55.6. origin: X ray induced in y+Y.
origin: Ultraviolet induced. discoverer: Brosseau.
discoverer: Edmondson, 1951. references: 1960, Genetics 45: 257—74.
synonym: ms2.8. phenotype: Male sterile. RK3.
references: 1952, DIS 26: 61. other information: Affects complementation groups
phenotype: Male sterile. Fertile in heterozygotes kl-2, kl-3, kl-4, and kl-5. One of four such induced
with fs(2)E3 (2-47.5), ms(2)E4 (2-47.9), fs(2)E4 changes in KL among 35.
(2-48.5), fs(2)E5 (2-50.4), fs(2)E6 (2-54.4), ms(Y)L3
ms(2)E5 (2-54.8), ms(2)E6 (2-54.8), ms(2)E7 location: F.
(2-54.8), fs(2)E7 (2-55.2), ms(2)E9 (2-57.0), and origin: X ray induced in y+Y.
fs(2)E8 (2-62.6). RK3. discoverer: Brosseau.
*ms(2)E9 references: 1960, Genetics 45: 257—74.
location: 2-57. 1960, DIS 34: 48.
origin: Ultraviolet induced. phenotype: Male nearly sterile. RK3.
discoverer: Edmondson, 1951. other information: Affects complementation group
synonym: ms2.9. kl-5. Three such changes of KL among 35 tested.
references: 1952, DIS 26: 61. ms(Y)L4
phenotype: Male sterile. Fertile in heterozygotes location: F.
with fs(2)E3 (2-47.5), ms(2)E4 (2-47.9), fs(2)E4 origin: X ray induced in y + F.
(2-48.5), fs(2)E5 (2-50.4), fs(2)E6 (2-54.4), discoverer: Brosseau.
ms(2)E5 (2-54.8), ms(2)E6 (2-54.8), ms(2)E7 references: 1960, Genetics 45: 257—74.
(2-54.8), fs(2)E7 (2-55.2), ms(2)E8 (2-55.6), phenotype: Male sterile. RK3.
fs(2)E8 (2-62.6), ms(2)E10 (2-66.5), ms(2)Ell other information: One of three noncomplementing
(2-68.0), and ms(2)El2 (2-68.2). RK3. changes among 35 induced changes of KL.
*ms(2)E10 ms(Y)L7
location: 2-66.5. location: F.
origin: Ultraviolet induced. origin: X ray induced in y + F.
discoverer: Edmondson, 1951. discoverer: Brosseau.
synonym: ms2.10. references: 1960, Genetics 45: 257—74.
references: 1952, DIS 26: 61. phenotype: Male sterile. RK3.
phenotype: Male sterile. Fertile in heterozygotes other information: Affects complementation groups
with fs(2)E6 (2-54.4), ms(2)E5 (2-54.8), ms(2)E6 kl-3, kl-4, and kl-5. Nine such changes among 35
(2-54.8), ms(2)E7 (2-54.8), ia(2)E7 (2-55.2), induced in KL.
ms(2)E8 (2-55.6), ms(2)E9 (2-57.0), fs(2)E8 ms(Y)L10
(2-62.6), ms(2)Ell (2-68.0), and ms(2)E12 (2-68.2). location: F.
RK3. origin: X ray induced in y+Y.
*ms(2)EU discoverer: Brosseau.
location: 2-68. references: 1960, Genetics 45: 257—74.
origin: Ultraviolet induced. phenotype: Male sterile. RK3.
discoverer: Edmondson, 1951. other information: Affects complementation groups
synonym: ms2.ll. kl-1 and kl-3. The only change induced in KL af-
references: 1952, DIS 26: 61. fecting nonadjacent complementation groups.
phenotype: Male sterile. Fertile in heterozygotes ms(Y)Ul
with fa(2)E6 (2-54.4), ms(2)E5 (2-54.8), ms(2)E6 location: F.
(2-54.8), ms(2)E7 (2-54.8), ia(2)E7 (2-55.2), origin: X ray induced in y*Y.
ms(2)Es (2-55.6), ms(2)E9 (2-57.0), fs(2)E8 discoverer. Brosseau.
(2-62.6), ma(2)E10 (2-66.5), and met(2)E12 (2-68.2). references: 1960, Genetics 45: 257-74.
RK3. 1960, DIS 34: 48.
*ms(2)E12 phenotype: Male sterile. RK3.
location: 2-68.2. other information: Affects complementation group
origin: Ultraviolet induced, kl-3. Four such changes in KL among 35.
discoverer: Edmonds on, 1951. ms(Y)U2
synonym: ms2.I2. location: F.
references: 1952, DIS 26: 61. origin: X ray induced in y+Y.
discoverer: Brosseau. ms(Y)S5

references: 1960, Genetics 45: 257—74. location: F.
phenotype: Male sterile. RK3. origin: X ray induced in y+Y.
other information: Affects complementation groups discoverer Brosseau.
kl-2 and kl-3. Two such changes in KL among 35. references: 1960, Genetics 45: 257—74.
ms(Y)L13 1960, DIS 34: 48.
location: F. phenotype: Male sterile. RK3.
origin: X ray induced in y+Y. other information: Affects complementation group
discoverer Brosseau. ks-2. Unique among 11 induced changes of KS.
references: 1960, Genetics 45: 257—74. *ms(Y)S14
1960, DIS 34: 48. location: F.
phenotype: Male sterile. RK3. origin: X ray induced in y+Y.
other information: Affects complementation group discoverer: Brosseau.
kl-1. Five of 35 KL changes were like ms(Y)L13. references: 1960, Genetics 45: 257-74.
ms(Y)L32 phenotype: Male sterile, bb deficient. RK3.
location: Y. other information: The only noneomplementing KS
origin: X ray induced in y+Y. change found among 11.
discoverer: Brosseau. ms2.: see ms(2)E
references: 1960, Genetics 45: 257—74. *msc: melanoscutellum
phenotype: Male sterile. RK3. location: 1-52.6.
other information: Affects complementation groups origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
kl-1, kl-2, and kl-3. Unique among 35 induced KL phenylalanine (CB. 3007).
changes. discoverer: Fahmy, 1954.
ms(Y)L36 synonym: ms (preoccupied).
location: Y. references: 1958, DIS 32: 72.
origin: X ray induced in y+Y. phenotype: Extra pigmentation confined to scu-
discoverer: Brosseau. tellum. One or more thoracic bristles duplicated.
references: 1960, Genetics 45: 257—74. Eyes slightly more oval than normal. Wings
1960, DIS 34: 48. slightly abnormal in shape and position. Charac-
phenotype: Male nearly sterile. RK3. ters not always penetrant. Viability and fertility
other information: Affects complementation groups good in both sexes. RK3.
kl-4 and kl-5. Unique among 35 induced changes of other information: One allele each induced by CB.
KL. 1506 and CB. 3025; two induced by CB. 3007.
ms(Y)L37 Msc: Multiple sex comb
location: Y. location: 3-48.0.
origin: X ray induced in y+Y. origin: Spontaneous.
discoverer: Brosseau. discoverer: Tokunaga, 64a.
1960, DIS 34: 48. phenotype: Extra sex combs on second and third
phenotype: Male sterile. RK3. legs of male. Fewer teeth on sex comb of first leg.
other information: Affects complementation group Homozygote lethal; heterozygote with Pc Sex sur-
kl-2. Unique among 35 KL changes. vives. RK1A.
ms(Y)L38 cytology: Associated with In(3R)Msc =
location; F. In(3R)84B;84F.
origin: X ray induced in y+Y. msh mi stormed
discoverer Brosseau. location: 2-55.2 (originally located at 55.6 but arbi-
references: 1960, Genetics 45: 257-74. trarily placed at 55.2 to be consistent with cyto-
1960, DIS 34: 48. logical indication that it is to the left of pk).
phenotype: Male sterile. RK3. discoverer: Bridges, 30b8.
other information: Affects complementation groups references: Curry, 1939, DIS 12: 46.
kl-3 and kl-4. Unique among 35 induced changes of phenotype: Eyes misshapen. Wings short and
KL. crumpled; legs shortened. Characteristics variable
ms(Y)S2: male sterile in short arm of Y and overlap wild type. RK3.
location: F. cytology: Placed between 41A and 42A3 on basis of
origin: X ray induced in y+Y. its inclusion in Df(2R)bwVE>°2LQyR = Di(2R)41A-
discoverer: Brosseau. B.-42A2-3 (Schultz). Certainly included in Dt(2RyU-
references: 1960, Genetics 45: 257—74. S2**11 z*Di(2R)40F-41Al;42A19'Bl (Morgan,
1960, DIS 34: 48. Schultz, Bridges, and Curry, 1939, Carnegie Inst.
phenotype: Male sterile. RK3. Wash. Year Book 38: 275).
other information: Affects complementation group *msg: missing
k*-I. One of nine such changes among 11 induced location: 2- (not located),
KS alterations. origin: Spontaneous.
MUTATIONS 169
discoverer: Mossige, 50b4. mw: mottler of white

references: 1951, DIS 25: 69. location: 1- (slightly to the right of ct).
phenotype: Bristles greatly reduced or missing. In origin: Spontaneous.
extreme cases, almost like sv. Female sterile; discoverer: Muller, 1946.
male sparingly fertile. RK2. references: 1946, DIS 20: 88-89.
tntA: see tu-bw phenotype: Normal by itself. A specific dilutor of
*mtb: matt brown wa and other intermediate alleles at the w locus.
location: 1-3.6. Eyes assume a lighter mottled appearance. Ex-
origin: Induced by e'thyl methanesulfonate (CB. pression not affected by dosage of Y chromosome
1528). (Oster, 1957, DIS 31: 150). RK1.
discoverer: Fahmy, 1956. cytology: Not associated with chromosome aberra-
references: 1959, DIS 33: 88. tion (Oster, 1957).
phenotype: Eye color flat and browner than normal
with greatly reduced reflection spots. Wing posi-
tion varies from slightly to completely outspread;
sometimes upheld. Male sterile, and viability
about 30 percent wild type. RK2.
*mu: mussed
location: 3-50.
discoverer: Mohr, 37121.
references: Mossige, 1939, DIS 12: 47.
phenotype: Wings thin textured. Dorsal surface of
thorax arched. RK1.
*mu-F: mutability factor from Florida mwh: multiple wing hairs
location: 2- (not located). Wing hairs. Left: wild type. Right: mwh.
origin: Spontaneous in Florida wild stock. A. Di Pasquale, unpublished.
references: 1937, Genetics 22: 469—78. mwh: multiple wing hairs
phenotype: Homozygote shows increase in lethal location: 3-0.0 (order with m and ve not tested).
and visible mutation rate. Factor acts during de- origin: Spontaneous.
velopment of germ cells in both male and female. discoverer: Di Pasquale, 501.
*mul: multiple 1952, Rend. 1st. Lombardo Sci. Lettere, Ser. B 85:
location: 1-0.0. 1-8.
origin: Spontaneous. phenotype: Wing cells contain groups of 2—5 hairs
discoverer: Neel, 41cl3. instead of one hair per cell as in wild type. Trans-
references: 1942, DIS 16: 51. plants of mutant wing disks to wild-type hosts de-
phenotype: Eyes rough and oval. Wings weak and velop autonomously (Ursprung and Hadom, 1962,
held out. Bristles occasionally missing or disar- Develop. Biol. 4: 40-60). RK1.
ranged. Body may show abnormal protuberances cytology: Salivary chromosomes apparently normal.
covered with hairs. Female sterile. After a few *mwhseml: multiple wing hairs-semi
generations in stock, only the eye abnormality origin: Spontaneous derivative of mwh.
showed. RK2. discoverer: Di Pasquale, 51e.
multiple wing hairs: see mwh phenotype: Like mwh except that the groups of wing
Multiple sex comb: see Msc hairs are restricted to wing margins. Wing surface
*mur: murrey between second and fifth longitudinal veins has
location: 1-14.3. single hair with only an occasional group.
origin: Spontaneous as one mosaic male. mwh*
mml
/mwh is likemwfisemi/mH'/i«e'n'. RK1.
discoverer: E. H. Grell, 57c. *mwi: misheld wings
phenotype: At 25°C, eye color reddish purple, origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
bristles very small, and body size reduced. At phenylalanine (CB. 3007).
17°C, eye color and body size normal, but bristles discoverer: Fahmy, 1954.
rather small. Original mosaic male transmitted references: 1958, DIS 32: 72.
only an X containing mur. He was mated to his phenotype: Wings diverge upward and outward at
daughters to produce homorygous mur females. various angles. Eye shape oval. Viability and fer-
mur/mur female and mut male are sterile. RK3. tility good in male but reduced in female. RK2.
murky: see mk
murrey, see mur nfah; see ml
mussed: see mu N: Notch
mutability factor from Florida: see mu~F location: 1-3.0.
phenotype: Wings of heterozygote incised at tips

and often along edges. Veins L3 and L5 thickened.
Thoracic hairs irregularly distributed. Male and
homozygous female lethal. Heterozygotes for any
two N alleles lethal. fa/N, spI/N, and nd/N hetero-
zygotes express both the N phenotype and the
phenotype of the recessive. N/fan° is lethal. For
developmental studies of N male embryo, see
Poulson, 1939, DIS 12: 64-65; 1940, J. Exptl.
Zool. 83: 271-325; Poulson and Boell, 1946,
Anat. Record 96: 508; Counce, 1961, Ann. Rev.
Entomol. 6: 295-312. RK1.
cytology: Heterozygosity for a deficiency including
salivary chromosome band 3C7 produces the N
phenotype (Demerec, Kaufmann, Fano, Sutton, and
41: 191). Many but not all N alleles are associ-
ated with chromosome rearrangements.
other information: The TV alleles and the recessives
ia, fano, spl, and nd belong to a pseudoallelic
complex (Welshons and Von Halle, 1962, Genetics
47: 743-59).
N8; Notch-8
From Mohr, 1924, Z. Induktive Abstammungs-
0.03 0.05 0.03 0.03 Vererbungslehre 32: 118.
fa fa spl nd
I I I
55ell 264-40 Nic 24\ 60g11
Co origin: Spontaneous.
264-103 discoverer: Mohr, 28k22.
phenotype: Like N. RK1A.
Map of the N region cytology: Associated with Df(l)N25. breakpoints not
Showing relative positions of dominant (below the line) known (Sutton).
and recessive (above the line) mutants at the Notch
locus. origin: Spontaneous.
From Welshons and Von Halle, 1962, Genetics 47: discoverer: Mohr, 28k29.
743-59. phenotype: Like N. RK1A.
cytology: Associated with Df(l)N26 = Df(l)3C4-
5;3C8-9 (Sutton).
discoverer: Mohr, 18j7. origin: Spontaneous.
references: 1919, Genetics 4: 275-82. discoverer: Mohr, 30115.
1923, Z. Induktive Abstammungs- Vererbungslehre phenotype: Like/V. RK1.
32: 108-232 (fig.). other Information: N27/w not white.
1932, Proc. Intern. Congr. Genet., 6th. Vol. 1J *N29
Mackensen, 1935, J. Heredity 26: 163-74. discoverer: Eker, 36el2.
Gottschewski, 1937, Z. Induktive Abstammungs- phenotype: Like N. RK1(A).
Vererbutigslehre 73: 131—42. cytology: Association with Df(l)N29 inferred from
Slizynska, 1938, Genetics 23: 291-99. its mutant interaction with w.
phenotype: Typical Notch. Hemizygous male lethal
in egg (Li, 1927, Genetics 12: 1-58). Nervous origin: Spontaneous.
system hypertrophied; ventral and cephalic hypo- discoverer: Mohr, 38b21.
derm missing; mesodermal organs absent; fore-gut phenotype: Like N. RK1.
rudimentary, and mid-gut incomplete [Poulson, other information: N3G/w not white.
1940, J. Exptl. Zool. 83: 271-325 (fig.); 1941,
Proc. Intern. Congr. Genet., 7th. pp. 240—4l]. origin: Spontaneous.
Cholinesterase activity higher in Ns than in normal discoverer: Ives, 33h29.
embryos (Poulson and Boell, 1946, Biol. Bull. 91: synonym: N*64'!*.
228; 1946, Anat. Record 96: 508). RK1A. references: Plough and Ives, 1934, DIS 1: 31.
cytology: Associated with Df(I)Ns 1934, DIS 2: 10, 34.
phenotype: Like N. RK1A.
MUTATIONS 171
cytology: Associated with Df(l)N33h = Df(l)3C6- cytology: Salivary chromosomes normal (Welshons).
7;3D2-3 (Sutton). other information: Located to the left of fa
(Welshons, Von Halle, and Scandlyn, 1963, Proc.
origin: X ray induced. Intern. Congr. Genet., 11th. Vol. 1: 1—2). Does
discoverer: Oliver, 34b3. not show mutant interaction with w, rst, dm, or ec.
phenotype: Like N. RK1(A). origin: Gamma ray induced.
cytology: Association with T(l;3)N34b suspected. discoverer: Ives.
Basis of suspicion not mentioned. phenotype: Like N. RK1.
other information: Recombines with fa and spl. Lo-
origin: Spontaneous. cated to the right of spl (Welshons and Von Halle,
discoverer: Curry, 38g. 1962, Genetics 47: 743-59). N60U0/w not white.
phenotype: Like /V. RK1A. H60gll
cytology: Associated with Df(l)N38g = Df(l)3C4- origin: Gamma ray induced.
5.-3C7-8 (Sutton). discoverer: Ives.
*N40j phenotype: Wings seldom notched; veins thickened;
origin: Spontaneous. deltas at tips. N60^1 */+ has rough eyes resem-
discoverer: Sismanidis, 40j. bling spl. N60611 /spl has extremely rough eyes.
references: Mather, 1942, DIS 16: 49. N60gll/[a e y e s like fam Semilethal with fa"°; poor
phenotype: Like N. RK1. viability with nd. RK2.
other information: N4®)/w not white. cytology: Salivary chromosomes normal (Welshons).
*N47i other information: Located to the right of Nc° and
origin: Ultraviolet induced. probably to the left of nd (Welshons, Von Halle,
discoverer: Meyer, 47i. and Scandlyn, 1963, Proc. Intern. Congr. Genet.,
references: 1952, DIS 26: 67. 11th. Vol. 1: 1-2).
phenotype: Expression less extreme than N; about N60h21
70 percent of heterozygotes wild type. N47i/spl origin: Gamma ray induced.
has wild-type eye, but bristles are like spl. discoverer: Ives.
N47i/fa has wild-type eye. Homozygous lethal. phenotype: Like N. Lethal in combination with
RK3. ta"°. RK1.
other information: Located to the right of spl
origin: X ray induced. (Welshons and Von Halle, 1962, Genetics 47:
discoverer: Lefevre, 5 0 k l l . 743-59). N60h21/w n o t w hite.
references: 1951, DIS 25: 71. N60jJ4
1952, DIS 26: 66. origin: Gamma ray induced.
Ratty, 1954, Genetics 39: 513-28. discoverer: Ives.
phenotype: Like N. RK1A. phenotype: Like N. N6°)*4/fano lethal. RK1.
cytology: Associated with T(l;3)NS0k11 = other information: Located to the right of apt
T(1;3)1E3-4;3C6-7;3C8-9;89A. 3C7 and 8 missing. (Welshons and Von Halle, 1962, Genetics 47:
*NSld 743-59).
origin: Ultraviolet induced. N6U19
discoverer: Byers, 51d. origin: Gamma ray induced.
references: Meyer and Edmonds on, 1951, DIS 25: discoverer: Ives.
73. phenotype: Like N. N61M9/fan° lethal. RK1.
Meyer, 1952, DIS 26: 67. other information: Located to the right of spl
phenotype: Like N, but whereas N^l ^/ia has char- (Welshons, Von Halle, and Scandlyn).
acteristic fa phenotype, N5id/spl has no spl char- N61hlO
acteristics. RK1. origin: Gamma ray induced.
NS4I9 discoverer: Ives.
origin: Spontaneous. phenotype: Like N. N<"t>*°/fa"° lethal. RK1.
discoverer: Mohler, 5419. other information: Located to the right of spl
references: 1956, DIS 30: 78. (Welshons, Von Halle, and Scandlyn).
phenotype: Weak Notch. Deltas of long veins re-
liable in classification when wing tips not origin: Gamma ray induced.
notched. RK2. discoverer: Ives.
phenotype: Like N. N<>2til 0/fano lethal. RKl.
origin: Spontaneous. other information: Located to the right of spl
discoverer: Mohler, 5 5 e l l . (Welshons, Von Halle, and Scandlyn).
references: 1956, DIS 30: 78. N62I
phenotype: Weak Notch. Deltas on wing veins most origin: Found among progeny of male treated with
reliable character for classification. Lethal when radiofrequency waves.
heterozygous with /a n o , N^°Stl, and Nc°. RK2. discoverer: Mickey, 6213.
references: 1963, DIS 38: 29. N264-10

phenotype: Like N. RK1. origin: X ray induced.
origin: X ray induced. phenotype: Heterozygous female like N/+. X/Y/Y
references: Lefevre and Wilkins, 1966, Genetics 53: male viable but sterile; has slight rst variegation.
175-87. X/Y male lethal (Schultz). RK1A.
phenotype: Typical Notch; inseparable from w63b. cytology: Associated with T(l;2yN264-10; break-
cytology: Associated with Df(l)N63b = Df(l)3C2- points not known.
3;3E2-3. N264.12
*N218 origin: X ray induced.
origin: X ray induced in R(l)2. discoverer: Demerec, 34a.
discoverer: Barigozzi. synonym: NaS.
references: 1939—40, Rend. 1st. Lombardo Sci. references: 1941, Proc. Intern. Congr. Genet., 7th.
Lettere, A 73: 382-87. pp. 99-103.
1940, DIS 13: 69. Judd, 1955, DIS 29: 126-27.
1942, Rev. Biol. (Perugia) 34: 59-72. phenotype: Expression weak but not variegated.
phenotype: Notching variable, N2i8/fa and N218/spl RK2A.
show variable expression for fa and spl, respec- cytology: Associated with T(1;4)N264.12 =
tively. Few X/Y males survive below 23°C and T(l;4)3C6-7;101F (Sutton).
are sterile. RK2A.
cytology: Associated with In(l)N2l8 _ ln(l)3C;20. origin: X ray induced.
Since inversion was induced in ring, position of discoverer: Demerec, 34a.
centromere uncertain. phenotype: Like N. RK1A.
N264.2 cytology: Associated with Df(l)N264-l3 - Df(l)3C6-
origin: X ray induced. 7;3C10-ll (Demerec and Hoover).
discoverer: Demerec, 33j. *N264.1S
references: Slizynska, 1938, Genetics 23: 291—99. origin: X ray induced.
phenotype: Like N. RK1A. discoverer: Demerec, 34c.
cytology: Associated with Df(l)N264-2 _ D%1)3C6- phenotype: Like N. RK1A.
7;3C7-8, cytology: Associated with Di(l)N264-l S = Df(l)3C6-
N264.6 7;3C7-8 (Sutton).
origin: X ray induced. N264-18; S e e N33h
discoverer: Demerec, 33k20.
phenotype: Heterozygous female like N/+; N264~6/Y N264.19
male usually lethal; N364~6/Y/Y male usually origin: X ray induced.
viable but sterile (Schultz). RK1A. discoverer: Demerec, 34k.
cytology: Associated with T(1;3)N264-* = references: Slizynska, 1938, Genetics 23: 291—99.
T(1;3)3C9-D1;62A; 73B;80C. phenotype: Like N. Embryonic development of
iy264~19 m a i e similar to N 8 (Poulson, 1941, Proc.
origin: X ray induced, Intern. Congr. Genet., 7th. pp. 240—41). RK1A.
discoverer: Demerec, 33k. cytology: Associated with D^1)N264-19 _ Df(l)3C6-
phenotype: Like AT. Lethal and cell lethal. RK1A.
cytology: Associated with In(l)N26*-7 = In(l)3C6' *N264.20
7;3C8-9;8C5-7. 3C7 and 8 missing (Hoover). origin: X ray induced.
discoverer: Demerec, 34g.
origin: X ray induced. phenotype: LikeJV. RK1A.
discoverer: Demerec, 33k. cytology: Associated with T(1;4)N264-20 =
references: Slizynska, 1938, Genetics 23: 291-99. T(l;4)3C4-5;3C7-8;101F; deficient for 3C5-7
phenotype: Like N, Developmental abnormalities of (Sutton).
male same as Na (Poulson, 1939, DIS 12: 64-65). *N2 64-23
cytology: Salivary chromosomes apparently normal discoverer: Demerec, 35h.
(Slizynska). references: 1941, Proc. Intern. Congr. Genet., 7th.
other information: w, rst, and ec not affected. pp. 99-103.
*N264-9 phenotype: Like N, but overlaps wild type. Occa-
origin: X ray induced. sional male with normal phenotype survives.
discoverer: Demerec, 3315. RK2A.
phenotype: Variegated for/V. X/Y male lethal. cytology: Associated with T(1;2)N264-23 =
X/Y/Y male viable and almost normal in appear- T(1;2)3C8-9;41A (Demerec and Hoover).
ance but sterile (Schultz). RK2A. *N2 64.24
cytology: Associated with T(l;2)N264-» » origin: X ray induced.
T(1;2)3C;41. discoverer: Demerec, 35h.
MUTATIONS 173
references: 1941, Proc. Intern. Congr. Genet., 7th. *N2 64.37

pp. 99-103. origin: X ray induced.
phenotype: Notch not expressed in all heterozygous discoverer: Demerec, 37b.
females. Males not observed. RK2A. references: Slizynska, 1938, Genetics 23: 291-99.
cytology: Associated with T(1;2)N264-24 = phenotype: Like N. RKlA.
T(l;2)3C8-9;40F (Demerec). cytology: Associated with DI(1)N264-37 = Df(i)3C6-
*N264.29 7;3C7-8.
origin: X ray induced. N264.38
discoverer: Demerec, 36d. origin: X ray induced.
references: 1941, Proc. Intern. Congr. Genet., 7th. discoverer: Demerec, 37b.
pp. 99-103. references: Slizynska, 1938, Genetices 23: 291—99.
phenotype: Notch not expressed in all heterozygous phenotype: Like N. Male embryo shows same abnor-
females. A few males with normal phenotype sur- malities as N8 (Poulson, 1941, Proc. Intern.
vive. RK2A. Congr. Genet., 7th. pp. 240-41). RKlA.
cytology: Associated with T(1;3)N264-29 = cytology: Associated with Df(l)N264-3S =Df(l)2D3-
T(l;3)3D4-5;80 (Hoover). 4;3E2-3.
N264.39
origin: Spontaneous in X carrying wch.
origin: X ray induced, discoverer: Slizynska, 1937.
discoverer: Demerec, 36d. references: 1938, Genetics 23: 291—99.
references: Slizynska, 1938, Genetics 23: 291—99. phenotype: Like N. N264~39/fano lethal. RK1(A).
phenotype: Like N. RKlA. cytology: Associated with Df(l)N264~39 = Df(l)3C6'
cytology: Associated with Df(l)N264-30 - Df(l)3A4- 7;3C7-8 (Slizynska, 1938; Welshons, 1958, Proc.
5;3C7-9. Natl. Acad. Sci. U.S. 44: 254-58). Later reexami-
nation of chromosomes of males from lines carrying
origin: X ray induced, wch and marked N264'39 revealed the presence of
discoverer: Demerec, 36d. 3C7 (Welshons).
references: Slizynska, 1938, Genetics 23: 291—99. other information: Recombines with both fa and spl
phenotype: Like /V. RKlA. and lies between them (Welshons, Von Halle, and
cytology: Associated with Df(iyN264"3* = Df(l)3B4- Scandlyn, 1963, Proc. Intern. Congr. Genet., 11th.
Cl;3D2-3. Vol. 1: 1-2).
N264-40
discoverer: Demerec, 36h. discoverer: Demerec, 37d.
references: Slizynska, 1938, Genetics 23: 291—99. phenotype: Like N. Male embryos have abnormali-
phenotype: Like N. RKlA. ties like Na (Poulson, 1939, DIS 12: 64-65).
cytology: Associated with Dt(l)N264'32 =Df(l)3C3- Lethal with /a"°. RK1.
5.-3C7-8. cytology: Salivary chromosomes apparently normal
*N264.33 (Hoover).
origin: X ray induced. other information: Located between fano and NNlc
discoverer: Hoover, 36h. (Welshons and Von Halle, 1962, Genetics 47: 743-
references: Slizynska, 1938, Genetics 23: 291—99. 59). w, rat, and dm not affected.
phenotype: Like N. RKlA. *N264.41
cytology: Associated with Df(l)N264-33 = Df(l)3C6- origin: Spontaneous in chromosome containing w.
7;3C7-8. discoverer: Slizynska, 37e.
phenotype: Like iy. RKlA.
origin: X ray induced. cytology: Associated with DfCljN264-4* =D%1)3C6-
discoverer Demerec, 37a. 7;3C8-9 (Sutton).
phenotype: Like N. Male shows same abnormalities discoverer. Demerec, 37e.
in embryonic development as Ns (Poulson, 1939, phenotype: Like N. RKlA.
DIS 12: 64-65). RKlA. cytology: Associated with Dt(l)N264-42 = Dt(l)3C4'
cytology: Associated with T(1;3W264~34 » 5;4B4~6 (Hoover).
7Xl;3)3C3-5;70e2-3 (Hoover). *N264.46
*N264.36 origin: X ray induced.
origin: X ray induced, discoverer: Demerec, 37f.
discoverer: Demerec, 37b. phenotype: Like N. RKlA.
references: Slizynska, 1938, Genetics 23: 291—99. cytology: Associated with D^lfi364'4^
phenotype; Like N. RKlA. 7;3C7-8.
cytology: Associated with DXiyt™4'36 » Df(l)3A3- H264.47
4;3D2-3. origin: X ray induced.
174 GENETIC VARIATIONS OF DROSOPHILA MELANOGA5TER
discoverer: Demerec, 37f.

phenotype: Like N. Lethal with iano. Male embryos origin: X ray induced.
show same developmental abnormalities as N& discoverer: Demerec, 38b.
(Poulson, 1939, DIS 12: 64-65). RK1. phenotype: Like N. RK1A.
cytology: Salivary chromosomes apparently normal cytology: Associated with Df(l)N264'54 = Df(l)3C3-
(Sutton). 5;3C7-8 (Hoover).
other informotion: Probably located to the right of */\/264-55
spl, but wild-type progeny from N264"47 /spl fe- origin: X ray induced.
males are frequently nonrecombinant (Welshons, discoverer: Demerec, 38b.
1958, Proc. Natl. Acad. Sci. U.S. 44: 254-58). references: 1941, Proc. Intern. Congr. Genet., 7th.
w, rst, and dm not affected. pp. 99-103.
+N264.48 phenotype: Variable Notch. N264~ss/fa variegates
origin: X ray induced. for fa. RK2A.
discoverer: Demerec, 37f. cytology: Associated with T(1;3)N264-55 =
references: 1941, Proc. Intern. Congr. Genet., 7th. T(l;3)3D4'5;80F9-81Fl.
pp. 99-103. */s/2 64-56
phenotype: Like N. RK1A. origin: X ray induced.
cytology: Associated with In(l)N264-4a =In(l)IB6- discoverer: Demerec, 38c.
7;lB10-ll;3C7-8. 1B7-10 missing (Hoover). phenotype: Probably variegated for N. RK2A.
cytology: Associated with T(1;3)N264-56 =
origin: X ray induced. T(l;3)3D4-5;80 (Sutton).
discoverer: Demerec, 37j. •f/264.57
phenotype: Like N but also slight Minute. RK1A. origin: X ray induced.
cytology: Associated with Df(l)N264-49 = Dt(l)3C4* discoverer: Demerec, 38d.
S;3E8'F1 (Sutton). references: 1941, Proc. Intern. Congr. Genet., 7th.
other information: Minute phenotype results from in- pp. 99-103.
clusion of M(1)3E in deficiency. phenotype: Notch variable. N264~S7/fa not facet.
-50 RK2A.
origin: X ray induced. cytology: Associated with In(l)N264-57 = In(l)3C9-
discoverer: Demerec, 37k. 11;2OD2-E1 (Hoover).
references: 1941, Proc. Intern. Congr. Genet., 7th. N264-58
phenotype: Notching of wings variable and not al- discoverer: Demerec, 38d.
ways penetrant. N264~S0/fa shows variegation for references: 1940, Genetics 25: 618-27.
fa. RK2A. phenotype: Like N. N264-58/fa variegates for fa.
cytology: Associated with T(l;2)N264-S0 = RK1A.
T(l;2)3C7-9;20Cl-F;22A2-3 (Hoover). cytology: Associated with T(1;3)N264-58 =
N2 64-51 T(l}3)3B2'3;3D6^7;80D-F (Sutton).
origin: Found among progeny of radium-treated male.
phenotype: Like N. RK1A. discoverer: Demerec, 38d.
cytology: Associated with references: 1941, Proc. Intern. Congr. Genet., 7th.
D%l)3C6-7;3C7-8 (Sutton). pp. 99-103.
phenotype: Weak Notch. N264~S9/spl variegates for
origin: X ray induced. spl. RK2A.
discoverer: Demerec, 38a. cytology: Associated with T(1;2^SI264-S9 =
references: 1941, Proc. Intern. Congr. Genet., 7th. T(l;2)3C8-9;40F (Hoover).
pp. 99-103.
phenotype: Notch in wings not always present. origin: X ray induced.
N364"S2/ia shows variegation for fa. RK2A. discoverer: Demerec, 38d.
cytology: Associated with In(I)N264'52 * In(t)3C3- phenotype: Like N. RK1.
5;2OB2-C1. cytology: Salivary chromosomes appear normal.
N2 64.53
other information: w, dm, and ec not affected.
origin: X ray induced. *N264-62
references: 1941, Proc. Intern. Congr. Genet., 7th. discoverer: Demerec, 38e.
pp. 99-103. references: 1941, Proc. Intern. Congr. Genet., 7th.
phenotype: Like N. N264'53/fa is not facet. Devel- pp. 99-103.
opmental abnormalities differ from other N alleies phenotype: Like /V. N264-62/fa variegates for fa.
(Poulson), RK1A. RK1A.
cytology: Associated with 7*7,- cytology: Associated with T(1;2)N364~62 =
T(1;2)3C6'7;34C7-D1. T(1;2)3C7~8;41A-B (Sutton).
MUTATIONS 175
*N264.63
discoverer: Demerec, 38e. discoverer: Demerec, 38k.
references: Sutton, 1940, Genetics 25: 534—40. references: 1941, Proc. Intern. Congr. Genet., 7th.
Demerec, 1941, Proc. Intern. Congr. Genet., 7th. pp. 99-103.
pp. 99-103. phenotype: Strong Notch. RK1A.
phenotype: Weak Notch. Overlaps wild type. RK2A. cytology: Associated with In(l)N264-71 =ln(l)3C6-
cytology: Associated with Tp(l)N264-63 = Tp(l)3C7- 7;20D-B (Sutton).
9;13C;19F (Hoover). HI 6 4.12
*N264.64 origin: X ray induced.
origin: X ray induced. discoverer: Demerec, 38k.
discoverer: Demerec, 38e. phenotype: Like /V. RK1A.
references: 1941, Proc. Intern. Congr. Genet., 7th. cytology: Associated with Df(iyN264-72 =Df(l)3C6-
pp. 99-103. 7;3C7-9 (Sutton).
phenotype: Overlaps wild type. N264~64/ia varie- *N264.73
gates for fa. RK2A. origin: X ray induced.
cytology: Associated with T(1;3)N264-64 = discoverer: Demerec, 381.
T(l;3)3E5-6}80C-F (Hoover). phenotype: Heterozygous females both Notch and
*N264-6S Minute. RK1A.
origin: X ray induced, cytology: Associated with Df(l)N264-73 =Df(l)3C3~
discoverer: Demerec, 38e. 4;4C6~7 (Demerec).
phenotype: Overlaps wild type. N264'65/ fa varie- other information: Minute phenotype results from in-
gates for ia. RK2A. clusion of M(1)3E and M(1)4BC in the deficiency.
cytology: Associated with T(1;3)N264-65 = *N264.74
T(l;3)2B10-16}3D4-5;8lF;96C4-5 (Hoover). origin: X ray induced.
N264.66 discoverer: Demerec, 38k.
origin: X ray induced. references: Sutton, 1940, Genetics 25: 534—40.
discoverer: Demerec, 38e. phenotype: Like N. N264"74/fa variegates for fa.
phenotype: Notching of wings weak and rarely vis- RK1A.
ible. N264-66/fa variegates for fa. Some males cytology: Associated with TXl;2;3yN264"74 =
viable; have cream-colored eyes with spots of T(1;2;3)3C10-ll;20D-E;40C-D;92E6-8 (Sutton).
normal red pigment. RK3A. *N264.76
cytology: Associated with T(1;2)N264-66 = origin: X ray induced.
T(1;2)3C6-7;41 + T(1;2)7C9-D1;53F (Hoover). discoverer: Demerec, 39b.
phenotype: Like N. Also slight Minute. RK1A.
*N264.68 cytology: Associated with Df(iyN264*76 =Df(l)3B4-
origin: X ray induced. Cl;3E4-5 (Sutton).
discoverer: Demerec, 38k. other information: Minute phenotype results from in-
phenotype: Like N but with slight Minute effect. clusion of M(1)3E in the deficiency.
RK1A.
cytology: Associated with DîyN264"68 = origin: X ray induced.
Df(l)3A10-Bl;3E8-Fl (Demerec). discoverer: Demerec, 39b.
other information: Minute phenotype results from in- phenotype: Like N. RK1A.
clusion of M(1)3E in the deficiency. cytology: Associated with Df(l)N264"77 = Df(l)3B4-
Cl;3C7-8 (Sutton).
origin: X ray induced. *N264.79
references: 1941, Proc. Intern. Congr. Genet., 7th. discoverer: Demerec, 39c.
pp. 99-103. phenotype: Like N but overlaps wild type. RK2A.
phenotype: Like JV. RK1A. cytology: Associated with Df(l)N264-79 =
cytology: Associated with T(1;2)N264~69 » D£Cl)2C10-Dl;3C6-7 (Sutton).
T(l;2)3C7-8;44C4-5 (Demerec). +N264.80
origin: X ray induced. discoverer: Demerec, 39d.
discoverer: Demerec, 33k. references: 1941, Proc. Intern. Congr. Genet., 7th.
references: Sutton, 1940, Genetics 25: 534—40. pp. 99-103.
phenotype: Wing notching overlaps wild type. phenotype: Like N. RK1A.
pj264-70/fm variegates for /a. Male viable and cytology: Associated with IXl^W264-*0 »
mottled for w and rmt. RK2A. T(l;2)3C6-7;36;40. An inversion with breakpoints
cytology: Associated with 7X1;3)N264-?O » in 11 and 20 induced at same time (Sutton).
Ttl;3)3C4-5;B0D-F + T(l;3)6F2-7Al;lWB2-3
(Sutton). origin: X ray induced.
discoverer: Demerec, 39d. cytology: Salivary chromosomes normal (Sutton).

phenotype: Like N. RK1A. other information: w, rst, and dm not affected.
cytology: Associated with Dt(l)N264"81 = Df(l)3C6-
7;3C7-8 (Sutton). origin: X ray induced.
discoverer: Demerec, 39j.
origin: X ray induced. phenotype: Like /V. Also slight Minute. RKlA.
discoverer: Demerec, 39d. cytology: Associated with Dt(l)N264-89 = Df(l)3B2-
phenotype: Like N. N264"82/fa variegates for fa. 3;3F2-3 (Sutton).
RK1A. other information: Minute phenotype results from in-
cytology: Associated with T(1;2)N264-82 = clusion of M(1)3E in deficiency.
T(1;2)3C3-4;41A + T(l;2)20A;57. Tip of 2L in *N264.90
chromocenter and may be involved (Sutton). origin: X ray induced.
43 discoverer: Demerec, 39j.
origin: X ray induced. phenotype: Like N. Also slight Minute. RKlA.
discoverer: Demerec, 39d. cytology: Associated with Df(l)N264-90 = Df(l)3C7-
references: 1941, Proc. Intern. Congr. Genet., 7th. 8;3E8-F1 (Sutton).
pp. 99-103. other information: Minute phenotype results from in-
phenotype: Like N. RK1A. clusion of M(1)3E in the deficiency.
cytology: Associated with T(1;3)N264-83 =
T(l;3)3C6-7;12F2-4;79E2-3 + In(3R)81;88. origin: X ray induced.
discoverer: Demerec, 39g.
N264.84 phenotype: Like N. RK1.
origin: X ray induced. cytology: Salivary chromosomes normal (Sutton).
discoverer Demerec, 39c. other information: w, rst, and dm not affected.
references: 1941, Proc. Intern. Congr. Genet., 7th. *N264.93
phenotype: Like N. N264'84/fa variegates for fa. discoverer: Demerec, 39k.
RK1A. phenotype: Like N. Slight Minute. RKlA.
cytology: Associated with In(l)N264-84 = In(l)3C6- cytology: Associated with Df(l)N264-93 = Df(l)3B4-
7;20A-B (Sutton). Cl;3F3-4 (Sutton).
f{2 64.85 other information: Minute phenotype results from in-
origin: X ray induced. clusion of M(1)3E in the deficiency.
discoverer Demerec, 39d. *N264.94
references: 1940, Genetics 25: 618-27. origin: X ray induced,
phenotype: Like N. N264'8S/fa variegates for fa. discoverer: Demerec, 39k.
RKlA. phenotype: Like N. RK1.
cytology: Associated with T(l;2;4y<264-ss = cytology: Salivary chromosomes normal (Sutton).
T(l;2;4)3B4-Cl;6A2-Bl;60A4-5;101 F-102A (Sutton, other information: w, rst, and dm not affected.
1940, Genetics 25: 534-40). *N264.9S
origin: X ray induced simultaneously with rst264'86. discoverer: Demerec, 39k.
discoverer: Demerec, 39i. phenotype: Like N. RK1.
references: 1940, Genetics 25: 618—27. cytology: Salivary chromosomes normal (Sutton).
Demerec and Sutton, 1940, Proc. Natl. Acad. Sci. other information: w, rst, and dm not affected.
U.S. 26: 532-36. *N2 64.96
Sutton, 1940, Genetics 25: 534-40. origin: X ray induced.
phenotype: Like N. N264'86/fa variegates for ia. discoverer: Demerec, 39k.
RK1A. phenotype: Like N. RKlA.
cytology: Associated with T(l;4^264"86 = cytology: Associated with Df(l)N264-96 = Df(l)3C6-
T(l;4)3C6-.7;3C7-8i3E5-6;l 01F. 7;3C7-8 (Sutton).
•/SJ264-87 *N264.97
discoverer: Demerec, 39j. discoverer: Demerec, 39k.
references: Sutton, 1940, Genetics 25: 534—40. phenotype: Like N. RK1.
phenotype: Like N. RK1A. cytology: Salivary chromosomes normal (Sutton).
cytology: Associated with T(l;2;3^l364m87 * other information: w, rst, and dm not affected.
T(l;2;3)3C7-9;10A2'Bl;45F-46A;59F-60A;97C- *N264-99
D;100E-F. origin: X ray induced,
discoverer. Demerec, 40a.
origin: X ray induced. phenotype: Like N. RKlA.
discoverer: Demerec, 39j. cytology: Associated with Df(iyN264'99 = Dt(l)2D2-
phenotyp*: Like N. RK1. 3;3C11-12 (Sutton).
MUTATIONS 177
*N264.100 phenotype: Heterozygous females Notch and slight

origin: X ray induced. Minute. RK1A.
discoverer: Demerec, 391. cytology: Associated with In(l)N264'108 =
references: 1940, Genetics 25: 618-27. In(l)3C3-5;3E7-8;20A4-5 (Sutton).
phenotype: Like N. N264'100/ta variegates for fa, other information: Minute phenotype results from
RK1A. absence of section 3C5-3E7, which contains
cytology: Associated with T(1;3)N264'100 = M(1)3E, from the inversion.
T(l;3)3B4-Cl;4B4-5;80 (Sutton, 1940, Genetics 25: N264.109
534_40). origin: X ray induced.
N26 4-iooT2: see In(3L)100r2 discoverer: Demerec, 40a.
*N264.101 phenotype: Like N, but semilethal with fano. RK1.
origin: X ray induced. cytology: Salivary chromosomes normal.
discoverer: Demerec, 40a. other information: Located to the right of spl
phenotype: Like N. RK1A. (Welshons, Von Halle, and Scandlyn). w, rst, and
cytology: Associated with Df(l)N264-1Q1 - dm not affected.
Df(l)3C4-5;3C7-8 (Sutton). JO
origin: X ray induced. discoverer: Demerec, 40a.
discoverer: Demerec, 391. phenotype: Like N. RK1A.
phenotype: Like N. RK1A. cytology: Associated with Df(l)N264-110 =
cytology: Associated with Tfl&yN264'1 02 _ Df(l)3B4-Cl;3D2-3 (Sutton).
T(l;2)3C6-7;50E;56C (Sutton). N2 64.111
N264.103 origin: X ray induced.
origin: X ray induced. discoverer: Demerec, 40b.
discoverer: Demerec, 40a. phenotype: Like N. RK1A.
phenotype: Like N. N264ml03/spl shows variable cytology: Associated with DîyN264"111 =
expression of spl, as though mottled. [y264~lO3/fa Df(l)3C3-5;3C12-Dl (Sutton).
is facet. RK1.
cytology: Salivary chromosomes appear normal. origin: X ray induced.
other information: Located between spl and Ni24 discoverer: Dotnerec, 40b.
(Welshons and Von Halle, 1962, Genetics 47: phenotype: Like N. RK1A.
743—59). pn, w, r$t, and dm not affected. cytology: Associated with ln(l)N264-112 =
+N264.104 In(l)3C6-7;3F5-6 (Sutton).
origin: X ray induced. *N264-113
discoverer: Demerec, 39j. origin: X ray induced.
phenotype: Like TV. RK1A. discoverer: Demerec, 40c.
cytology: Associated with T(1;3)N264-104 = references: Sutton, 1940, Genetics 25: 628—35.
T(1;3)3C7-9;87D1-E1 + In(l)lB4-5;18-19 (Sutton). phenotype: Variegates for N and apl. RK2A.
N264-10S cytology: Associated with T(1;4)N264-113 ~
origin: X ray induced. TX1;4)3C10-D1;101 (Sutton).
discoverer: Demerec, 40a. *N264-114
phenotype: Like /V. RK1A. origin: Spontaneous.
cytology: Associated with Df(l)N264-105 - discoverer: Kaufmann, 40d.
D£(l)3C6~7;3D2-3 (Sutton). phenotype: Like N. RK1A.
+N264.106 cytology: Associated with Dt(l)N364-114 =
origin: X ray induced. Df(l)3C6-7;3D4~S (Sutton).
discoverer: Demerec, 40a. *H264.11S
phenotype: Like/V. RK1A. origin: X ray induced,
cytology: Associated with Df(l)N264"106 = discoverer: Sutton, 40e.
Df(l)3C6-7;3C7-8 (Sutton). phenotype: Like N. RK1A.
N264-/07 cytology: Associated with Dl(l)N264"115 =
origin: Spontaneous. D$1)3C3-5;3E2'3 (Sutton).
discoverer: Demerec, 40a.
phenotype: Like AT. RK1. origin: X ray induced,
cytology: Salivary chromosomes apparently normal. discoverer: Sutton, 40e.
other information: Locus seems to lie to the right of phenotype: Like N. RK1A.
•p/. Analysis complicated by a lethal between w« cytology: Associated w ~li6
and N264-*07 (Welshons, 1958, Proc, Natl. Acad, In(l)2C8-10;3C7-9 (Sutton).
Sci. U.S. 44: 254-58). rmt and tin not affected. *N264-117
*H264.10S origin: X ray induced,
origin; X ray induced, discoverer: Demerec, 40g.
discoverer: Demerec, 40a, phenotype: Like N. RK1A.
cytology: Associated with Df(l)N264"117 = cytology: Associated with Df(l)N264"127 =

Df(l)3A6-7;3E2-3 (Sutton). Df(l)3C6-7;3C7-8 (Sutton).
*N264-118
discoverer: Demerec, 40h. discoverer: Demerec, 41b.
phenotype: Like N. RK1A. phenotype: Like iV. RK1A.
cytology: Associated with Df(l)N264-118 = cytology: Associated with Di(l)N26*-128 =
Df(l)3C6-7;3C7-9 (Sutton). Df(l)3C6-7;3C7-8 (Sutton).
*N264.119 *N264.129
discoverer: Demerec, 40i. discoverer: Demerec, 41c.
phenotype: Like N. RK1. phenotype: Like N. RK1.
cytology: Salivary chromosomes normal (Sutton). cytology: Salivary chromosomes normal (Sutton).
other information: kz, w, rst, and dm not affected. other information: w, rst, and dm not affected.
*N264-120 +N2 64-130
discoverer: Demerec, 40j. discoverer: Neel, 41c.
phenotype: Like N. RK1A. references: 1942, Genetics 27: 530.
cytology: Associated with Df(l)N264-120 _ phenotype: Like N. RK1A.
Dt(l)3C6-7;3D2-3 (Sutton). cytology: Associated with Df(l)N264'i30 =
*N264.121 Di(l)3C6-7;3C7-8 (Sutton).
discoverer: Demerec, 40j. origin: X ray induced.
phenotype: Like N. RK1A. discoverer: Demerec, 41c.
cytology: Associated with T(1;3)N264.121 _ phenotype: Like N. RK1.
T(1;3)3C7'9;81F;86B6'C1 (Sutton). cytology: Salivary chromosomes normal (Sutton).
*N264.122 other information: w, rst, and dm not affected.
origin: X ray induced. /Va*: see N264-12
discoverer: Demerec, 40j. *N*: Notch of Aronson
phenotype: Like N. RK1. origin: Spontaneous.
cytology: Salivary chromosomes normal (Sutton). discoverer: Aronson, 57gll.
other information: kz, w, rst, dm, and ec not references: 1958, DIS 32: 67.
affected. phenotype: Like N. RK1A.
*N264.123 cytology: Several bands to the right of 3C4 de-
origin: Ultraviolet induced. ranged.
discoverer: Demerec, 40k. *WS; Notch of Bernstein
phenotype: Like N. RK1. origin: Spontaneous.
cytology: Salivary chromosomes normal (Sutton). discoverer: Bernstein, 28a7.
other information: w, rst, dm, and ec not affected. phenotype: Like N. RK1A.
*N264.U4 cytology: Associated with Df(l)NB = D((1)3C4-
origin: X ray induced. 5;3C12-D1 (Sutton).
discoverer: Demerec, 41a. Nc°: Notch-Confluens
phenotype: Like/V. RK1. origin: Spontaneous.
cytology: Salivary chromosomes normal (Sutton). discoverer: Welshons, 1955.
other information: w, rst, and dm not affected. references: 1956, DIS 30: 79.
*N264.125 1958, Cold Spring Harbor Symp. Quant. Biol. 23:
origin: X ray induced. 171-76.
discoverer: Demerec, 41a. phenotype: Wing tips seldom notched; veins thick-
phenotype: Like N. RK1A. ened; deltas present at juncture of longitudinals
cytology: Associated with Df(l)N2e>'t"12S m and marginal vein. Acrostichal rows irregular.
Df(l)3C4-5;3C7~8 (Sutton). Nc°/fa and NCo/$pl are facet and split, respec-
*N264.126 tively. NCo/lan wings more deeply notched than
origin: Spontaneous, /V Co /+. NCo/tano lethal; rare survivors sterile and
discoverer: Bishop, 401. weak. NCo/Dp(t;l)Co similar to Dp(l;l)Co/
phenotype: Like N. RK1A. Dp(l;l)Co. RK1.
cytology: Associated with DffiyN264'126 = cytology: Salivary chromosomes appear normal.
Df(l)3C3-S;3D4-5 (Sutton). other Information: Located between Ni24 and
N&Ogll (Welshons, 1958; Welshons and Von Halle,
*N264-127 1962, Genetics 47: 743-59).
discoverer: Demerec, 41b. origin: Spontaneous.
phenotype: Like N. RK1A. discoverer: Morgan, 1929.
MUTATIONS 179
references: 1937, DIS 7: 7. phenotype: Wings notched apically. Sometimes over-

phenotype: Like N. RK1A. laps wild type. Homozygous lethal. RK2A.
cytology: Associated with Df(l)NEZ =Df(l)3C6- cytology: Associated with In(2R)G = In(2R)50E;54D
7;3C7-8 (Sutton). (T. Hinton). May be inseparable.
*HG: Notch of Goidschmidt *N-b: Notch-b
origin: Found among progeny of heat-treated flies. location: 2- (not located).
discoverer: Goidschmidt. origin: Spontaneous.
references: Gottschewski, 1935, DIS 4: 15, 16. discoverer: Mann, 1921.
1937, Z. Induktive Abstammungs- Vererbungslehre synonym: Notch J2.
73: 131-42 (fig.1), references: 1923, Genetics 8: 27—36.
phenotype: Like N. RK1. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
cytology: Salivary chromosomes normal. Genet. 2: 232.
phenotype: Resembles Notch. Wing nicked in about
origin: Spontaneous (as cluster of two maternal 10 percent of heterozygous flies. Homozygote
chromosomes carrying fa). probably lethal. RK3.
discoverer: Welshons, 1955. other information: Possibly a vg allele.
synonym: N22 (Welshons, 1958). na: narrow abdomen
references: 1958, Cold Spring Harbor Symp. Quant. location: 1-45.2.
Biol. 23: 171-76. origin: X ray induced.
phenotype: Like N. fa Ni24/fa" ° lethal. RK1. discoverer: H. M. Miller, 34c.
cytology: Salivary chromosomes normal (Welshons). references: 1934, DIS 2: 9.
other information: Located between N264-103 a n ( j 1935, DIS 4: 9.
NCo (Welshons and Von Halle, 1962, Genetics 47: phenotype: Abdomen long and cylindrical in both
743-59). iv not affected. sexes. Viability low; female fertility low. Ovaries
in juvenile condition (Brehme). RK2.
M
: Notch of M/sc/iaikow *na2
origin: Spontaneous in the In(l)dl-49, y w f com- origin: Ultraviolet induced.
ponent of C(1)DX. discoverer: Edmonds on, 51g.
discoverer: Mischaikow, 561. references: 1952, DIS 26: 60.
references: Cicak and Oster, 1957, DIS 31: 80. phenotype: Like na. RK2.
phenotype: Wings notched at tips and occasionally
at sides. Veins thickened, with deltas. Eyes narrow: see nw
slightly smaller than normal; occasionally, one eye narrow abdomen: see na
extremely small. RK1. narrow eye: see ney
: Notch of NicoIetti narrow scoop: see nrs
discoverer: Nicoletti. nd: notchoid
phenotype: Like N. N^tc/fano lethal. RK1. location: 1-3.0.
cytology: Salivary chromosomes normal (Welshons). origin: Induced by trie thy lenemelamine (CB. 1246).
other information: Located between spl and N264'40 discoverer: Fahmy, 1951.
(Welshons and Von Halle, 1962, Genetics 47: 743- synonym: ntah; notch-Fahmy*
59). wnot affected. references: 1958, DIS 32: 72.
NP: Notch from P*2 phenotype: Wings notched; veins thickened. At low
origin: Induced by P 3 2 . temperature, wings not notched. Viability and fer-
discoverer: Bateman, 1950. tility of both sexes excellent, nd heterozygotes
phenotype: Like N. RK1A. with N2<S4-39t NCOi OTN264-40 h a v e extremely
cytology: Associated with In(l)Np = ln(l)3C;8E notched and strap like wings, small rough eyes,
(Darby). low viability and fertility (Welshons). fano/nd has
N*: Notch of Williams slightly thickened wing veins, with deltas. About
origin: Spontaneous in the In(l)scs, I component of 10 percent of fa/nd flies have small notches in one
C(1)DX. or both wings, spl/nd lacks a few bristles, like
discoverer: Williams, 56j. spl/+* Eyes sometimes smaller than normal and
references: Cicak and Oster, 1957, DIS 31: 80. roughened, spl nd males have rough eyes, ncf-like
phenotype: Like N. RK1. wings, and irregular, bushy sex combs. RK1.
N-: see 1(2)S cytology: Salivary chromosomes normal (Fahmy).
N-2G: Notch-2 from Gallup Placed in band 3C7, on basis of interaction with N.
location: 2-72.0. other information: Member of Notch pseudoallelic
origin: Spontaneous. series; located at or to the right otN60^11
discoverer: Ives, 41117. (Welshons and Von Halle, 1962, Genetics 47:
synonym: /V-2. 743-59).
references: 1943, DIS 17: 50. nd*
1957, DIS 31: 83. discoverer: R. M. Valencia, 62dl6.
phenotype: nd^/nd2 and nd2/nd like nd/nd. Normal other information: Not an allele of ix (Maynard
in combination with other visible mutations at the Smith).
N locus. RK1. *ney: narrow eye
other information: Close to but to the right of nd location: 1- (rearrangement).
(Welshons). origin: X ray induced.
*ne: nicked eye discoverer: Becker, 1950.
location: 2- (not located). references: 1952, DIS 26: 69.
discoverer: Kfil. phenotype: Homozygote has narrow eyes haliway be-
references: 1946, DIS 20: 66. tween B and wild type. Heterozygote usually
phenotype: Eye margin nicked. Overlaps wild type. normal. RK1A.
RK3. cytology: Associated with In(l)ney = In(l)10A;16D.
other information: Probably an allele of L. *ni: nicked
location: 3-40 (35 to 45).
discoverer: Neel, 41c26.
phenotype: Small notches or nicks in wing tips of
60—90 percent of homozygous males and 80—100
percent of homozygous females. RK3.
*ni-2: nicked on chromosome 2
net: net discoverer: Travers, 1955.
Edith M. Wallace, unpublished. references: Clarke, 1957, DIS 31: 80.
phenotype: Wing tips deeply emarginate between L2
net: net and L4 and occasionally between L4 and L5. Pen-
location: 2-0.0 [to the left of al; order with l(2)gl etrance and viability good. RK3.
not known]. nicked: see ni
origin: Spontaneous. nicked eye: see ne
discoverer: Bridges, 31cl0. no-wings: see ap3
phenotype: Wing veins form plexus like net; first Notch: see N
posterior cell between L3 and L4 widens toward Notch 2: see N-b
tip; branch missing from posterior crossvein; all notchoid: see nd
veins fused at base of wing, like bi. According to notchy: see ny
Waddington [l940, J. Genet. 41: 75-139 (fig.)], Notopleural: see Np
spaces form between epithelial layers owing to in-
adequate contraction during pupal period; spaces
later fuse and form extra veins. RK1.
cytology: Locus of net lies between 21A1 and 21C1
(Lewis, 1945, Genetics 30: 137-66).
*net*
discoverer: Braun, 1937.
phenotype: Like net. RK1.
discoverer: Williams, 56f.
phenotype; Wings have extreme plexus of veins, but
otherwise less abnormal than nef. RK1.
origin: Probably spontaneous.

discoverer: Meyer, 56c.
phenotype: Like net3; less extreme than net, RK1.
*JMW; neuter Np: Notopleural
location: Autosomal. From Bridges, Skoog, and L i , 1936, Genetics 21: 788-
origin: Spontaneous. 95.
discoverer: Travers, 1955.
reference*: Clarke, 1957, DIS 31; ISO. *Np; Notopleural
phenotype: Homozygous female intersex; homozy- location: 2-58.7 to 60.2 (between en and en; insep-
gous male normal. RK3. arable from Wo).
MUTATIONS 181
origin: Spontaneous. phenotype: Wings very small, opaque, curved spoon-

discoverer: Nichols-Skoog, 33b20. like up or down; inflated at eclosion. Wing margins
references: Bridges, Skoog, and Li, 1936, Genetics interrupted. Only one vein (L2 or L3) present.
21: 788-95 (fig.). Halters somewhat reduced. Viability excellent.
Li, 1936, Peking Nat. Hist. Bull. 11: 39-48. RK1.
phenotype: Notopleural, humeral, presutural, and cytology: Not included in Di(2L)64j = Df(2L)34E5-
pretarsal bristles shorter and blunter than normal. F1.-35C3-D1 (E. H. Grell).
Wings short and broad. Female produces few or no nub*
progeny. Viability fair. Development retarded. origin: Probably X ray induced.
More extreme at 19°C than at 25°, also more ex- discoverer: R. F. Grell, 56fl.
treme in female. Lethal over T(2;3)dp, Homozy- references: 1956, DIS 30: 71.
gous lethal. RK2A. phenotype: Wings small and spoonlike but less ex-
cytology: Locus lies between 44F1 and 45E2, on treme than nub. Patches of dried blood on wings.
basis of its association with Df(2R)Np = Veins LI to L4 almost indiscernible; L5 and alula
Df(2R)44Fl-2;45El-2 (Bridges). frequently absent. Viability and fertility excellent.
nts: narrow scoop RK1.
location: 1-54.2.
origin: Induced by 2-chloroethyl methanesulfonate origin: X ray induced.
(CB. 1506). discoverer Seiger, 62d.
discoverer: Fahmy, 1956. references: 1963, DIS 37: 53.
references: 1959, DIS 33: 88. Abbadessa and Burdick, 1963, DIS 37: 54.
phenotype: Wings narrow and slightly shorter than phenotype: Wings very small and spoonlike. RK1.
normal; frequently scooped. Slightly thinner Nude: see Nu
bristles. Eyes large and dull red. Eye and body *nw: narrow
colors darken with age. Viability and fertility location: 2-83.
good in male; fertility low in female. RK2. origin: Spontaneous.
Ns: Nasobemia discoverer: Bridges, 16b7.
location: 3-48.0. (No recombinants with pP among references: Morgan, Bridges, and Sturtevant, 1925,
1472 flies.) Bibliog. Genet. 2: 231.
origin: Spontaneous. phenotype: Wings long, narrow, and somewhat
discoverer: Gehring. pointed. Low viability and fertility in both sexes.
phenotype: In extreme cases Ns/+ forms, in place of At 25°C may overlap wild type; at 19° nearly all
an antenna, a complete leg including sternopleura, flies approach wild type but have longer wings.
coxa, trochanter, femur, tibia, and tarsus. Antennal RK2.
leg has no sex comb in male, and bristle pattern is
that of a middle leg. Eyes smaller; whole head
tends to be malformed. Expression variable but
penetrance complete. Hotnozygous lethal.
iVs ss"/+ ss" indistinguishable from Ns/+.
Ns/AntpB viable; phenotype like extreme Ns/+.
RK1.
cytology: Salivary chromosomes appear normal,
other information: Allelism with Antp not excluded
since all Antp alleles are associated with inver- nw 2 : narrow-2
sions that eliminate recombination in this region. From Payne, 1924, Genetics 9: 327-42.
However, all heterozygotes of Antp alleles are
lethal, unlike Ns/AntpB. origin: Spontaneous.
*Nu: Nude discoverer: Payne, 1615.
location: 2- or 3- (rearrangement). synonym: l&nce.
origin: X ray induced. references: Payne, 1924, Genetics 9: 327-42 (fig.).
discoverer: Sutton, 41a27. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
phenotype: Many bristles missing from head and Genet. 2: 227.
thorax; post scute liars, notopkurals, verticals, and phenotype: Wings like nw. Classification easier in
postverticals usually present. Homozygous lethal. females. Slight notching or tufting of marginal
RK2A. hairs on tip of wings. Both sexes nearly sterile.
cytology: Associated with T(2;3)Nu - Tf2;3)24;J6- Ovaries tumorous at eclosion [King, Burnett, and
37;39-40;73-74;7$-76;77-78;8l-82;8S-86;89-90. Staley, 1957, Growth 21: 239-61 (fig.); King, 1964,
nub: nubbin Roy. Entomol. Soc. (London) Symp. Insect Repro-
location: 2-47.0. duction pp. 13—25J. Oogonia proliferate asynchron-
origin: Spontaneous. ously within ovariole; follicle development in-
discoverer: Mickey, 48elO. hibited [Beatty, 1949, Proc. Roy. Soc. Edinburgh,
references: 1949, DIS 23: 61. B 63: 249-70 (fig.)]. RK2.
nwD: narrow-Dominant female viability about 40 percent wild type and

origin: X ray induced. fertility reduced. RK2.
discoverer: E. H. Grell, 59f. other information: One allele induced by CB. 1506
references: 1962, DIS 36: 37. oblique: see ob
phenotype: Wings of heterozygote longer and nar- oblique wings: see obl
rower than normal. Expression variable and some- obt: obtuse
times approaches wild type. Viability oi nwD/+ location: 3-77.5.
low. Homozygous lethal, as is nwD/nw2. RK2. discoverer: E. M. Wallace, 35gl.
NX: Notch Xasta phenotype: Wings shorter and blunter, but overlap
location: 3- (between st and Dfd; 44.0-47.5). wild type slightly. Thorax somewhat humpy; body
origin: X ray induced. chunky; eyes slightly bulging. RK3.
discoverer: Ohnishi, 49116.
1951, DIS 25: 79.
Schalet, 1960, DIS 34r 55.
phenotype: Resembles Notch but more extreme. Ho-
mozygote resembles Xasta. Viability of heterozy-
gote fair; homozygote semilethal. Enhanced by Dl
and suppressed byH. Combination of NX and apXa
produces small wings, like vg, and lower viability.
RK2 as heterozygote. oc: ocelliless
oc ocelliless
location: 1-23.1.
discoverer: Bedichek, 30cl5.
phenotype: Ocelli completely absent. Bristles in
ocellar area and on top of head irregular and more
numerous; postverticals usually absent. Eyes
ny: notchy somewhat reduced and body size dwarfed. Viability
From Griinberg, 1929, Biol. Zentr. 49: 680-94. about 90 percent wild type. Females sterile.
According to Beatty Ll949, Proc. Roy. Soc.
ny: notchy Edinburgh, B 63: 249-70 (fig.)J, oocytes often mis-
location: 1-32. shapen, eggs abnormal in appearance, and par-
origin: X ray induced. ovaria nearly always absent. RK2.
discoverer: Gruneberg, 28j29. cytology: Salivary chromosome studies by Demerec
references: 1929, Biol. Zentr. 49: 680-94 (fig.). and Sutton show locus to lie between 7C4-5 and
1934, DIS 2: 8. 8C1-2 (Demerec, Kaufmann, Fano, Sutton, and
phenotype: Wing tips slightly nicked. Expression Sansome, 1942, Carnegie Inst. Wash. Year Book
variable; overlaps wild type in some females and 41: 191). Further restricted to 7E1 through 8C2,
most males. Viability about 70 percent wild type. on the basis of its exclusion from Df(l)sn =
RK3. D%1)7B2-3;7D22-E1 (Hinton and Welshons, 1955,
DIS 29: 125-26).
*ob: oblique
location: 1-37.2. Oce: Ocellarless
discoverer: Neel, 41f30. origin: Induced by triethylenemelamine (CB. 1246).
references: 1942, Genetics 27: 532. discoverer. Fahmy, 1953.
1942, DIS 16: 51. references: 1958, DIS 32: 72.
phenotype: Wings obliquely truncated from inner phenotype: One or both ocellar bristles and fre-
margin outward. Venation disturbed. Viability quently postverticals missing; other bristles,
about 20 percent wild type. RK3. especially the scutellars, sometimes absent.
obl: oblique wings Wings frequently positioned abnormally, with
location: 1-60.1. incised margins; effect more marked in homozygous
origin: Induced by triethylenemelamine (CB. 1246). females. Bristle effect dominant. Good viability
discoverer: Fahmy, 1953. and fertility in both sexes. RK1.
references: 1958, DIS 32: 72. other information: One allele induced by each of the
phenotype: Wings slightly upheld and outspread; following: CB. 3025, CB. 1592, CB. 1540, and
small blister occasionally present. Body color CB. 1528.
slightly darker. Male viability and fertility good; ocelliless: see oc
MUTATIONS 183
*ocr: ochracea omm: ommatoreductum

location: 2-0. location: 1-12.8.
discoverer: Serebrovsky, 40g25. origin*. Induced by triethylenemelamine (CB. 1246).
references: 1941, DIS 15: 19. discoverer. Fahmy, 1953.
phenotype: Eye color lighter at eclosion, darkening references: 1958, DIS 32: 72.
with age. RK1. phenotype: Some peripheral ommatidia absent, fre-
od: see os° quently in an irregular manner, giving a rough eye
OdhF: Octanol dehydrogenase-Fast and a notched border. Shape of head abnormal;
location: 3-49.2. head bristles deranged or absent. Palps absent or
origin: Naturally occurring allele. deformed. Thoracic bristles deranged. Wings often
discoverer: Ursprung. unexpanded. Good viability and fertility in both
references: Ursprung and Leone, 1965, J. Exptl. sexes. RK2.
Zool. loV): 147-54. other information: One ^llele each induced by
Courtright, 1966, DIS 41: 59. CB. 1246, CB. 1522, CB. 1592, CB. 1528; two
Courtright, Imberski, and Ursprung, 1966, Genetics alleles induced by CB. 3026.
54: 1251-60. ommatidia: see o/n
phenotype: Produces octanol dehydrogenase that ommatoreductum: see omm
migrates more rapidly to cathode in agar gel elec- *0n: Open
trophoresis at pH 8.7 than Odhs. OdhF/Odhs het- location: 3-26.
erozygote produces enzyme of intermediate mo- origin: X ray induced.
bility in addition to fast and slow types. Hexanol discoverer: Tanaka, 36c26.
and heptanol, as well as octanol, are substrates references: 1937, DIS 7: 21.
for the enzyme. RK3. 1937, DIS 8: 11.
Odhs: Octanol dehydrogenase-Slow phenotype: Wings spread, Homozygous viable. RK2.
origin: Naturally occurring allele. *op: opaque
discoverer: Ursprung. location: 1-50.
references: Ursprung and Leone, 1965, J. Exptl. origin: X ray induced.
Zool. 160: 147-54. discoverer: H. M. Miller, 33k.
Courtright, 1966, DIS 41: 59. references: 1934, DIS 2: 9.
Courtright, Imberski, and Ursprung, 1966, Genetics 1935, DIS 3: 14.
54: 1251-60. 1935, DIS 4: 10.
phenotype: Produces octanol dehydrogenase that phenotype: Wings opaque and whitish, usually diver-
migrates less rapidly to cathode in agar gel elec- gent and slightly convex. Viability and fertility
trophoresis at pH 8.7 than OdhF. RK3. good in male, poorer in female. RK3.
odsy: see os *opb: opaque broad
Of: see Diof location: 1-28.3.
*Off: Off origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
location: 2-82. phenylalanine (CB. 3025).
origin: Spontaneous. discoverer: Fahmy, 1955.
discoverer: Bridges, 23el4. references: 1959, DIS 33: 88.
references: Morgan, Bridges, and Sturtevant, 1925, phenotype: Short, broad, and opaque wings, with
Bibliog. Genet. 2: 232. slightly convex or concave membranes. Slightly
phenotype: Some bristles missing in heterozygotes, brownish eye color. Legs short with long segments
especially from side of abdomen; basal rings frequently bowed. Abdomen slightly abnormal in
remain as in H. Homozygote lacks more bristles. shape, and genitalia deformed. Males fertile;
Eyes large, creased, and roughened. RK2. viability about 10 percent wild type. Females
other information: Agrees with abr in locus and sterile. RK3.
description; may have been an allele. Open: see On
ol-2: see sp *op/i; ophthalmopedia
olive-2; see sp location: 2-45.
olvD: see dp° origin: Spontaneous.
*om; ommatidia discoverer: Gordon, 1934.
location: 1-0.1 (to the right of sc). references: 1936, J. Genet. 33: 25—60.
origin: X ray induced in (or with) ac3. 1941, DIS 14: 39.
discoverer: Muller. phenotype: In extreme form, an appendage grows
references: Muller, Prokofyeva, and Raff el, 1935, from eye; in less extreme form, eye is kidney
Nature 135: 253-55. shaped. Expression sensitive to genetic and
Muller, 1935, DIS 3: 30. environmental modification. Effect caused by
phenotype: Ommatidia disarranged, giving a slight enlargement and abnormal folding of eye-forming
eye roughness difficult to classify. RK3. portion of optic disk in late larvae IWaddingtan and
cytology: Thought by Muller to be in or very close Pilkington, 1943, J. Genet. 45: 44-50 (fig.)]. RK3.
to 1CI. opt: see
or: orange references: 1930, J. Genet. 22: 303 (fig.).

location: 2-107.2 (to the left of Fo). 1935, DIS 3: 30.
origin: Spontaneous. Verderosa and Muller, 1954, Genetics 39: 999.
discoverer: Mossige, 1942. phenotype: Wings extremely divergent, often at right
references: 1950, DIS 24: 61. angles to body. os°/os8 is wild type. RK1.
phenotype: Eye color bright orange, or/pd wild type oss: outstretched small eye-small eye
(Von Halle). RK1. origin: Spontaneous.
discoverer: Bridges, 19g3.
origin: Spontaneous in In(2L)Cy + In(2R)Cy, synonym: sy.
Cy en3 spa, probably simultaneously with bw4Sa. references: Morgan, Bridges, and Sturtevant, 1925,
discoverer Ives, 45a. Bibliog. Genet. 2: 236.
references: 1951, DIS 25: 70. phenotype: Eyes small and rounded, high on the
phenotype: Eye color like or, RK1. head, but not bulging. RK1.
or-m *osh: outshifted
origin: Spontaneous, location: 1-33.0 (no crossover with v in 997 chromo-
discoverer: Ives, 49h31. somes).
references: 1951, DIS 25: 70. origin: Induced by 2-chloroethyl methanesulfonate
phenotype: Eye color like or. RK1. (CB. 1506).
os: outstretched small eye discoverer: Fahmy, 1955.
origin: X ray induced. phenotype: Wings shortened and often slightly
discoverer: Abrahams on, 1953. divergent. Body and wings pale in color. Eyes
synonym: odsy. somewhat smaller and browner than normal. Via-
references: Verderosa and Muller, 1954, Genetics bility and fertility good in both sexes. RK2.
39: 999. ot: outhefd
phenotype: Wings held virtually at right angles to location: 1-65.7.
body. Eye small and rounded. os/os° has wing origin: Induced by triethylenemelamine (CB. 1246).
effect but eyes normal, os/os8 has eye effect but discoverer Fahmy, 1952.
wings normal. RK1. references: 1958, DIS 32: 73.
cytology: Placed in region 16E-17A, on the basis of phenotype: Wings held horizontally; inner margin
its being to the right of Dt(l)C-PL = Dl(l)15F;16E slightly cut away in many males. Ocellar bristles
and to the left of Dp(l;l)Bxr4*k =Dp(l;l)17A;17C. usually absent or reduced; effect variable. Hairs
osbdw; outstretched small eye-bending wings sparse, especially in posterior midthoracic region.
origin: X ray induced. Males sterile; viability about 20 percent wild type.
discoverer: Halfer, I960, RK3.
synonym: bdw. outshifted: see ash
phenotype: Wings divergent and drooping; size and outstretched small eye: see os
shape normal. Males sterile. RK2A. outstretched: see os°
cytology: Associated with T(l;3)osbdw = *ov: oval
T(l;3)16E;80C. location: 1-17.5.
discoverer: Steinberg, 37hl5.
phenotype: Eye somewhat oval and quite rough.
RK1.
ovaless: see ov/
over overetherized
discoverer Plaine and Aubele, 64b.
phenotype: Wings held vertically within 1 hr after
eclosion; vibrate feebly but are incapable of sup-
porting flight. Movements of first two pairs of legs
uncoordinated. Viable and fertile although ove
male often unsuccessful in mating with ove + fe-
male. RK2.
os°: outstretched small eye-outstretched Overflow: see Dl°t
Edith M. Wallace, unpublished. *ov7: ovioculus
location: 1-0.9.
os°: outstretched small eye-outstretched origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
origin: X ray induced. phenylalanine (CB. 3007).
discoverer: Muller, 1930. discoverer Fahmy, 1953.
synonym: od. references: 1958, DIS 32: 73.
ONS 185
notype: Eyes small, egg shaped, and rough. discoverer: Alexander.

Ings spread or elevated to varying degrees; edges references: Ward and Alexander, 1957, Genetics 42:
cised, especially inner margin. Eclosion slightly 42-54.
ilayed. Males sterile. Viability 20—60 percent phenotype: Eye color pink. RK1A.
Lid type. RK2. cytology: Associated with In^Rfe1 00.290 -
ivaless In(3R)85B3-4;85D12'15.
ation: 2- (not located). p<"; see Pu&*
I in: Spontaneous. pp: pink-peach
coverer: Bridges, 21a3. discoverer: Bridges, 13a24.
srences: Morgan, Bridges, and Sturtevant, 1925, references: Bridges and Morgan, 1923, Carnegie
Bibliog. Genet. 2: 232. Inst. Wash. Publ. No. 327: 82 (fig.).
notype: Rough eyes. Males fertile; females phenotype: Eye color lighter and more orange than
itirely sterile. Small groups of cells in place of p. Eyes have 9 percent normal red and 15 percent
r
aries, but ducts and genitalia normal. Abdomen normal brown pigment (Nolte, 1959, Heredity 13:
female grayish and translucent. RK3. 233—41); become brown with age. In combination
>k with en, eyes orange-red in young flies, darkening
ation: 3-48-.0. toward deep red with age; with bw, eyes light
I in: Spontaneous. reddish yellow to rose-brown, darkening with age;
:overer: Morgan, lOg. color autonomous in larval optic disk transplanted
irences: Bridges and Morgan, 1923, Carnegie into wild-type host (Beadle and Ephrussi, 1936,
[nst. Wash. Publ. No. 327: 44 (fig.). Genetics 21: 230). Larval Malpighian tubes color-
notype: Eye color dull ruby with purplish tone, less (Beadle, 1937, Genetics 22: 587-611). Fe-
res contain 40 percent normal red and 33 percent males heterozygous for pP and for a white allele
>rmal brown pigment (Nolte, 1959, Heredity, 13: (e.g., w, wh, wbf) have brownish eye (Judd, 1955,
•3—41). Larval Malpighian tubes colorless DIS 29: 126). RK1.
irehme and Demerec, 1942, Growth 6: 351—56). *pPS6
K2. origin: Spontaneous.
>logy: Tentatively placed in region 85A6-B3, on discoverer: Williams, 56c.
e basis of position of the breakpoint common to references: 1956, DIS 30: 80.
(3)pl 00.48 =ln(3)80Sl;85A6-Bl and phenotype: Eye color light ruby with orange tone.
(3R)p1(>0.290 = in(3R)85B3-4;85D12-15 (Ward and RK1.
Lexander, 1957, Genetics 42: 42—54). P: Pale
location: 2- or 3- (rearrangement).
I in: Spontaneous. origin: Spontaneous.
coverer: Thoday, 53h. discoverer: Bridges, 17jl6.
srences: 1954, DIS 28: 78. references: Bridges and Morgan, 1923, Carnegie
notype: Like pP. RK1. Inst. Wash. Publ. No. 327: 184 (fig.).
phenotype: Heterozygote a specific dilutor of the w°
|in: Spontaneous. series of white alleles; tends to darken eye color
coverer: Williams, 56h. of w" series. Homozygous lethal. RK2A.
trences: 1956, DIS 30: 80. cytology: Associated with T(2;3)P = T(2;3)58E3-
notype: Like pP. RK1. F2;60D14-B2;96B5-Cl (Morgan, Bridges, and
48 Schultz, 1934, Carnegie Inst. Wash. Year Book 33:
jin: X ray induced. 278).
coverer Alexander. pa; patulous
»rences: Ward and Alexander, 1957, Genetics 42: location: 2-101,0.
tfiotype: Pink eye color. RK1A. discoverer: Edmonds on and Meyer, 49d.
ology: Associated with ln(3)pi00.48 = in(3)8Q- references: 1949, DIS 23: 61.
;85A6-B1. phenotype: Wings spread wide apart. Excellent via-
88 bility; fair fertility. RK1.
jin: X ray induced. cytology: Placed to the right of 58F2, on the basis
coverer: Alexander. of its being covered by Dp(2;3)P from T(2;3}P »
srences: Ward and Alexander, 1957, Genetics 42: T(2;3)58E3-F2;60D14-E2;96B5-Cl.
42-54. p&: see pt
notype: Eye color pink. Homozygous semilethal. *pads: pads
K2A. location: 2-55.
ology: Induced with ln{3)plOO.i8 m 1^3)80- origin: Spontaneous.
';94D11-EI, which does not involve the pink discoverer: Bridges, 17e9.
gion. references: Morgan, Bridges, and Sturtevant, 1925,
290 Bibliog. Genet. 2: 212 (fig.), 232.
jin: X ray induced. Stern, 1934, DIS 1: 36.
phenotype: Wings malformed; often remain in condi- phenotype: Transforms posterior metathoracic seg-
tion of those of newly emerged flies. RK2. ment into a posterior mesothoracic structure.
Transformation suppressed by Cbx. bx3 pbx ho-
origin: Spontaneous. mozygotes show virtually complete mesothoracic
discoverer: Mohr, 20bl5. transformation of the metathorax. bx3 +/+ pbx is
references: 1929, Z. Induktive Abstammungs- wild type. bxd pbx/+ + is wild type but bxd +/+
Vererbungslehre 50: 126. pbx shows moderate p6x-like transformation. RK3.
phenotype: Like pads. RK2. cytology: Locus probably in 89E3-4 (Lewis).
pads-b: see pu other information: The rightmost member of the
Pale: see P pseudoallelic series including, from left to right,
pale ocelli: see po bx, Cbx, Ubx, bxd, and p6x.
pale wing: see plw Pc: Poly comb
pallid: see pld location: 3-48 (0.3 unit to the left of Sex).
parted: see a&2 origin: X ray induced.
*pat: patchytergum discoverer: P. H. Lewis, 1947.
origin: Induced by triethylenemelamine (CB. 1246). Lewis, 1956, DIS 30: 76.
discoverer: Fahmy, 1952. Hannah-Alava, 1958, Genetics 43: 870-905.
references: 1958, DIS 32: 73. phenotype: Presence of sex combs (1—4 teeth) on
phenotype: Wings divergent. Pigmentation of an- second and third legs of male is most conspicuous
terior border of fifth tergite patchy. Ocelli light. effect. Other effects are elevated, divergent, or
Male sterile; viability about 10 percent wild type. crinkled wings, bent humeral and anterior noto-
RK3. pleural bristles, abnormal sternopleurals, terminal
other information: One allele induced by CB. 3007. gaps in L4, and leglike antennae — all are less
* patch: patched extreme in male than in female (or are absent in
location: 2- (not located). male). Homozygous lethal and lethal with Pc2 but
origin: Spontaneous. not with Sex. Enhances the Antennapedia pheno-
discoverer: Bridges, 13k25. type when mutually heterozygous with AntpYu and
references: Bridges and Morgan, 1919, Carnegie AntpB; in the latter, antennal leg is completely
Inst. Wash. Publ. No. 278: 241. expressed only in Pc ssa/AntpB ssB compound
phenotype: Abdominal sclerites fewer or sharply cut (Stern). Possibly lethal with Antp49 but not with
into triangular segments obliquely fitted together. Antp50. Expression of Pc enhanced in male het-
Overlaps wild type. RK3. erozygous for bx, bxd, and Ubx; enhancement more
patchytergum: see pat extreme when mutants (at least bx and bxd) are in
patulous: see pa coupling than in repulsion (Hannah-Alava, 1964,
pb: proboscipedia Z. Vererbungslehre 95: 1-9). RK2.
location: 3-47.7.
origin: Spontaneous, origin: X ray induced.
discoverer: Bridges, 31d27. discoverer: Puro, 61j.
references: Bridges and Dobzhansky, 1933, Arch. phenotype: Similar to Pc, but sex combs of male are
Entwicklungsmech. Organ. 127: 575—90 (fig.). larger and resemble those of Sex. Other pleiotropic
phenotype: Oral lobes changed to tarsus like or effects more extreme than in Pc. Enhances expres-
arista like appendages. Cold (15°C) shifts expression of Antp49 and Antp50; reduces viability of
sion toward arista like, heat (29°) toward tarsus like Antp50. RK2.
[Villee, 1944, J. Exptl. Zool. 96: 85-102 (fig.)]. Pch: see pyd
Temperature sensitive period in last larval instar pd: purpleoid
[Vogt, 1946, Z. Naturforsch. 1: 469-75 (flg.)L location: 2-106.4.
Very short lived because adults cannot feed. Male origin: Spontaneous.
fertile; female sterile. Ovaries normal but few if discoverer: Bridges, 16h31.
any eggs formed (Beatty, 1949, Proc. Roy. Soc. references: 1937, Cytologia (Tokyo), Fujii Jub. Vol.
Edinburgh, B 63: 249-70). RK2. 2: 745-55.
phenotype: Eye color dark pink or maroon, like pt
pbx: postbithorax but less extreme; 20 percent normal red pigment
location: 3-58.8 (to the right of bxd). and 61 percent normal brown pigment (Nolte, 1955,
origin: X ray induced (arose simultaneously with J. Genet. 53: 1—10). Semidominant; eye color of
Cbx). heterozygote duller than wild type; color autono-
discoverer: E. B. Lewis. mous in larval optic disk transplanted into wild-
references: 1954, Proc. Intern. Congr. Genet., 9th. type host (Beadle and Ephrussi, 1936, Genetics
Pt. 1: 100-5. 21: 230). Malpighian tubes wild type (Beadle,
1954, DIS 28: 76. 1937, Genetics 22: 587-611). RK2.
1955, Am. Naturalist 89: 73-89. cytology: Placed in region between 59E2 and 60B10
1963, Am. Zoologist 3: 33-56 (fig.). by Bridges (1937), on basis of its being to the right
MUTATIONS 187
of In(2R)bwVDel = In(2R)41B2-Cl;59E2-4 and to phenotype: Wings spread; fluid often accumulates

the left of Df(2R)Px = Df(2R)60B8-10;60Dl-2. between membranes. Degree of wing divergence
pdf: pod foot inversely correlated with temperature; wings more
location: 1-57.0. divergent in male. In transfers from 19°C to 31°,
origin: X ray induced. temperature-effective period begins 6—8 hr before
discoverer: Welshons, 57h6. eclosion in male and 4—6 hr before eclosion in fe-
references: 1960, DIS 34: 54. male and ends with eclosion. In transfers from
phenotype: Terminal tarsus swollen in one or more 31 °C to 19°, the temperature-sensitive period be-
legs. Classification, viability, and fertility good. gins 8—10 hr before eclosion and ends 2—4 hr be-
RK2A. fore eclosion (P. H. Baker, 1950). RK2.
cytology: Associated with In(l)pdf = In(l)16B;19F- *pg: prong
20A. Tentatively placed in 16A and at 57.0 since location: 2-40.
pdf is covered by BSY, but not by Ymal +2. discoverer: Mohr, 19e.
Pdr: Purpleoider Vererbungslehre 32: 218.
location: 3-46. phenotype: Extra crossveins distal to anterior
origin: Spontaneous. crossvein; usually incomplete. Overlaps wild type
discoverer: Bridges, 22f20. in at least 10 percent of flies. RK3.
phenotype: The combination pd/pd; Pdr/+ gives pg: see pig
lighter, yellower eye color than pd alone. pd/+; Pgd*: Phosphogluconate dehydrogenase-A
Pdr/Pdr has eye color like pd/pd. pd/pd; Pdt/Pdr location: 1-0.9.
is lethal. Pdr/Pdr is rosier than wild type. origin: Naturally occurring allele.
Pdr/Pdr and pd/pd; Pdr/+ JMalpighian tubes normal discoverer: Young.
(Brehme and Demerec, 1942, Growth 6: 351—56). references: Kazazian, Young, and Childs, 1965,
RK3. Science 150: 1601-2.
*pe: petit Young, 1966, J. Heredity 57: 58-60 (fig.).
location: 3- (not located). phenotype: Produces phosphogluconate dehydro-
origin: Spontaneous in In(3Li)P. genase that migrates faster in starch gel than that
discoverer: Mohr, 38k30. produced by PgdB. PgdA/PgdB produces, in addi-
references: 1939, DIS 12: 47. tion to the fast and slow bands, a hybrid band of
phenotype: Body small. Eyes small and rough. Via- intermediate mobility; hybrid enzyme may also be
bility good, but female fertility low. RK2A. produced in vitro. Male and female produce equiv-
Pearl: see PI alent enzyme levels. RK3.
peb: pebbled PgdB
location: 1-7.3 (0.4 unit to the right of bi). origin: Naturally occurring allele.
discoverer: Dubinin. discoverer: Young.
phenotype: Eyes markedly rough at 28—30°C, references: Kazazian, Young, and Childs, 1965,
slightly rough (like S) at 25°, and wild type at 19°. Science 150: 1601-2.
RK2 (28-30°C). Young, 1966, J. Heredity 57: 58-60 (fig.),
cytology: Placed in salivary chromosome region 4C7 phenotype: Produces a slow-migrating phosphoglu-
through 4D2 (Demerec, Kaufmann, Fano, Sutton, conate dehydrogenase. Enzyme level same in male
and Sansome, 1942, Carnegie Inst. Wash. Year and female. RK3.
Book 41: 191). pi: pied
pebbled: see rg* location: 2-17.
pentagon: see ptg origin: Spontaneous.
pers: persimmon discoverer: Harnly, 38k31.
location: 3- (left arm). phenotype: Eyes like 5 but more extreme, smaller
origin: X ray induced. and rougher, with facets jumbled. Wings larger,
discoverer: Demerec, 3712. flimsy, arched, and fringed. Male usually sterile,
references: 1940, DIS 14: 40. with abnormal genitalia. Viability erratic, varying
phenotype: Eye col6r dull orange. Larval Mal- from 20 to 80 percent. RK3.
pighian tubes colorless (Brehme, 1942, Genetics pic: piccolo
27: 133). Viability and fertility good. RK2A. location: 3-52.1.
cytology: Associated with In(3L)pmm • In(3L)63C2- origin: X ray induced in a kar2 chromosome.
5;73B2-5. discoverer: Sena let.
petit: see pe references: Schalet, Kernaghan, and Chovnick,
Pfd; Pufdi 1964, Genetic® 1261-68.
location: 2-70.8. phenotype: Bristles short and fine; fergite mor-
discoverer: Brkwley, 1935, phology abnormal as In fab. Inviable in combination
references: Shall, 1937, DIS 8: 10. with pic21 and pic31, Homosygottt sterile. RK2.
1938, Proc. Michigan Acad. ScL 23: 647—49. pic*h picco/o-2 lethal
Baker, 1950, Am. Naturalist 84: 51-70. origin: X ray induced in a kar% drome*orae.
discoverer: Sena let. lethal. bw + Y; Pin2/+ has bristles intermediate in

references: Schalet, Kernaghan, and Chovnick, length between Pin2/+ and wild type. RK1.
1964, Genetics 50: 1261-68. : Pin-Tack
phenotype: Lethal homozygous and in combination origin: Spontaneous.
with pic and pic^t. RK3. discoverer: Weiskettel, 571.
pic* I synonym: Tac.
origin: X ray induced in a kar2 chromosome. references: Kadel, 1958, DIS 32: 80.
discoverer Schalet. phenotype: At 22°C, thoracic bristles very small;
references: Schalet, Kernaghan, and Chovnick, other bristles not so small. At 18°, phenotype is
1964, Genetics 50: 1261-68. nearly wild type* Older female holds wings in ab-
phenotype: Lethal homozygous and in combination normal position. Homozygous lethal. RK1.
with pic and pic21. RK3. *: Pin-Yellow tip
pied: see pi origin: Spontaneous as one-half of a mosaic male.
*pig: pigmy discoverer: E. H. Grell, 57e.
location: 1-29. synonym: Ylt.
discoverer. Muller, 2618. phenotype: Distal third of thoracic bristles pale
synonym: pg (preoccupied). yellow, thin, and slightly twisted. Lethal homozy-
references: 1935, DIS 3: 30. gous and in combination with Pin and Pin2) sur-
phenotype: Fly small and melanotic. Viability about vives in combination with Df(2R)Px and resembles
25 percent wild type. RK3. PinY*/+. RK1.
Pigmentless: see Ps pink: see p
pigmy: see pig pink wing: see pw
*pil-3: pilosus in third chromosome pink-wing: see ltPk
location: 3- (near or identical with Ira). pinkish: see pkh
discoverer Goldschmidt. pinkaid: see ltPk
references: 1953, J. Exptl. Zool. 122: 53-96 (fig.). pk: prickle
phenotype: Produces setae on sixth sternite of male location: 2-55.3 (between ap and tuf).
or transformed female. Semidominant. Enhanced origin: Spontaneous.
by pil-X. RK3. discoverer: Ives, 38k.
*pil-X: pilosus in X references: 1947, DIS 21: 68-69.
location: 1- (left of w). phenotype: Posterior acrostichals irregularly erect
discoverer: Goldschmidt. and whorled. Lateral costal hairs of wing regularly
references: 1953, J. Exptl. Zool. 122: 53-96 (fig.). slanted anteriorly instead of posteriorly. Flies
phenotype: Produces setae of varying numbers and slightly larger than wild type. Occasional extra
sizes on the sixth sternite of male and of X/X; dorsocentral and scute liar bristles appear at tem-
tra/tra female. Effect enhanced by presence of peratures above 23°C. RK1.
pil-3 and also by Y chromosome of tra stock. RK3. cytology: Placed in salivary chromosome region
Pin: Pin 42A3-19, on the basis of its inclusion in
location: 2-107.3 (to the right of sp). In(2R)Cy = In(2R)42A2-3;58A4-Bl and its being to
origin: Spontaneous, the left of tuf, which is within Df(2R)MS2y6lt =
discoverer Ives, 39a9. Df(2R)40F-41Al;42A19-Bl (Sturtevant, 1949, DIS
references: 1940, DIS 13: 50. 23: 98).
phenotype: Thoracic bristles, especially dorso- *pkh: pinkish
centrals and scutellars, shortened and thick at location: 2-100.
base but tapering sharply. Strong in homozygote; discoverer: Bridges, 14g27.
heterozygote reliably classified. RK1. references: 1919, J. Exptl. Zool. 28: 365.
cytology: Located between 60C5 and 60D2, on the Bridges and Morgan, 1919, Carnegie Inst. Wash.
basis that Pin3 is lethal in combination with Publ. No. 278: 247 (fig.),
Df(2R)Px - Df(2R)6QB8-10rfODl-2 and Df(2R)Px2 = phenotype: Specific dilutor of we. RK3.
D%2R)60CS-6f60D9-10. pi: pleated
location: 1-47.9.
discoverer: E. H. Grell, 57b. discoverer: Moore, 31cl5.
phenotype: Thoracic bristles of heterozygote very phenotype: Wings folded lengthwise in pleats. Over-
short. At low temperature (17°C), heterozygote laps wild type at 25°C, more extreme at 19°. RK3.
appears normal. Homozygote usually lethal; rare cytology: Placed in salivary chromosome region
survivors have virtually no thoracic bristles. 13B2-F17, on basis of its being included in
Pin2/Pin has smaller bristles than Pin2/+ and low Dp(l;f)A12 ^Dp(l;f)lB-C;13Bl-S but not in the
viability. Pin2/PinYt is lethal. Df(2R)Px/Pin2, proximal part of the X derived from T(1;4)A4 =
D%2R)Px2{Pin2, undD§C2R)Px4/Pin2 are also T(l;4)13F6-14Al;102F (inferred from Patterson,
MUTATIONS 189
1938, Am. Naturalist 72: 193-206, also frontispiece

of Texas Univ. Publ. 4032). origin: X ray induced.
pi: see pld discoverer: Demerec, 28f30.
*PI: Pearl synonym: se-like 62.
location: 2-6. phenotype: Eye color like pn but lighter and more
origin: Spontaneous, ruby. Eyes contain 18 percent normal red pigment
discoverer: Rosin, 1948. and 114 percent normal brown pigment (Nolte,
references: 1951, DIS 25: 75. 1959, Heredity 13: 233-41). Larval Malpighian
1952, Rev. Suisse Zool. 59: 261-68. tube color normal (Beadle, 1937, Genetics 22:
Nef, 1958, Z. Vererbungslehre 89: 272-319 (fig.). 587-611). RK1.
phenotype: Heterozygote has pearl-like nodes in pn~
wings. Wing margins often snipped; venation dis- discoverer: Weinstein.
turbed. Bristle pattern defective. Eyes small and phenotype: Like pn. RK1.
rough. At 28°C, at least one of these characters
always present; at 18°, phenotype virtually normal. origin: X ray induced.
Viability good; fertility of male slightly reduced. discoverer: Glass, 1929.
Fraction of cells die in all imaginal disks. In wing references: 1934, DIS 2: 7.
disks, dead cells surrounded by epithelial cells 1935, DIS 3: 14.
and produce pearl-like structures in adult wing. phenotype: Like pn. RK1.
Homozygote dies as pupa (Tschanz). RK2. other information: Induced in, but separable from,
platinum: see pt In(l)dl-49.
*pld: pallid *pn26-20
location: 1-0. origin: X ray induced.
origin: Found in progeny of flies treated with Janus discoverer: Sobels, 57j.
green. references: 1958, DIS 32: 85.
discoverer: Muller, 28e20. phenotype: Eye color like pn. RK1.
synonym: pi. pn27.9
references: 1935, DIS 3: 30. origin: Induced by mustard gas.
phenotype: Body and wings pale. Viability about 10 discoverer: Sobels, 57j.
percent wild type. RK3. references: 1958, DIS 32: 84.
pleated: see pi phenotype: Like pn. RK1.
Plexate: see Px *pn27.22
plexus: see px origin: Induced by mustard gas.
Plum: see bwvi discoverer: Sobels, 57j.
*p/w: pale wing references: 1958, DIS 32: 84.
location: 1-37.2. phenotype: Like pn. RK1.
origin: Spontaneous, 5Ib
V
discoverer: Fahmy, 1952. origin: Induced by P 3 2 .
references: 1959, DIS 33: 88. discoverer: R. C. King, 51b.
phenotype: Body, wings, and bristles pale silvery references: 1952, DIS 26: 65.
yellow. Eclosion delayed; viability low. RK3. phenotype: Like pn. RK1.
Pm: see pnSTh8
Pra*: see origin: X ray induced.
PmD1: see bwA discoverer: W. K. Baker, 51 h8.
Pro*1; see Pu* references: 1956, DIS 30: 69.
pn: prune phenotype: Like pn. RK1.
location: 1-0.8. pn55
discoverer: Bridges, 16dl4. origin: Spontaneous.
phenotype: Eye color of newly emerged fly trans- discoverer: Kivett, 1955.
parent brownish red, darkening with age to references: Clancy, 1959, DIS 34: 48.
brownish purple. Lethal with K-pn. Eye color phenotype: Like pn. RK1.
autonomous in larval optic disks transplanted into pn 62
wild-type hosts (Beadle and Ephrussi, 1936, origin: X ray induced in YSX-YL, In(l)EN+dl-49,
Genetics 21: 230). Eyes contain 26 percent normal y v t.
red and 110 percent normal brown pigment (Nolte, discoverer: Petty, 62d,
1959, Heredity 13: 233-41). Larval Malpighian phenotype: Like pn. K-pn sensitive. RK1A.
tube color normal (Brehme and Demerec, 1942, H0Ac4
pn
Growth 6: 351-56). RK1. origin: X ray induced in Jn(i>cSJl^ c «K+d3-49.
cytology: Salivary chromosome locus placed at discoverer: Muller, Valencia, and Valencia, 1946-53.
2D5-6 by Deraerec and Sutton (Deroerec, Kaufmann, references: Valencia, 1966, DIS 41: 58.
Fano, Sutton, and Sansome, 1942, Carnegie Inst. cytology: Associated with
Wash. Year Book 41: 191) and by J. I. Valencia. Di(l)2C8-9;3Al-2 (J. I. Valencia).
po: pate ocelli origin: Spontaneous.

location: 2-65.2. discoverer: Piternick, 1944.
origin: Spontaneous. references: Goldschmidt, Hannah, and Piternick,
discoverer: Bridges, 38dl. 1951, Univ. Calif. (Berkeley) Publ. Zool. 55:
phenotype: Ocelli virtually colorless; some pigment 67-294.
bordering inner margins. Eye color slightly phenotype: Wings transformed into leglike struc-
brighter than wild type. RK2. tures. Penetrance of 15 percent in selected stocks
is increased by presence of YL. RK3.
origin: Spontaneous. poi: see svrPoi
discoverer: Bridges, 20jl3. Pointed wing: see Pw
synonym: do: dilute ocelli. Pointedaid: see BxJ
references: Morgan, Bridges, and Sturtevant, 1925, pot: see spePoi
Bibliog. Genet. 2: 224. poliert: see spaPo1
phenotype: Ocelli pale. RK3. polychaetoid: see pyd
pod foot: see pdf polychaetous: see pys
*pod-G: podopiera of Goldschmidt Polycomb: see Pc
location: Multifactorial. polymorph: see pym
origin: Spontaneous. polyphene: see pyp
discoverer Goldschmidt, 1943. polyphenic: see pph
references: 1945, Science 101: 389-90. *pop: popeye
1945, J. Morphol. 77: 71-103 (fig.). location: 1-0.4.
Goldschmidt, Hannah, and Piternick, 1951, Univ. origin: Induced by p-NN-di-(2-chloroethyl)amino-
Calif. (Berkeley) Publ. Zool. 55: 67-294. phenylbutyric acid (CB. 1348).
phenotype: Wing transformation into legs varies from discoverer: Fahmy, 1952.
almost wild type to three-jointed leglike append- references: 1958, DIS 32: 73.
ages. Penetrance of 1—2 percent was increased to phenotype: Eyes small, round, bulging, and rough.
2—4 percent by selection. Scalloped, blistered, Often some central ommatidia protrude. Small
and unexpanded wings and various abnormalities of body. Wings short, broad, and frequently blistered.
legs are pleiotropic effects. RK3. Male sterile; viability less than 10 percent wild
other information: Podoptera may be similar to type. RK3.
tetaltera effects. *port: port
*pod-H: podoptera of Hannah location: 3- (not located).
location: Multifactorial (principal factor on chromo- discoverer: Morgan, 14c.
some 2). references: Bridges and Morgan, 1923, Carnegie
origin: Spontaneous. List. Wash. Publ. No. 327: 125.
discoverer: Hannah, 1943. phenotype: Eye color slightly diluted. RK3.
references: Goldschmidt, Hannah, and Piternick, *port-b: port-b
1951, Univ. Calif. (Berkeley) Publ. Zool. 55: location: 3- (not located).
67-294. discoverer: Bridges, 19ill.
phenotype: Wings transformed into leglike append- references: Bridges and Morgan, 1923, Carnegie
ages. Legs characteristically changed, with parts List. Wash. Publ. No. 327: 214.
often duplicated. Average penetrance of 2.5 per- phenotype: Eye color maroon. RK3.
cent increases to 5 percent in selected lines. So- postbithorax: see pbx
matic elimination o(X chromosome produces more postverticalless: see pvt
than 2 percent gynandromorphs. RK3. *pph: polyphenic
other Information: Claimed to have a maternally in- location: 1-60.8 (originally located at 61.0 but ge-
herited component. netic location arbitrarily interchanged with that of
*pod-K: podoptera of Kellen-Pitemick sby for consistency with cytological observations).
location: Multifactorial. origin: Induced by D-l:6-dimethanesulfonyl mannitol
origin: Spontaneous, (CB. 2511).
discoverer. Kellen-Piternick, 1944. discoverer. Fahmy, 1959.
references: Goldschmidt, Hannah, and Piternick, synonyms pph-61: polyphene 61.
1951, Univ. Calif. (Berkeley) Publ. Zool. 55: references: 1964, DIS 39: 58.
67-294. phenotype: Body small. Eyes brighter than normal.
phenotype: Like pod-G. Wing sometimes replaced by Wing size and shape slightly altered. Scute liar
palpus like structure. Average penetrance 30 per- bristles occasionally kinked. Both sexes viable;
cent in X/X/Y females and X/Y males. Females fertility of homozygous female low. RK3.
without Y or YL do not show podoptera phenotype. cytology: Not included in deficiency for 18A4
Rough eyes, notched wings, and absence of post- through 18B8 produced by combining left end of
verticals occur. RK3. In(l)y4 **In(l)lA8-Bl;18A3-4 and right end of
*pod~M: podoptera in M(3)»-124 In(7>sc* **In(l)lB2-3;l8B8-9 (Norton and Valencia,
I ©cot ion: MultifactorlaL 1965, DIS 40: 40).
MUTATIONS 191
pr: purple
location: 2-54.5.
discoverer: Bridges, 12b20.
references: 1919, J. Exptl. Zool. 28: 264-305.
Bridges and Morgan, 1919, Carnegie Inst. Wash.
Publ. No. 278: 169 (fig.).
Sturtevant and Beadle, 1939, An Introduction to
Genetics, Saunders, p. 64 (fig.).
phenotype: Eye color ruby at hatching, darkening to
purplish ruby with age; orange in combination with
st, reddish brown in combination with bw (Mainx,
1938, Z. Induktive Abstammungs- Vererbungslehre Pr: Prickly
75: 256—76). Eye color autonomous in larval optic From Muller, 1930, J. Genet. 22: 299-334.
disks transplanted into wild-type hosts (Beadle Pr: Prickly
and Ephrussi, 1936, Genetics 21: 230). Larval location: 3-90.0.
Malpighian tubes normal (Beadle, 1937, Genetics origin: X ray induced.
22: 587-611). A lethal interaction of pr and ey discoverer: Muller, 27el7.
reported by Clemente (1941, Proc. Intern. Congr. references: 1930, J. Genet. 22: 299-334 (fig.).
Genet., 7th. p. 90) could not be confirmed by Green 1935, DIS 3: 30.
(1955, DJB 29: 121). RK1. phenotype: Bristles very short; tips thin and
cytology: Placed in salivary chromosome region twisted. Postdorsocentrals and scutellars usually
37B2 through 40B2, on the basis of its being missing; dark granule present beneath normal
within the deficiency from T(Y;2)H = T(Y;2)37B1- bristle location. Homozygote has low viability.
2;40B2-3. RK1.
2 P/-J-; Prickly-Long
V
discoverer: L. V. Morgan. origin: Spontaneous derivative of Pr.
references: Morgan, Bridges, and Sturtevant, 1925, discoverer: E. H. Grell, 65f.
Bibliog. Genet. 2: 233. phenotype: Bristles of Pr L /+ one-third as long as
phenotype: Eye redder than in pr. RK1. wild type; longer than Pr/+. Enhanced by H/+ so
*pr42d that it resembles Pr/+. Homozygote viable, with
origin: Spontaneous. small vestiges of bristles. RK1.
discoverer: Nolte, 42d. *pra: prawny abdomen
phenotype: Eye color somewhat more transparent origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
than pr and with a redder tone; less brown pigment phenylalanine (CB. 3007).
than pr (Nolte, 1955, J. Genet. 53: 1-10). RK1. discoverer: Fahmy, 1954.
prfew; purple-brown references: 1959, DIS 33: 88.
origin: Spontaneous. phenotype: Thorax narrow. Abdomen slender, often
discoverer: Bridges, 38d20. flexed between fourth and fifth segments. Wings
phenotype: Eye color brownish pink; lighter in fe- short, rather broad, and often held atypically.
male. RK2. Eciosion delayed. Viability about 15 percent wild
prIM60: purple-lethal of Meyer type. RK3.
origin: Spontaneous. prickle: see pk
discoverer: Meyer, 60g. Prickly: see Pr
references: 1963, DIS 37: 51. proboscipedia: see pb
phenotype: Homozygous lethal. pr1M60/pr has prong: see pg
purple eye. RK2. Protein 7: see Pt-1
other information: May be a small deficiency. prune: see pn
*prM60 Ps: Pigmentless
origin: X ray induced. location: 2-57.5 (inseparable from en),
discoverer: Meyer, 60f. origin: X ray induced.
references: 1963, DIS 37: 51. discoverer: Krivshenko, 56115.
phenotype: Eye color dark brown in p r ^ ^ / p r ; light references: 1959, DIS 33: 95.
apricot in pr^60 en homozygote at eciosion. RK1. phenotype: Black stripes on last abdominal seg-
*pr*; purple-sterile ments of female reduced; expression variable.
origin: Spontaneous. Male unaffected. Homozygous lethal. RK2.
discoverer: Ives, 38k. cytology: Salivary chromosomes apparently normal.
references: 1937, DIS 13: 50. pt: platinum
phenotype: Eye color weak pr. Eggs of pr'/pr* fe- location: 1-23.1.
male do not hatch; eggs of heterozygote and prm/pt origin: Deuteron induced.
female develop normally. Viability good. Male discoverer: Hildreth, 51h.
fertile. RK2. synonym: pa (preoccupied).
references: 1953, DIS 27: 56. Pt-8"

phenotype: Body color very pale yellow, almost location: 3- (not located).
colorless. Bristles colorless and translucent ex- origin: Naturally occurring allele.
cept for dark bases. Male sterile and short lived. discoverer: Duke.
Tyrosinase forms in adult (Horowitz and Fling). references: 1966, Genet. Res. 7: 287-94 (fig.).
RK2. phenotype: Homozygote apparently lacks detectable
pt: see abz amount of one of a number of protein fractions de-
Pt-V-Oh Protein 7 with mobility of 7.07 monstrable by starch gel electrophoresis of larval
location: 3- (10 crossovers with gl3 among 43 lymph (fraction 8). RK3.
tested). other information: May be the same locus as Pt-1
origin: Naturally occurring allele. described by Hubby (1963, Genetics 48: 871-79).
discoverer: Hubby.
references: 1963, Genetics 48: 871-79 (fig.). Pt-9"
phenotype: Protein 1 is one of about 10 bands found location: Autosomal.
after electrophoresis on acrylamide gel of the 40— origin: Naturally occurring allele.
50 percent ammonium sulfate cut of whole fly ho- discoverer: Duke.
mogenates. Protein 1 produced by Pt-11'01 has a references: 1966, Genet. Res. 7: 287-94 (fig.).
relative electrophoretic mobility of 1.01 under con- phenotype: Homozygote apparently lacks detectable
ditions used by Hubby (1963). RK3. amount of one of a number of protein fractions de-
other information: May be the same locus as Pt-8 monstrable by starch gel electrophoresis of larval
described by Duke (1966, Genet. Res. 7: 287-94). lymph (fraction 9). RK3.
Pf-71.13 Pt-13"
origin: Naturally occurring allele. location: Autos omal.
discoverer: Hubby. origin: Naturally occurring allele.
references: 1963, Genetics 48: 871—79 (fig.). discoverer: Duke.
phenotype: Pt-11'13/Pt-11'13 produces protein 1 references: 1966, Genet. Res. 7: 287-94 (fig.).
with electrophoretic mobility 1.13. Pt-11-01/ phenotype: Homozygote apparently lacks detectable
Pt-11'13 produces both protein types but none with amount of one of a number of protein fractions de-
intermediate mobility. RK3. monstrable by starch gel electrophoresis of larval
Pt-4": Protein ^negative lymph (fraction 13). RK3.
location: 1- (not located). pta: see sldPta
origin: Naturally occurring allele. Ptd: see BxJ
discoverer: Pantelouris and Duke. *pte: pterygion
references: 1963, Genet. Res. 4: 441-45 (fig.). location: 1-1.4.
phenotype: Homozygote apparently lacks detectable origin: Induced by l:4-dimethanesulfonoxybut-2-yne
amount of one of a number of protein fractions de- (CB. 2058).
monstrable by starch gel electrophoresis of larval discoverer: Fahmy, 1951.
lymph [fraction A according to Pantelouris and references: 1958, DIS 32: 73.
Duke (1963); fraction 4 according to Duke (1966, phenotype: Wings shortened, usually spread, and
Genet. Res. 7: 287-94)]. RK3. slightly drooping. Eyes misshapen and somewhat
Pt-5* rough. Abdomen disproportionately large. Eclosion
location: 2- (not located). slightly delayed and viability about 20 percent
origin: Naturally occurring allele. wild type. RK3.
discoverer: Pantelouris and Duke. ptg: pentagon
references: 1963, Genet. Res. 4: 441-45 (fig.)- location: 1-23.2.
phenotype: Homozygote apparently lacks detectable discoverer: Bridges, 2218.
amount of one of a number of protein fractions de- phenotype: Thoracic trident darker than wild type,
monstrable by starch gel electrophoresis of larval especially the pentagonal spot just ahead of scu-
lymph [fraction B of Pantelouris and Duke (1963); telium; more extreme at 19°C. Hard to classify in
fraction 5 of Duke (1966, Genet. Res. 7: 287-94)]. young flies. RK3.
RK3. cytology: Located in salivary chromosome region
Pt-Spn: Protein 5 prime-negative 7C4-8C2 (Demerec, Kaufmann, Fano, Sutton, and
location: 2- (not located). Sansome, 1942, Carnegie Inst. Wash. Year Book
origin: Naturally occurring allele. 41: 191). Further restricted to 7E1 through 8C2,
discoverer: Pantelouris and Duke. on the basis of its exclusion from Dt(l)sn -
references: 1963, Genet. Res. 4: 441—45 (fig.). Di(l)7B2-3;7D22-El (Hinton and Welshons, 1955,
phenotype: Homozygote apparently lacks detectable DIS 29: 125-26).
amount of one of a number of protein fractions de- ptg2
monstrable by starch gel electrophoresis of larval discoverer: L. V. Morgan, 24j21.
lymph [fraction C of Pantelouris and Duke (1963); references: 1935, DIS 3: 14.
fraction 5'of Duke (1966, Genet. Res. 7: 287-94)]. phenotype: Pentagonal spot darker and sharper than
RID. in ptg. Scute Hum often dark and prongs of trident
MUTATIONS 193
sometimes so. Best classification at lower temper- phenotype: Eye color rosy purple. With st, eyes
atures. RK2 at 19°C. show patchwork of light to deep orange areas.
ptg3 Homozygous lethal. RK2A.
discoverer: Kaliss, 351. cytology: Associated with T(2;3)PUGT ~
synonym: cro; crown. TX2;3)57C;81F (Lewis, 1956, DIS 30: 130).
references: 1937, DIS 7: 6, 18. PuK: Punch of Krivshenko
Felsenstein, 1937, DIS 7: 21. origin: X ray induced.
phenotype: Trident darker than in ptg; dark color ex- discoverer: Krivshenko, 53k24.
tends to head, sides, and abdomen. RK2. synonym: Pni^-.
other information: Occasionally reverts to wild type references: 1954, DIS 28: 75.
or weak ptg. Allelism with ptg shown by Bridges. Rowan, 1966, DIS 41: 166-67.
ptg4 phenotype: Like PuGr. RK1A.
origin: Spontaneous in In(l)AM. cytology: Associated with In(2R)PuK =
discoverer: Curry, 38b8. In(2R)41;S7E-F.
phenotype: Darkness of pentagon intermediate be- *Pu™: Punch-reversed
tween that of ptg and ptg2. RK2A. origin: X-ray-induced derivative of Pu.
pu: pupal discoverer: Oliver, 32127.
location: 2-51. references: 1941, Proc. Intern. Congr. Genet., 7th.
discoverer: Duncan, 20d. p. 228.
synonym: pads-b. phenotype: Eye color appears wild type at 25°C;
references: Morgan, Bridges, and Sturtevant, 1925, frequently homogeneous brownish shade in young
Bibliog. Genet. 2: 232. flies at 16°. Pu/PuTV flies viable but sterile; eye
phenotype: Wings unexpended or incompletely ex- color like Pu/+'f often wings are opaque, bristles
panded. More extreme at 19°C. RK2. thin, trident dark, eyes rough, dark, and sometimes
cytology: Placed in region between 34E5 and 35D1 variegated. Homozygous lethal. RK3A.
on basis of its inclusion in Df(2L)64j = cytology: Reportedly associated with T(2;3)Purv =
Df(2L)35E5-Fl;35C3-Dl (E. H. Grell). T(2;3)S7B5-C1;79F superimposed on T(2;3)Pu -
other information: Not a lie lie to pada. T(2;3)40F-41Al;70D-E + T(2;3)57B5-C1;79F,
*Pu: Punch P« w : Punchline
location: 2-97 (location of Pu2). origin: X ray induced.
origin: X ray induced. discoverer: E. B. Lewis, 55h.
discoverer: Oliver, 28k4. phenotype: Like Pu with variegated appearance.
references: Muller, 1930, J. Genet. 22: 326 (fig.). Homozygous lethal. RK2A.
Oliver, 1932, Z. Induktive Abstammungs- cytology: Associated with T(2;3)Puw = T(2;3)S7B-
Vererbungslehre 61: 484. C;80.
1935, DIS 3: 14. pub: pubescent
phenotype: Eye color dilute purple. Gives normal location: 1-63.
eye color when heterozygous with T(2;4)A34 = origin: Induced by P 3 2 .
T(2;4)56F6-7 (Oliver, 1943, Anat. Record 87: 461). discoverer: Bateman, 1950.
Homozygous lethal. RK2A. references: 1950, DIS 24: 55.
cytology: Associated with T(2;3)Pu = T(2;3)40F- phenotype: Hairs and bristles M-like; black pigment
41A;70D-E + T(2;3)57B5-C1;79F. Tentatively on terminal abdominal segments nearly absent;
placed in region 57B-C, on basis of breakpoint male sterile. Tendency toward short, fat, gnarled
common to T(2;3)Pu, T(2;3)PuOr - T(2;3)57C;81F, legs, shortened L2, and posterior nicking of wings.
and T(2;3)Puw = T(2;3)57B-C;80. After several generations, only bristle effect and
male sterility remained. RK3.
origin: Spontaneous. *Pub: Pub
discoverer: E. H. Grell, 57b. location: 1- (rearrangement),
references: 1960, DIS 34: 50. discoverer. P. Famsworth.
phenotype: Heterozygote has purplish eye color re- references: Lefevre, 1954, DIS 28: 75.
sembling pr. Homozygous lethal. Pu2/Pu, phenotype: Eye size of heterozygote variably re-
pu2/puOrt a n d Pu2/puW a l s o lethal. RK1. duced, ranging from something like Bty+ to wild
cytology: Apparently not associated with a chromo- type. Eyes of homozygote greatly reduced, similar
somal rearrangement. to double Bar. Interacts with B to give small,
glazed, almost facetless eyes. RK2A.
.. Punch-Grape cytology: Associated with In(l)Pub; breakpoints un-
origin: X ray induced, known.
discoverer: Muller, 291. pubescent: see pub
synonym: pOr: pink-Grape. put: puff
references: Glass, 1933, J. Genet. 28: 69—112 location: 2-58.
(fig.). origin: Spontaneous.
1934, Am. Naturalist 68: 111. discoverer: Nichols-Skoog, 35kl9.
phenotype: Wings puffed or blistered, effect center-

ing in third posterior cell; wings warped and
creased longitudinally along vein L3. Penetrance
usually 90—100 percent in female and 20—40 per-
cent in male. RK3.
Pufdh s e e Pfd
puff: see puf
pun: puny
location: 1-41.1.
origin: Induced by triethyleriemelamine (CB. 1246).
phenotype: Body small. Wings slightly shorter than
normal. Eyes occasionally deformed. Eclosion
delayed. Both sexes fertile; viability about 50 per-
other information: One allele each induced by CB.
1356 and CB. 3025.
Punch: see Pu
puny: see pun
pupal: see pu
purple: see pr
purpleoid: see pd
Purpleoider. see Pdr Pw: Pointed wing
*pvf: postverticalless Publ. No. 327: 239.
location: 1-20.9.
origin: Induced by ethyl methanesulfonate (CB. px: plexus
1528). location: 2-100.5.
discoverer: Fahmy, 1956. discoverer: Bridges, 14h20.
references: 1959, DIS 33: 88. references: Bridges and Morgan, 1919, Carnegie
phenotype: Wings either divergent or slightly held Inst. Wash. Publ. No. 278: 251 (fig.).
up. Thoracic hairs sparse, and one or both post- Morgan, Bridges, and Sturtevant, 1925, Bibliog.
vertical bristles almost invariably absent. Shape Genet. 2: 212 (fig.), 233.
of head and eyes varies from almost normal to phenotype: Wings have network of extra veins,
anteroposterior flattening of head and deep groov- especially toward tips and margins; LA bent near
ing of eyes. Male viable and fertile; female tip. Semidominant with some Minutes. Suppressed
sterile. RK2. by S (Bedichek, 1936, DIS 5: 24). Venation effect
*pw: pink wing caused by inadequate contraction of wing during
location: 2-14. pupal stage, leaving spaces between epithelial
discoverer: Bridges, 2Obi7. layers (Waddington, 1940, J. Genet. 41: 75-139).
references: Morgan, Bridges, and Sturtevant, 1925, RK1.
Bibliog. Genet. 2: 213. cytology: Placed in 58F, on basis of its inclusion
1931, Eos 7: 229^»8. in Dl(2R)M-l m Df(2R)57Fll-58Al;5SF8'59Al and
phenotype: Eye color like pink. Wings shorter than Dp(2;3)P from T(2;3)P = T(2;3)58E3-F2;60D14-
normal and crumpled. Viability low. RK3. E2;96BS-C1 [Bridges, 1937, Cytologia (Tokyo),
Fuji! Jub. Vol. 2: 745-55],
*Pw: Pointed wing
location: 3-94.1.
Inst. Wash. Publ. No. 327: 238 (fig.),
phenotype: Wings narrowed slightly at tips; extra
venation near tips of L3 and L4. Homozygous
lethal. RK3.
other information: Not an allele of Bd (3-93.8).
pw-c: pink wing-c px; plexus

location: 2-79. Edith M. Wallace, unpublished.
discoverer: Bridges, 31cl8. px*
phenotype: Eye color lighter than normal. Wings origin: Spontaneous.
short and blunt. Overlaps wild type. RK3, discoverer: Villee, 40a.
MUTATIONS 195
references: 1942, Univ. Calif. (Berkeley) Publ. phenotype: Like Px. Px2/Px is lethal. Homozygote
Zool. 49: 125-84. dies as embryo. RK1A.
phenotype: Like px. RK1. cytology: Associated with Df(2R)Px2 =
px3id: see bs^ Df(2R)60C5-6;60D9-10 (Bridges, 1937).
pXS29 Px*
origin: X ray induced. origin: Synthetic.
discoverer: Iyengar, 52g. discoverer: Thompson, 56f.
references: Iyengar and Meyer, 1956, DIS 30: 73. references: Burdick, 1956, DIS 30: 69.
phenotype: Like px. RK1A. phenotype: Wing venation like Px. Thickening of L5
cytology: Induced simultaneously with at posterior crossvein produces a vesicle, as in
In(2LR)pxS2&\ breakpoints not determined. 6s. More extreme in female. Homozygous lethal.
px54h RK1A.
origin: Spontaneous. cytology: Associated with In(2LR)Px4 -
discoverer: Meyer, 54h. In(2LR)22A3-Bl;60B-CL;21 C8-Dl;60Dl-2R derived
references: 1954, DIS 28: 77. from single recombinant between ln(2LR)bwvl and
phenotype: Like px. RK1. SMI, which is deficient for 60B-60D1.
pxSSk *PxS
origin: Spontaneous. origin: Spontaneous in In(2LR)bwvl.
discoverer: Williams, 55k. discoverer: Thompson, 1957.
phenotype: Like px. RK1. phenotype: Sacs or vesicles in wing but little irreg-
bs
px : see ularity of venation. Lethal in homozygote and in
heterozygote with other Px alleles. RK1A.
other information: 6s and 6a affected but not sp.
pyd: polychaefoid
location: 3-39.
discoverer: Spencer, 39h31.
synonym: Pch.
1937, DIS 7: 15.
Neel, 1939, Genetics 24r 81.
Px: Plexate 1941, Genetics 26: 52-68.
Edith M. Wallace, unpublished. 1943, Genetics 28: 49-68.
phenotype: Extra bristles present in homozygote at
Px; Plexate or near almost all normal bristle locations but most
location: 2-107.2 (107.0-107.4 inclusive). frequently in dorsocentral and scutellar regions.
origin: Spontaneous. Heterozygote in some stocks occasionally shows
discoverer: Bridges, 22f6. extra bristles, especially vibrissae. Character ex-
references: 1937, Cytologia (Tokyo), Fujii Jub. pressed better at low temperatures and in large
Vol. 2: 745-55. flies. Combinations with h and Hw generally
phenotype: Wing veins of heterozygote have plexus- superadditive for bristle number. RK3.
like or deltalike thickenings, most often near pos-
terior cross vein, and free fragments of veins, most *pym: polymorph
often in third posterior cell; L4 bent near margin. location: 2- (not located).
Wings smaller and narrower than wild type and origin: Spontaneous.
dusky textured. Closely resembles 6s. Expression discoverer: Bryson, 1939.
more extreme in female and enhanced by cold references: 1940, DIS 13: 49.
(19°C). Homozygote lethal in egg stage (Li, 1927, phenotype: Eye color translucent dull ruby. Wings
Genetics 12: 1-58). RK1A. small; may be absent. Bristles slightly Minute.
cytology: Associated with Dt(2R)Px - Df(2R)60B8- Posterior crossvein often missing. Both sexes
10;60Dl-2. Locus placed in salivary chromosome sterile. Viability low, especially in female. RK3.
region 60C6 through 60D1, on basis of the region *pym2
of overlap of Di(2R)Px and Df(2R)Px^ « origin: Spontaneous.
Dg2R)60CS-6}60D9'10 (Bridges, 1937). discoverer: Neel, 1941.
other information: May be part of a pseudoallelic references: 1942, Am. Naturalist 76: 630-34.
complex with 6a and 6s. phenotype: Eyes ruby; ocelli pale. Body small and
abnormally shaped. Wings small, thin, and wavy;
origin: X ray induced. second crossvein often interrupted or missing;
discoverer: Schultz, 3211. plexus often present near wing tip or in third pos-
references: Bridges, 1937, Cytologia (Tokyo), Fujii terior cell; marginal hairs irregular, shallow inci-
Jub. Vol. 2: 745-55. sions present in posterior margin. Bristles slender
and either increased or decreased in number. Ex- references: 1915, Am. Naturalist 49: 240—50.
ternal genitalia normal in both sexes, but internal Morgan and Bridges, 1916, Carnegie Inst. Wash.
genitalia abnormal. Viability about 80 percent Publ. No. 237: 25 (fig.).
normal. RK3. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
Genet. 2: 24, 56-57, 234 (fig.).
*pyp: polyphene phenotype: Wings obliquely truncated to about tip of
location: 1-53.5. abdomen; marginal hairs sparse and ruffled; veins
origin: Spontaneous. L4 and L5 generally shortened. Wings usually arc-
discoverer: Bridges, 37126. like and often blistered. Viability irregular. Fe-
phjenotype: Wings spread, yellowish, and with male usually sterile in cross with r male but occa-
uneven surface. Trace of extra vein in third pos- sionally gives a few offspring (mostly daughters)
terior cell, near second crossvein. Eyes rough, in outcross; for morphology of female sterility see
pitted, bulging, and smaller than wild type. Trident t3$. RK2.
more darkly pigmented in male. Female sterile. other information: The r locus was subdivided into
Viability about 70 percent wild type. RK3. six complementation groups by Fahmy and Fahmy
pys: polychaetous (1959, Nature 184: 1927-29) and into at least
location: 2-52. three groups, on the basis of both complementation
discoverer Curry, 37kl5. and recombination analysis, by Green (1963,
phenotype: Extra or double bristles present; most Genetica 34: 242-53). Unfortunately, the two
easily seen are scutellars, dorsocentrals, orbitals, analyses were performed on different groups of
and vibrissae. Extra bristles on scutellum curve mutants, and the maps cannot be correlated. Com-
upward. Overlaps wild type at 19°C, but classifi- plementation varies from partial to complete and
cation good at 28-30°. RK3. the degree may be related to distance apart on the
complementation map.
*Q: Queer wing V
location: 2- (not located). discoverer: Bridges, 14g.
discoverer: E. M. Wallace, 1931. references: 1916, Genetics 1: 151.
phenotype: Wings irregularly incised; marginal phenotype: Like r. RK2.
bristles irregular. Heterozygote has low pene- *r2 L: rudimentary of Lancefield
trance; homozygote better. RK3. discoverer: Lancefield.
Qd: Quadroon references: 1918, Am. Naturalist 52: 264-69.
location: 1-6.8. phenotype: Like r. RK2.
origin: Spontaneous. *f3
discoverer: Thompson, 58k. discoverer: Sturtevant, 17J30.
references: 1959, DIS 33: 99. phenotype: Like r except that about one-third of fe-
phenotype: Broad dark band on margins of all ab- males are fertile (Lynch, 1919, Genetics 4: 501—
dominal tergites, giving abdomen superficial 33). RK2.
appearance of uniform darkness. Viability of heter- r?: see r»
ozygous female normal, of homozygous female 40
percent normal, and of male 30 percent normal.
RK2.
Queer wing: see Q
X/2
X10
X9
X8
X7
X13 X6
X5 X3
X2 XI
sS r9: rudimentary~9
$8 Edith M. Wallace, unpublished.
s6 s3
X/4 r*
39k s2
discoverer: Bridges, 20b3.
Map of r locus synonym: r7.
From Green, 1963, Genetfca 34: 242-53. references: Morgan, Bridges, and Sturtevant, 1925,
*r: rudimentary phenotype: Wings truncated. Veins sometimes in-
location: 1-54.5. complete; slight deltas at crossveins; marginal
discoverer: Morgan, lOf. hairs uneven. Female usually fertile. RK1.
MUTATIONS 197
rU discoverer: Fahmy, 531.

origin: Spontaneous. references: 1959, Nature 184: 1927-29.
discoverer: E. M. Wallace, 22k8. phenotype: Typical r. RK2.
references: Morgan, Bridges, and Sturtevant, 1925, other information: Occupies complementation group
Bibliog. Genet. 2: 234. III of the Fahmys.
phenotype: Much less extreme than r and somewhat ,54 c
less so than r9; female more fertile. Overlaps wild origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
type in female. RK3. phenylalanine (CB. 3026).
discoverer: Fahmy, 54c.
discoverer. Bridges, 24d4. references: 1959, Nature 184: 1927-29.
references: Morgan, Bridges, and Sturtevant, 1925, phenotype: Typical r. RK2.
Bibliog. Genet. 2: 234. other information: Occupies complementation group I
phenotype: Formerly more extreme than r; later less of the Fahmys.
extreme. RK2. r54d
*r35 origin: Induced by L-p-NN-di-(2-chloroethyI)amino-
origin: Spontaneous. phenylalanine (CB. 3025).
discoverer: Gottschewski, 1935. discoverer: Fahmy, 54d.
phenotype: Strong allele of r. RK2. references: 1959, Nature 184: 1927-29.
*r3Sa phenotype: Typical r. RK2.
origin: X ray induced. other information: Occupies complementation groups
discoverer Oliver, 35alO. I and II of the Fahmys. Complementation group II
references: 1939, DIS 12: 48. inferred from its interaction with rSSa.
phenotype: Like r, but less viable. RK3. rS4f
* r 39 origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
origin: Induced by mustard gas. phenylalanine (CB. 3007).
discoverer: Auerbach, 1951. discoverer: Fahmy, 54j.
references: Counce, 1956, Z. Induktive references: 1959, Nature 184: 1927-29.
Abstammungs- Vererbungslehre 87: 482—92. phenotype: Typical r. RK2.
phenotype: Wings like r. Homozygous female sterile other information: A noncomplementing allele. Four-
in cross to r39 male; fertile in out cross, producing teen such alleles found among 31 tested by the
mostly daughters (1—3 percent of progeny are Fahmys.
male). Homozygous female produces many mal- rSSa
formed eggs and unfertilized eggs with normal mor- origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
phology. Ovarian development often retarded or phenylalanine (CB. 3025).
fails. Yolk deposition affected. Lethal effect in discoverer. Fahmy, 55a.
progeny results from generalized disturbance in references: 1959, Nature 184: 1927-29.
differentiation 13-16 hr after fertilization at 25°C. phenotype: Typical r. RK2.
Surviving embryos hatch late and may produce other information: Occupies complementation groups
larvae that neither move nor feed. RK2. II and m of the Fahmys.
r3 9k rSSk
origin: Recovered among progeny of cold-treated fe- origin: Induced by p-NN-di-(2-chloroethyI)amino-

male. phenylethylamine (CB. 3034).
discoverer. L. V. Morgan, 39k9. discoverer: Fahmy, 55k.
synonym: r*1 (Green, 1963, Genetica 34: 242—53). references: 1959, Nature 184: 1927-29.
references: 1940, DIS 13: 51. phenotype: Typical r. RK2.
phenotype: Wing short and crumpled; legs weak. other information: Occupies complementation group
Homozygous female sterile, but r/r 3 5 * female par- V of the Fahmys. Seven such alleles among 31
tially fertile. RK2. tested by the Fahmys.
other information: Complements completely with r*2, rS6d
partially with i*7', r**, rX9, rx^0i a n d jX14 a n d origin: Induced by ethyl methanesulfonate (CB.
slightly with r**. Genetically to the left of r* 2 , 1528).
rX7t TX8f rX9t rXl0> a n d ,JU4 (Green, 1963). discoverer: Fahmy, 56d.
v» references: 1959, Nature 184: 1927-29.
origin: Induced by mustard gas. phenotype: Typical r. RK2.
discoverer: Auerbach, 1951. other information: Occupies complementation groups
references: Counce, 1956, Z. Induktive IV and V of the Fahmys.
rS6j
phenotype: Like r39, but slightly more extreme. origin: Induced by methyl methanesulfonate (CB.
RK2. 1540).
rS3l discoverer: Fahmy, 56j.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- references: 1959, Nature 184: 1927-29.
phenyialanine (CB. 3025). phenotype: Typical r. RK2.
other information: Occupies complementation groups *r*4

I through IV of the Fahmys. origin: Spontaneous.
rS6k discoverer: Green, 59b.
origin: Induced by methyl methanesulfonate (CB. references: 1963, Genetica 34: 242-53.
1540). phenotype: Like r. RK2.
discoverer: Fahmy, 56k. other information: A noneomplementing allele.
references: 1959, Nature 184: 1927-29. *rs5
phenotype: Typical r. RK2. origin: Spontaneous,
other information: Occupies complementation group discoverer: Green, 60i7.
VI of the Fahmys. Four such alleles among 31 references: 1963, Genetica 34: 242-53.
tested by the Fahmys. phenotype: Like r. RK2.
r 58o other information: A noncomplementing allele lo-
origin: X ray induced. cated to the right of r39k by recombination.
discoverer: M. Burdick, 1958. *rs6
references: A. B. Burdick, 1961, DIS 35: 45. origin: Spontaneous.
phenotype: Like r and t9. Females sterile. RK2. discoverer: Green, 60112.
r63c references: 1963, Genetica 34: 242-53.
origin: Spontaneous. phenotype: Like r. RK2.
discoverer: Clancy, 63c. other information: A noncomplementing allele lo-
references: 1964, DIS 39: 65. cated to the left of r*2 by recombination.
phenotype: Like r. Female conditionally sterile. rs7
other information: Does not complement with i39k Of discoverer: Green, 60112.
T*. references: 1963, Genetica 34: 242—53.
*rG: rudimentary of Goldschmidt phenotype: Like r. RK2.
origin: Spontaneous. other information: A noncomplementing allele.
discoverer: Goldschmidt. *r,8
synonym: rPx *>•*, origin: Spontaneous.
references: 1945, Univ. Calif. (Berkeley) Publ. discoverer: Green, 61g2.
Zool. 49: 501-3. references: 1963, Genetica 34: 242—53.
phenotype: Like r. RK2. phenotype: Like r. RK2.
*rH other information: Complements like r39k. Genet-
origin: Induced by formaldehyde. ically to the right of r*-* 0.
discoverer: Auerbach, 1951. *rs: rudimentary-Swiss
references: Counce, 1956, Z. Induktive origin: Spontaneous.
Abstammungs- Vererbungslehre 87: 482—92. discoverer: Hadorn, 59d.
phenotype: Like r39. RK2. references: Rohr, 1962, DIS 36: 39.
phenotype: Like r. Wings smaller in homozygous fe-
*rK: rudimentary of Krivshenko male than in male. Duplicated bristles often
discoverer: Krivshenko. arising from same socket. Lateral marginal wing
references: Agol, 1936, DIS 5: 7. bristles of r^r 1 2 , but not rs/t39k or r S/ r 9 ( o c c a .
phenotype: Wings like r. Semilethal; female sterile. sionally normal. Female sterile. RK2.
RK2A. *rSn; rudimentary of Silberman
cytology: Associated with/nfl^r^; proximal break in discoverer: Silberman.
chromocenter. references: Agol, 1936, DIS 5: 7.
rPx bl: see rG phenotype: Like r. RK2.
rsl: see r^9k r* 1: rudimentary from X irradiation
r*2; rudimentary-spontaneous origin: X ray induced.
origin: Spontaneous. discoverer: Green, 60bl3.
discoverer: Green, 59k22. references: 1963, Genetics 34: 242—53.
references: 1963, Genetica 34: 242-53. phenotype: Like r. RK2.
phenotype: Like r. RK2. other information: A noncomplementing allele lo-
other information: Complements with r35*, r"8, rxs, cated to the right of r39k by recombination.
, an< j r^*. Genetically to the right of r39*, * r X2
discoverer: Green, 60cl5.
origin: Spontaneous, references: 1963, Genetica 34: 242-53.
discoverer Green, 58b. phenotype: Like r. RK2.
references: 1963, Genetica 34: 242—53. other information: A noncomplementing allele lo-
phenotype: Like r. RK2. cated to the left of rs? by recombination.
other information: Complements slightly with r X3
Genetically to the right of origin: X ray induced.
MUTATIONS 199
discoverer: Green, 60c 15. *rX12

references: 1963, Genetica 34: 242—53. origin: X ray induced.
phenotype: Like r. RK2. discoverer: Green, 60k27.
other information: Complements slightly with r39k, references: 1963, Genetica 34: 242-53.
Ta8i TX5f a r u j rX13m Genetically to the right of phenotype: Like r. RK2.
other information: A noncomplementing allele lo-
*rX4 cated to the right of r39k by recombination.
origin: X ray induced. *rX13
discoverer: Green, 60cl5. origin: X ray induced.
references: 1963, Gfenetica 34: 242—53. discoverer: Green, 60k27.
phenotype: Like r. RK2. references: 1963, Genetica 34: 242-53.
other information: A none complementing allele. phenotype: Like r. RK2.
*rX5 other information: Complements completely with r s 2 ,
origin: X ray induced. r*«, rX9, and r*-*<\ partially with t*l*, and
discoverer: Green, 60cl5. slightly with r^3. Genetically to the left of t*2.
references: 1963, Genetica 34: 242—53. r X14
phenotype: Like r. RK2. origin: X ray induced.
other information: Complementation behavior like discoverer: Green, 62j7.
r39k. Genetically to the left of r*2 and rÔ. references: 1963, Genetica 34: 242-53.
r X6 phenotype: Like r. RK2.
origin: X ray induced. other information: Complements completely with ra^,
discoverer: Green, 60dl. partially with r39k, r««, r*-*, r*«, T*9, &*<>, and
references: 1963, Genetica 34: 342—53. r*13, and slightly with t*3 and raS. Genetically
phenotype: Like r. RK2. between r39k and ra2.
other information: A noncomplementing allele lo- R: Roughened
cated to the right of r39k by recombination. location: 3-1.4.
* r X7 discoverer E. M. Wallace, 35i.
origin: X ray induced, phenotype: Eyes of R/+ rough with some large dark
discoverer: Green, 60e24. facets. Male genitalia frequently rotated and male
references: 1963, Genetica 34: 242—53. sometimes sterile; viability about 80 percent wild
phenotype: Like r. RK2. type. Homozygote se mile thai; wings spread.
other information: Shows partial complementation Thorax short; acrostichal hairs deranged, some
with r 35 *, ra8, and rX5. Genetically to the right of missing; eyes small. Homozygous female fertile.
RK1.
*rX8 R3(+): see T(2;3;4)+3
origin: X ray induced, *RS1b
discoverer: Green, 58a. origin: Recovered among progeny of female treated
references: 1963, Genetica 34: 242—53. as embryo with cold shock.
phenotype: Like r. RK2. discoverer: Mickey, 51b21.
other information: Complements completely with references: 1951, DIS 25: 74.
rX13, partially with r***, r««, r**, and r**'. Ge- 1951, Genetics 36: 565-66.
netically to the right of r5**. phenotype: Eyes of heterozygote small, oblong, and
*rX9 rough; facets and eye hairs irregular. Viability
origin: X ray induced. good. Homozygote lethal. Rsib/R has very small
discoverer: Gloor, 57a. eyes; much fusion of facets; resembles gl and Gl.
references: Green, 1963, Genetica 34: 242—53. RK1.
phenotype: Like r. RK2. ra: rase
other information: Complementation pattern like r^®. location: 3-97.3.
Genetically to the right of t39k. origin: Spontaneous.
*rX10 discoverer: Beadle, 34d.
discoverer: Gloor, 57a. phenotype: Bristles and hairs small; irregularly
references: Green, 1963, Genetica 34: 242—53. absent, especially from head and thorax. Viability
phenotype: Like r. RK2. good; developmental time normal. RK2.
other information: Complementation pattern like rxs. *ra2
Genetically to the right of r*9*. o-ri gin: Spontane ou* in In(3R)P.
*rXll discoverer: Mossige, 36k21.
origin: X ray induced. synonym: bd; bald.
discoverer: Green, 60k27. references: 1937, DIS S: 9.
references: 1963, Genetica 34: 242—53. phenotype: Homozygote lacks all head bristles and
phenotype: Like r. RK2. some scutellars* Heterozygote has extra anterior
other information: A noncomplementing allele. scute liars in about 30 percent of flies. RK2A.
cytology: Occurred in and probably inseparable from phenotype: Eye color of male maroon, light and
In(3RyP = In(3R)89C2'4;96A18-19. translucent immediately after eclosion but becomes
*rab: rabbit nearly wild type after 1 day. Female wild type.
origin: Induced by P 3 2 . ras4
discoverer Bateman, 1950. origin: Spontaneous.
references: 1950, DIS 24: 55. discoverer: Ives, 38f.
phenotype: Hairs on mesonotum near dorsocentral phenotype: Like ras, but female sterile. RK2.
bristles turned inward toward midline. Air bubbles rasy: raspberry-variegated
occasionally in thorax, beneath dorsocentrals, and origin: Fast neutron induced.
scutellum. Wiags rarely held up. Viability and fer- discoverer: E. B. Lewis, 1953.
tility normal. RK2(A). references: Brokaw, 1954, DIS 28: 73.
other information: Slight disturbance of crossing phenotype: Variegates for ras. Homozygous viable.
over proximally. RK2A.
radius incompletus: see ri cytology: Associated with T(l;3)ras^ =
T(1;3)9E;13C;81F (Lewis).
*rag: ragged ras4: see ra
location: 3-37 (Steinberg). raspberry: see ras
discoverer: Charles, 1932. rauhig: see gl3
references: Dunn, 1934, DIS 1: 30. raven: see rv
phenotype: Hairs missing from sections of wing rb: ruby
margin. RK3. location: 1-7.5.
*rai: raisin discoverer: Bridges, 14;18.
location: 3-17 (Stanley). phenotype: Eye color clear ruby, white in combina-
origin: Spontaneous. tion with wa, orange with st, and brownish red with
discoverer Hersh. bw (Mainx, 1938, Z. Induktive Abstammungs-
references: 1953, DIS 27: 55. Vererbungslehre 75: 256—76). Development of pig-
phenotype: Eye color deep brown, like se. Eclosion ment autonomous in rb eye disks transplanted into
delayed 1 or 2 days. RK2. wild-type hosts (Beadle and Ephrussi, 1936,
raised: see rsd Genetics 21: 230). Larval Malpighian tubes pale
raised wing: see rw yellow (Beadle, 1937, Genetics 22: 587-611).
raisin: see rai RKI.
cytology: Salivary chromosome location between
ras: raspberry 4C8 and 4D1 (Demerec, Kaufmann, Fano, Sutton,
location: 1-32.8. and Sansome, 1942, Carnegie Inst. Wash. Year
origin: Recovered among progeny of heat-treated Book 41: 191; J. I. Valencia). Located at 4C7-8 or
flies. possibly 4C6 by Hannah.
discoverer: Muller, 28dl7.
phenotype: Eye color dark ruby; 25 percent normal discoverer Fox, 48a7.
red pigment, 114 percent normal brown pigment references: 1948, DIS 22: 53.
(Nolte, 1959, Heredity 13: 233-41). Color autono- 1949, Genetics 34: 647-64.
mous in larval optic disks transplanted into wild- phenotype: Like rb. Fly lacks an antigen produced
type hosts (Beadle and Ephrussi, 1936, Genetics by wild type; the same antigen removed by v48*.
21: 230). Larval Malpighian tubes nearly wild rfr**a fly has no antigen not shared with wild type
type, not useful for classification (Brehme and or v*«*. RKI.
Demerec, 1942, Growth 6: 351-56). RK1. cytology: Salivary chromosomes normal.
cytology: Placed in 9E-F, on the basis of its being
included in the section of the X translocated into rbm48aH5: ruby-mottled
the base of 3R by 7X1,-3)1*8* « T(1;3)9E;13Q;81F origin: X ray induced in ln(l)&cslL8c8R+dl-49.
and its genetic position to the left of v in 10A1-2. discoverer: Muller, Valencia, and Valencia, 1946-53.
ras* references: Valencia, 1966, DIS 41: 58.
discoverer: Grossman, 1932. cytology: Associated with In(l)rb°"*8aH5 x*in(l)3E3-
references: Dtinn, 1934, DIS 1: 30. 4;11A7-8;2OF. Euchromatic section of X inserted
phenotype: Eye color translucent ruby, lighter than into JO? in reverse order,
ras; darkens less with age. Eyes contain 15 per- rt>RlSBH3
cent normal red pigment and 103 percent normal origin: X ray induced in R(l)2.
brown pigment (Nolte, 1959, Heredity 13: 233—41). discoverer: Muller, Valencia, and Valencia, 1946-53.
RKI. references: Valencia, 1966, DIS 41: 58.
ras3 cytology: Associated with
origin: Spontaneous. Df(l)4B4-5;4D5-6 O- I. Valencia).
discoverer: Iveii, 37bl8. rbc: see re
MUTATIONS 201
re: red cells cytology: Placed in region between 35C3 and 36B5
location: 2-36.8 (between d and / ) . on the basis of being to the right of Df(2L)64j =
origin: Spontaneous. Dt(2L)34E5'Fl;35C3-Dl and to the left of the de-
discoverer: E. B. Lewis, 1946. ficiency from T(Y;2)G = T(Y;2)36B5-C1;4OF.
synonym: rbc: zed blood cells.
Jones and Lewis, 1957, Biol. Bull. 112: 220-24
(fig.).
Grell, 1961, Genetics 46: 925-33.
phenotype: rc/rc normal; in lys rc/lys re, fat cells
of head and thorax acquire brownish red pigment.
Effect most prominent in one or more rows of pig-
mented cells along mid-dorsal line of thorax just
beneath chitin. Pigment is ommochrome since lys
re bw cells are pigmented, whereas v; lys re cells
are colorless except in kynurenine-fed flies. RK3.
discoverer: R. F. Grell, 1957.
references: Grell, 1961, Genetics 46: 925—33.
phenotype: Wild type at 25°C on standard medium;
at 17° a few red fat cells are visible. Early third
instar larvae placed on glucose-agar medium pro-
duce flies with numerous red cells, lys re2 has red
cells under any condition. RK3. re/*: reduced-scraggly
rd*: reduced-scraggly
discoverer: Bridges, 18j2.
phenotype: More bristles reduced and remaining
ones more irregular than in rd. More extreme in
male. Abdominal banding abnormal in female. Both
sexes fertile. RK1.
RD(1): Recovery Disrupter (1)
location: 1-62.9 [lO percent of the distance between
car and suf/)].
origin: Found in a chronically irradiated population
obtained from B. Wallace.
discoverer: Hanks, 1957.
references: Novitski and Hanks, 1961, Nature 190:
989-90.
Erickson and Hanks, 1961, Am. Naturalist 95:
rd: reduced 247-50.
Prom Bridges and Brehme, 1944, Carnegie Inst. Wash. phenotype: Males containing this factor, RD(2), and
Publ. No. 552: 157. certain other factors produce approximately 67 per-
cent female and 33 percent male progeny. The ef-
rd: reduced fect is not produced by zygotic mortality but by a
location: 2-51.2. mechanism that operates during meiosis, leading to
origin: Spontaneous. fragmentation of the Y chromosome, and production
discoverer: Bridges, 17gl5. of fewer than 64 sperm heads per sperm bundle
references: Morgan, Bridges, and Sturtevant, 1925, (Erickson, 1965, Genetics 51: 555-71). The effect
Bibliog. Genet. 2: 233. is maximal at 25°C and less pronounced at both
phenotype: Bristles, in varying numbers and posi- 18° and 27°. Viability good but fertility reduced
tions, strongly reduced in size; others unaffected. both sexes. RK3.
Reduced bristles usually curved and pointing in RD(2)
odd directions. Sternopieurals best criterion. Male location: 2- (not located).
more extreme than female. Female usually sterile origin: Found in a chronically irradiated population
(Lynch, 1919, Genetics 4: 501-33). RK2. obtained from B. Wallace.
discoverer: Hanks, 1960. references: 1940, Dokl. Acad. Nauk SSSR 27: 1030-
references: Novitski and Hanks, 1961, Nature 190: 32.
989-90. phenotype: Eye size reduced from the normal 750 to
phenotype: Males with this factor, RD(1), and cer- about 180 facets. Reduction more extreme in com-
tain other factors produce about 67 percent female bination with £?; some flies have no facets and are
progeny. RK3. sterile. RK2.
*rdi>: reddish brown re: see rey
location: 1-21.7. re*: see rey^
origin: Induced by methyl methanesulfonate (CB. *re-b: reduced eyes-b
1540). location: 3-45.
references: 1959, DIS 33: 89. discoverer: Whittinghill, 53g.
phenotype: Eye color deep reddish brown. Wings references: Schacht, 1954, DIS 28: 78.
frequently curve slightly upward at tips. Body phenotype: Eyes reduced in 80 percent of homozy-
somewhat small. Male sterile. Viability about 30 gotes. Expression varies independently in each
percent wild type. RK3. eye from absence of facets to wild type. RK2.
*rdm: reduced macros other information: Possibly allelic to re.
location: 1-59.8. rea: rearranged tergites
origin: Induced by 2-fluoroethyl methanesulfonate location: 1-25.4.
(CB. 1522). origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
discoverer: Fahmy, 1957. phenylalanine (CB. 3007).
references: 1959, DIS 33: 89. discoverer: Fahmy, 1954.
phenotype: Most bristles thin and short. Eye shape references: 1958, DIS 32: 73.
slightly abnormal. Body short; wings short, broad, phenotype: Tergites highly abnormal, partly
and frequently pleated. Male fertile. Viability missing, and different segments united. Expres-
about 10 percent wild type. RK3. sion variable. Viability and fertility inversely re-
lated to tergite abnormality. RK2.
rdo: reduced ocelli other information: One allele induced by CB. 3025.
location: 2-53. Recovery Disrupter: see RD
origin: Spontaneous. red: red Malpighian tubules
discoverer E. M. Wallace, 37113. location: 3-53.6.
phenotype: Ocelli small and colorless, often origin: Spontaneous.
missing, leaving top of head smooth and sometimes discoverer: Muller, 49a.
pigmented. Hairs between ocelli fewer than wild synonym: bw-1: brown-like.
type. Eye surface irregular. RK2. references: Oster, 1954, DIS 28: 77-78.
Aslaksen and Hadorn, 1957, Arch. Julius Klaus-
origin: Spontaneous. Stift. Vererbungsforsch. Sozialanthropol.
discoverer Bridges, 38bl 0. Rassenhyg. 32: 464—69.
phenotype: Like rdo. RK2. phenotype: Malpighian tubes of larva and adult rusty
*rdp: reduplicated red. Eye color brown, darkening with age. Mal-
location: 1-34.7. pighian tubes of v; red and en; red colorless; bw;
discoverer: Hoge-Richards, 12k. red tubes red; pigment therefore an ommochrome.
references: Hoge, 1915, J. Exptl. Zool. 18: 241-97. Eyes contain less drosopterin and isoxanthopterin
phenotype: At low temperatures, most flies have but more of the other pteridines than normal. Eye
malformed or branched legs, often with mirror color autonomous in red eye disks transplanted
image reduplication. At 25°C most flies normal. into wild-type hosts. Wild-type Malpighian tubes
RK3. acquire some red pigment after transplantation into
*rdt: reduced thorax red hosts. RK1.
location: 1-54.4. cytology: Placed in region 88A through 88C, based
origin: Induced by p-NN-di-(2-chloroethyl)atnino- on its inclusion in duplication derived from
phenylethylamine (CB. 3034). T(l;3)OS = T(1;3)4F2-3;62B-C;88A-C;92C-D
discoverer: Fahmy, 1955. (Lindsley and Grell, 1958, DIS 32: 136) and its
references: 1959, DIS 33: 89. genetic position to the left of cv-c (C. Hint on),
phenotype: Head and thorax disproportionately small which has been placed in region 88A-C.
compared to abdomen. Wings short, reaching only red blood cells: see re
to tip of abdomen; frequently incompletely ex- red cells: see re
panded or misheld. Male inviable and usually red Malpighian tubules: see red
sterile. RK3. reddish brown: see rdb
re: reduced eyes reduced: see rd
location: 3- (not located). reduced eyes: see re
origin: Spontaneous. reduced macros: see rdm
discoverer: Rapoport. reduced ocelli: see rdo
MUTATIONS 203
reduced pigment: see rg/ synonym: re (preoccupied).

reduced size: see rsi references: 1942, DIS 16: 52.
reduced tarsi: see rta phenotype: Eyes small and rough. RK3.
reduced thorax: see rdt rey; see rey?
reduplicated: see rdp *rey2
reduplicated sex combs: see rsc origin: Spontaneous.
re/: refractaire discoverer: Sturtevant, 1948.
location: 2-52.8. synonym: Described as rey.
origin: Spontaneous. references: 1948, DIS 22: 55-56.
discoverer: Ohanessian-Guillemain, 53b. phenotype: Eyes extremely small and rough in male,
references: 1953, DIS 27: 59. less extreme in female. Areas of thorax often un-
phenotype: Morphologically normal. Growth of the developed; sometimes hemithoracic. RK1.
carbon dioxide-sensitivity virus inhibited in other information: Allelism inferred from similarity
ref/ref. RK3. of phenotype and location to rey.
refringent: see rfr rey3
Resistancef ): see Rst( ) origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
*ret: reticulated phenylalanine (CB. 3025).
location: 1- (rearrangement). discoverer: Fahmy, 1953.
origin: Induced by L-p-NN-(2-chloroethyl)amino- synonym: re 2 : rougheye'like.
phenylalanine (CB. 3025). references: 1958, DIS 32: 73.
discoverer: Fahmy, 1953. phenotype: Eyes small and rough. Homozygous fe-
references: 1958. DIS 32: 73. male viable but infertile. RK2.
phenotype: Wing veins increased to anastomosing other information: One allele induced by CB. 3007.
reticulated areas. Wings shortened, deformed, and Allelism to rey inferred from phenotype and genetic
blistered. Eyes large and rough. Postvertical position.
bristles usually absent. Male sterile; viability *rf: roof wings
about 20 percent wild type. RK2A. location: 2-81.
cytology: Associated with T(l;2)ret - T(l;2)20A5- discoverer: Bridges, 1921.
B2;2R. references: Morgan, Bridges, and Sturtevant, 1925,
Rev: Revolute Bibliog. Genet. 2: 233.
location: 2- (rearrangement), phenotype: Wings rotated on long axis so that inner
origin: X ray induced, margins are raised and costal margins lowered.
discoverer Dobzhansky, 31b5. Overlaps wild type. RK3.
phenotype: Wings of heterozygote spread at 45° from rV
midline; edges curled, giving spoon shape. Sense origin: Spontaneous.
organs along veins enlarged. Eyes mottled in discoverer. Redfield, 1926.
Rev/It. Homozygote viable and fertile; somewhat references: Franke, 1933, Ph.D. Thesis, Univ.
more abnormal than heterozygote. Phenotype sup- Berlin.
pressed by extra Y'&; probably a variegated posi- phenotype: Like rf. RK3.
tion effect. RK2A. Rf: Roof
cytology: Associated with In(2LR)Rev = location: 3-59.
In(2LR)40F;52D10-El (Bridges and Li, in Morgan, origin: Spontaneous.
Bridges, and Schultz, 1936, Carnegie Inst. Wash. discoverer: Waddington, 38a.
Year Book 35: 293). references: 1939, DIS 12: 48-49.
Rev*: Revolute of Bridges phenotype: Wing position normal at eclosion; be-
origin: Spontaneous as a single homozygous female comes roof like in 12-hr imagos. RK1.
in a culture with no heterozygote. *Rf-c: Roof-c
discoverer: Bridges, 36e22. location: 3- (to the left of se).
synonym: Rvd: Revotutoid. discoverer: Bridges, 20al.
references: Morgan, Bridges, and Schultz, 1936, references: Bridges and Morgan, 1923, Carnegie
Carnegie Inst. Wash. Year Book 35: 293. Inst. Wash. Publ. No. 327: 228 (fig.).
phenotype: Wings spread and curved. Extreme Rev phenotype: Wings slanted at rooflike angle. RK3.
allele. Homozygous lethal but Rev/Rev® viable rfr: refringent
(E. B. Lewis). RK2A. location: 1-67.9.
cytology: Associated with In(2LR)RevB •» origin: Induced by D-p~NN-di-(2-chloroethyl)amino-
In(2LR)40;52C-E (Lewis). phenylalanine (CB. 3026).
Revolutaick see Rev& discoverer: Fahmy, 1955.
*rey: rough eye synonym: ret (preoccupied).
location: 1-0.6 (from combined measurements on rev, references: 1959. DIS 33: 89.
rey3, and rey3)* phenotype: Wing surface yellowish and iridescent;
origin*. Spontaneous. occasionally, one or both wings held out; inner
discoverer: Neel, 41g7. margins may be incised. Expression more extreme
in male than female. Male viable and fertile; fe- origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
male has reduced viability and is sterile. RK2. phenylalanine (CB. 3025).
other information: One allele each induced by CB. discoverer: Fahmy, 1954.
3026 and CB. 3034. references: 1959, DIS 33: 89.
rg: rugose phenotype: Characteristic pigmentation of fifth
location: 1-11.0. tergite reduced or absent in male. Body color
discoverer: Demerec, 28f23. yellowish. Eyes bright red. Male sterile. RK2.
synonym: rough-64. rh: roughish
phenotype: Eyes rough. Wings thin, with margins location: 2-54.7.
somewhat frayed. Viability excellent. RK2. origin: Spontaneous.
cytology: Locus at 4E1-3 (Demerec, Kaufmann, discoverer: Bridges, 21a3.
Fano, Sutton, and Sansome, 1942, Carnegie Inst. phenotype: Eyes moderately rough. At 19°C,
Wash. Year Book 41: 191). bristles slightly wavy and wings broad. RK2.
*rg rh: see gt3
synonym: roughish*
phenotype: Eyes uniformly rough. Viability 70 per-

synonym: pebbled.
phenotype: Eyes slightly rough; occasionally over- ri: radius incompletus
laps wild type. Viability 80 percent wild type. Edith M. Wallace, unpublished.
RK3. ri: radius incompletus
*rg* location: 3-47.0; to the left of the centromere,
discoverer: Ives, 33g22. based on mapping to left of Dp(l;3)sn13al =
synonym: rg33*. Dp(l;3)6C;7C9-Dl;79D2-El (Muller, 1958, DIS 32:
references: Plough and Ives, 1935, Genetics 20: 140) and Dp(l;3)N264-58 - Dp(l;3)3B2-3;3D6-7;
42-69. 80D-F (Gersh, 1966, DIS 41: 89).
1934, DIS 2: 34. origin: Spontaneous.
phenotype: Eyes somewhat rough. Viability excel- discoverer: Tshetverikov, 1926.
lent. RK2. phenotype: Vein L2 interrupted. Wings slightly
rg
warped and' blunt. Acts during contraction period
origin: Spontaneous. in Droaophila simulans, inhibiting fusion of small
discoverer: Bridges, 38c9. spaces into a vein (Waddington, 1940, J. Genet.
phenotype: More extreme than rg. Viability low. 41: 75-139). RK1.
RK2. cytology: Tentatively placed salivary region 77
7 (Hannah, Arajarvi, and Puro).
origin: X ray induced. *,/2
discoverer: Cantor, 46d20. discoverer: Nordenskiold, 36c5.
phenotype: Eyes rough and smaller than wild type. phenotype: Like ri. RK1.
Eclosion delayed. Viability and fertility excellent.
RK1A. origin: Spontaneous,
cytology: Associated with In(l)rg7 =*ln(l)4E;7A discoverer: Meyer, 51k.
(J. I. Valencia). references: 1952, DIS 26: 67.
see t£* phenotype: Less extreme than ri, RK1
3i
*rgp: rugose from P32 origin: Spontaneous.
origin: Induced by P 3 2 . discoverer: Meyer, 53j.
discoverer. Bateman, 1950. references: 1953, DIS 27: 58.
synonym: tes: facetious. phenotype: Like ri. RK1.
references: 1950, DIS 24: 54. RIDDT: seeRst(2)DDT
1951, DIS 25: 77-78. RI": see Rst(2)DDT
phenotype: Eyes small and rough; body pale; wings rickets: see rk
often curled upward. RK2A. rimy: see rm
cytology: Associated with ln(l)rgp În{l)3C;4E rk: rickets
(Darby). location: 2-48.2.
*rgt: reduced pigment origin: Ultraviolet induced.
! ©cation: 1-11.5. discoverer. Edmonds on, 48h.
MUTATIONS 205
references: 1948, DIS 22: 53. in bal, but less deformed. Subnormal viability, fer-
phenotype: L e g s , e s p e c i a l l y hind o n e s , flattened tility good. RK2.
and bent. Femora and tibiae bowed in middle; first rl: rolled
two tarsal joints shortened, bent and flattened; location: 2-55.1 [between centromere and stw
last three tarsal joints almost a unit, shortened (Sturtevant); 0.03 unit to the left of stw (Tano,
and flattened; tarsal claws disarranged. Wings not 1966, Japan. J. Genet. 41: 299-308)].
expanded, sometimes partially extended, some- discoverer: Bridges, 22f23.
times drooping. Postcutellar bristles c r o s s e d . references: Morgan, Bridges, and Sturtevant, 1925,
Body small. Viability about 90 percent wild type. Bibliog. Genet. 2: 233.
RK2. phenotype: Wing edges rolled downward, margins
cytology: Placed in region between 34E5 and 35D1 somewhat frayed, L4 interrupted distal to posterior
on basis of its inclusion in Df(2L)64j = cross vein. Eyes small, dark, and rough. Most ex-
Df(2L)35E5-Fl;35C3-Dl (E. H. Grell). treme at 25°C, less extreme above and below that
*rk* temperature (Lakovaara, 1963, Proc. Intern. Congr.
origin: Ultraviolet induced. Genet., 11th.Vol. 1: 175). RK2.
discoverer: Erickson, 50a. cytology: Placed in 41A, on basis of its inclusion
references: Meyer, Edmonds on, Byers, and in Df(2R)M-S21 ° =Df(2R)41A (Morgan, Schultz, and
Erickson, 1950, DIS 24: 60. Curry, 1941, Carnegie Inst. Wash. Year Book 40:
phenotype: Tarsi weak, but usually not deformed. 284).
At 21°C, wings curved downward and held out *r/G29; rolled of Goldschmidt
slightly; may be crumpled or unexpanded. Expres- origin: Recovered among progeny of heat-treated
sion more extreme at 27°C; overlaps wild type at flies.
17°. Viability fair at 17°, low at 27°. rk2/rk inter- discoverer: Goldschmidt, 1929.
mediate between the two homozygotes. RK2. references: 1929, Biol. Zentr. 49: 437-48.
*rk3 1939, Am. Naturalist 73: 547-59.
origin: Ultraviolet induced. phenotype: Like rl. RK2.
discoverer: Meyer, 54d. *rlu: rolled up
references: 1955, DIS 29: 74. location: 1- (rearrangement).
phenotype: Wings unexpanded; legs warped; body origin: Spontaneous in In(l)scsl + dl-49.
small; bristles fine. Viability low. RK2. discoverer: Reddi.
rk* references: 1963, DIS 37: 53.
origin: Spontaneous. phenotype: Wings rolled. Good viability and fer-
discoverer: Jackson, 54c. tility. RK2A.
synonym: cq: creeper. rm: rimy
Meyer, 1958, DIS 32: 83. origin: Induced by 2-chloroethyl methanesulfonate
phenotype: Wings unexpanded, spread, and drooping. (CB. 1506).
Posterior legs malformed. Both sexes fully viable discoverer: Fahmy, 1956.
and fertile, rk4 male mates with wild-type female references: 1958, DIS 32: 74.
only if wings removed from female. Viability 60 phenotype: Eyes often dull brownish red with con-
percent wild type. RK2. spicuous white hairs between ommatidia. Wings
*rk* longitudinally pleated. Viability and fertility good.
origin: Spontaneous. RK2.
discoverer: Mischaikow, 59a. other information: One allele each induced by CB.
references: 1959, DIS 33: 98. 1540 and CB. 1592.
phenotype: Less extreme than rk. No leg abnor- TJQ: s e e rmp
mality. Wings sometimes fully expanded but held *rmp: rumpled
out. RK3. location: 1-14.4.
*rk6 origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
origin: X ray induced. phenylalanine (CB. 3025).
discoverer: Thomas, 60g. discoverer Fahmy, 1955.
references: Meyer, 1963, DIS 37: 51. synonym: rm.
phenotype: Legs weak. Wings unexpanded. Via- references: 1959, DIS 33: 89.
bility higher at higher temperature. RK2. phenotype: Wings variably unexpanded. Bristles
*rfeey': rickets-cylindrical deranged; postvertica Is frequently crossed. De-
origin: Spontaneous. rangement of bristles correlated with degree of
discoverer Stroher, 1958. wing abnormality. Viability and fertility good in
synonym: cyl. both sexes. RK2.
references: Mainx, 1958, DIS 32: 82. m: rotund
phenotype: Abdomen cylindrical; terminal segments location: 3-47.7 [from location of rn3 (Carlson,
thickened. Posterior scute liars erect. Wings fail 1956, DIS 30: 109)].
to expand; halteres small and melanotic. Legs as origin: X ray induced.
discoverer: Glass, 1929. Roi: Rough eye

references: 1934, DIS 2: 8. location: 2- [left arm, not separated from In(2L)t\.
phonotype: Wings shortened but of normal width; origin: Spontaneous in In(2L)t.
nearly round. Tarsi three jointed. Sex combs ab- discoverer: Ives, 47kl8.
sent. Both sexes sterile; ovaries, follicles, references: 1952, DIS 26: 65.
oocytes, and eggs small (Beatty, 1949, Proc. Roy. 1956, DIS 30: 72.
Soc. Edinburgh, B 63: 249-70). Viability 50 per- phenotype: Eye facets of Roi/+ irregularly rounded,
cent normal. RK3A. sometimes enlarged; eyes sometimes bulge.
cytology: Associated with T(2;3)rn; breakpoints not Roi/Roi lethal, Roi/S viable. Acts as a partial
determined but probably chromocentral. suppressor of B (E. H. Grell). Viability good.
RK2A.
rolled: see rl
rolled up: see rlu
Roof: see Rf
roof wings: see rf
rose: see rs
rosy: see ry
Rosy: see hwy*
rotated abdomen: see rt
rotated penis: see rp
rotund: see rn
rough: see ro
rough eye: see rey
Rough eye: see Roi
rough III: see dli
Rough wing: see Rw
rough-64: see rg
roughened eye: see roe
Roughened: see R
roughest: see rst
roughestlike: see rstl
roughex: see rux
rougheye-likd: see rey3
roughish: see rgo
rn: rotund roughish: see rh
From Bridges and Brehme, 1944, Carnegie Inst. Wash. roughoid: see ru
Publ. No. 552: 159. *rp: rotated penis
location: 3-41.7.
origins Spontaneous,
origin: Spontaneous. discoverer: Bridges, 29cl5.
discoverer: Carlson. references: Morgan, Sturtevant, and Bridges, 1929,
references: 1956, DIS 30: 70, 109. Carnegie Inst. Wash. Year Book 28: 339.
phenotype: Wings round and tarsi small, like rn. Ul phenotype: As viewed from behind, external geni-
usually interrupted. Male sterile; female fertility talia of male rotated counterclockwise from 0° to
low. Viability good. RK2. 270°, usually about 180°; overlaps wild type in 30
ro: rough percent of flies. Eyes rough. Ply small; legs
location: 3-91.1. weak; tergites ridged; abdomen narrowed. Male
discoverer: Muller, 13f. sterile, even when genitalia not rotated. RK3.
references: Bridges and Morgan, 1923, Carnegie *rs: rose
Inst. Wash. Publ. No. 327: 93 (fig.). location: 3-35.0.
phenotype: Eyes rough, with facets irregular in size origin: Spontaneous.
and arrangement. Eyes slightly smaller and discoverer: Bridges, 23c 10.
narrower than wild type. RK1. references: Morgan, Bridges, and Sturtevant, 1925,
ro-63: see tm3 Bib Hog. Genet. 2: 234.
phenotype: Eye color translucent purplish pink but
me: roughened eye approaches wild type. Often sterile, especially
location: 3-47.6. male. Viability 80 percent wild type. RK2.
origin: Spontaneous. r*2
discoverer: Maxkowitz, 61 g. origin: Spontaneous.
references: 1963, DIS 38: 31. discoverer. Bridges, 38d5.
phenotype: Eyes rough and slightly smaller than phenotype: Eye color translucent pink. Viability
wild type. RK1. and fertility excellent. Larval Malplghian tubes
MUTATIONS
207
pale yellow (Brehme and Demerec, 1942, Growth 6: references: 1935, DIS 3: 27.
351-56). RK1. 1935, J. Genet. 31: 163-84.
*rsc; reduplicated sex combs 1937, J. Genet. 34: 169-89.
location: 1- (between y and cv). phenotype: Eyes rough; more extreme in male than
origin: X ray induced, female. Less extreme than other rst alleles.
discoverer: Yanders, 56f6. RK2A.
referencei: 1957, DIS 31: 85. cytology: Associated with In(l)rst3 =In(l)3C3-
phenotype: Sex combs present on all six legs of 4;20B (Emmens, 1937, J. Genet. 34: 191-202;
males. Overlaps wild type in crowded cultures. Kaufmann, 1942, Genetics 27: 537-49).
Wings droop. Male fertile but viability only 15 per- *rst264.S7
cent wild type; female lethal. RK2. origin: X ray induced simultaneously with N264-57,
rsd: raised discoverer: Demerec, 38d.
location: 3-95.4. references: 1941, Prdc. Intern. Congr. Genet., 7th.
origin: Spontaneous, pp. 99-102.
discoverer: Ives, 40i5. phenotype: Described only as not variegated.
references: 1945, DIS 19: 46. RK3A.
1947, DIS 21: 69. cytology: Associated with In(iyN™4-57 =ln(l)3C9-
phenotype: Wings held straight up, nearly meeting 11;2OD2-E1 (Hoover).
over thorax. Viability and fertility normal. RK1. *rsf264-86
other information: Possibly an allele of tx: taxi origin: X ray induced simultaneously with N264-86m
(3-91). discoverer: Demerec, 39i.
rsi: reduced size Demerec and Sutton, 1940, Proc. Natl. Acad. Sci.
location: 1-0.6 (no crossovers with br in 1038 flies). U.S. 26: 532-36.
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- Sutton, 1940, Genetics 25: 534-40.
phenylalanine (CB. 3026). phenotype: Like rst. RK2A.
discoverer: Fahmy, 1954. cytology: Associated with T(1;4)N264~86 -
references: 1959, DIS 33: 89. T(l ;4)3 C6-7;3C7-8;3E5-6}101F.
phenotype: Body small; eclosion delayed; viability Rst: Resistance
reduced. RK3, A term used to denote genes that confer resistance
other information: One allele each induced by CB. to the killing effects of insecticides. The symbol
1506 and CB. 3026. Rst is followed by parenthetical designation of the
*rst: roughest chromosomal location of the gene and then by an
location: 1-1.7. indication of the insecticide. Both dominant and
origin: X ray induced. recessive genes for insecticide resistance are con-
discoverer: Ball, 32b25. ceivable. Several investigators have exposed popu-
phenotype: Eyes rough and bulging; facets irregular lations to insecticides for numerous generations
in size and arrangement. Body small. Viability 70 and selected resistant lines. In most cases, the
percent wild type. Male sterile. RK2A. genetic basis of resistance is polygenic, and these
cytology: Associated with T(l;3)rst; breakpoint un- strains are not included in this list.
known in chromosome 3, X chromosome breaks near Rst(2)DDT: Resistance{2) DDT
w and bb (Beadle), rat locus in 3C4 (Slizynska, location: 2-65 (64.5-66).
1938, Genetics 23: 291—99; confirmed by Demerec, origin: Naturally occurring allele..
Kaufmann, Fano, Sutton, and Sansome, 1942, discoverer: Tsukamoto and Ogaki, 1953.
Carnegie Inst. Wash, Year Book 41: 191). synonym: RlDDT: Resistance to Insecticide-DDT;
RI11; Resistance to Insecticide on chromosome 2.
origin: Spontaneous. references: 1954, Botyu-Kagaku 19: 25.
discoverer: Bridges, 33d7. Tsukamoto, 1958, DIS 32: 87.
references: Gersh, 1965, Genetics 51: 477—80. Kikkawa, 1961, Ann. Rept. Sci. Works, Fac. Sci.,
phenotype: Eyes extremely rough and bulging; Osaka Univ. 9: 1-20.
facets irregular in size and arrangement. Body phenotype: Median lethal dose of DDT for
small. Some bristles absent; hairs sparse and ir- Rst(2)DDT lines is about 4000 ptg/cc of medium;
regular. Viability about 50 percent wild type. Fer- that for sensitive lines is 50—100 ptg/cc. Also re-
tility low. RK2A. sistant to BHC (benzene hexachloride) and organo-
cytology: Associated with Di(l)mta *aD£(l)3C3- phosphorus insecticides such as parathion and
4;3C6~7 (Schultz; confirmed by Gersh, 1965). malathion. Median lethal dose of parathion is 2
other information: Di(l)mt3 deficient for loci of both pptn for resistant line and 0.08 ppm for sensitive.
mt and vt; bristle abnormalities of rmt3 associated Sensitive to phenylthiourea (Ogita, 1958, Botyu-
with tfa* deficiency for vt (Gersh, 1965). Kagaku 23: 188-204). Shows maternal effect in
that progeny of Rat(2)DDT/+ female crossed to +/+
©ri-fln: X my Jtadoced.
male are more resistant than those of reciprocal
discoverer: 33116. cross. Larva more resistant than adult. RK3.
other information: Strains selected for resistance to ru: roughoid

DDT found to be resistant to parathion and to carry location: 3-0.0 [Actually about 4 units to the right
a factor for resistance in the same region of 2R as of the end of the chromosome, based on the loca-
resistance factors found in strains from the same tion of y * in T(l;3)scJ*].
population selected for resistance to parathion and discoverer: Sturtevant, 19bl4.
subsequently shown to be resistant to DDT. Thus references: Strong, 1920, Biol. Bull. 38: 33—37.
the resistance factors selected by exposure to Bridges and Morgan, 1923, Carnegie Inst. Wash.
DDT and parathion have been judged to be the Publ. No. 327: 212 (fig.).
same. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
*Rsi(3)ns: Resistance(3) nicotine sulfate Genet. 2: 215 (fig.), 234.
location: 3-49.5. phenotype: Eyes small and rough, with irregular
origin: Spontaneous. facets and hairs; black specks from erupted facets.
discoverer: Tsukamoto, 1954. Expression variable; sometimes overlaps wild
references: 1955, Botyu-Kagaku 20: 73. type. RK2.
1956, Botyu-Kagaku 21: 71. cytology: Placed in 61F5-62A3, on basis of its in-
1958, DIS 32: 87. clusion in Df(3L)ru-K2 = D%3L)61F4-5;62A10-Bl
phenotype: Median lethal dose to homozygote is 600 (Krivshenko, 1958, DIS 32: 81) and
pptn of nicotine sulfate added to culture medium Df(3L)ru300'23* = Df(3L)61E;62A2-4 (Ward and
(from first instar larva through eclosion); to hetero- Alexander, 1957, Genetics 42: 42—54).
zygote, it is 300 ppm; to susceptible strains, 40
ppm. RK3. origin: Spontaneous.
*rsth roughest I ike discoverer: Steinberg, 40k.
location: 1- (rearrangement), references: 1942, DIS 16: 54.
origin: X ray induced, phenotype: More extreme than ru. RK1.
discoverer: Oliver, 29d3. *ru100.392
synonym: lz-1: lozenge-like. origin: X ray induced.
references: 1935, DIS 3: 28. discoverer. Alexander.
phenotype: Eyes rough; more extreme than lz. Via- references: Ward and Alexander, 1957, Genetics 42:
bility low. RK2A. 42-54.
cytology: Associated with In(l)rstl; breakpoints un- cytology: Associated with Df(3L)ml00.392 =
known. Df(3L)61Ei62A10'Bl.
*rt: rotated abdomen
location: 3-37 (based on location of it2). *ru100.393
discoverer: Bridges, 18g28. origin: X ray induced.
references: Bridges and Morgan, 1923, Carnegie discoverer: Alexander,
Inst. Wash. Publ. No. 327: 190 (fig.). references: Ward and Alexander, 1957, Genetics 42:
Morgan, Bridges, and Sturtevant, 1925, Bibliog. 42-54.
Genet. 2: 54 (fig.). cytology: Associated with Df(3L)Tu100'393 =
phenotype: Abdomen twisted clockwise through 60° Df(3L)61F2-3;62A4-6.
to 90°, as viewed from behind. Both sexes sterile. *ru300.234
Viability low. RK2. origin: X ray induced.
rf2 discoverer: Alexander.
origin: Spontaneous, references: Ward and Alexander, 1957, Genetics 42:
discoverer: Bridges, 25114. 42-54.
phenotype: Abdomen twisted as is rt. Viability cytology: Associated with Df(3L)ru300.234 -
erratic, usually about 50 percent wild type. Male Dt(3L)6lE;62A2-4.
fertile; female not tested. RK2. ru9
*rt*: rotated abdomen of Vfallbrunn origin: Spontaneous.
origin: Gamma ray induced. discoverer Glass.
discoverer: Wallbrunn, 61126. references: 1934, DIS 2: 8.
references: 1964, DIS 39: 59. phenotype: Eyes small and extremely rough. More
phenotype: Like rt. RK2. reliable in classification than ru. RK1.
*rta: reduced tarsi
locations 1-4.5. rub: ru broad
origin: Induced by methyl methanesulfonate (CB. location: 2-5.0 (to the right of ho).
1540), origin: Spontaneous.
discoverer: Fahmy, 1956. discoverer: Mohr, 31k20.
references: 1959, DIS 33: 89. phenotype: Eyes rough and kidney shaped. Wings
phenotype: Tarsi short and sometimes deformed. broad and somewhat arc like. Abdomen short and
Body small. Eyes and wings small and abnormal. bloated; tergites irregular. External genitalia of
Bristles often waved or bent, postscuteliars often male rotated in varying degrees. Overlaps wild
held upright. Male sterile. RK2. type. RK3.
MUTATIONS 209
discoverer: Rolfes, 1960.

references: Hollander, 1960, DIS 34: 50.
phenotype: Eyes variably roughened; little reduction
in size. Both sexes fertile; viability about 50 per-
other information: Males of rux60<t stock mated to
attached-.^ females produced 17 homozygous rux60d
exceptional daughters among 9447 progeny
(Hollander and Festing, 1962, DIS 36: 79). This
production of equational exceptions has been
shown to be caused by a factor near f, probably an
allele of eq, by Thompson.
rv: raven
location: 1-4.4.
phenotype: Body small and heavily melanized. Eye
color dark. Wings short and frequently divergent or
not fully expanded. Male fertile but viability re-
duced; female more inviable and infertile. RK2.
rub: rubroad Rvd: see Rev&
Edith M. Wallace, unpublished. rw: raised wing
location: 2-93.2.
origin: Spontaneous, discoverer: Gomes, 55a.
discoverer: Chute, 48d. references: Bur dick, 1955, DIS 29: 70.
references: Sturtevant, 1948, DIS 22: 56. phenotype: Wings held vertically; venation normal.
phenorype: Like rub but a l s o wings show slight net- Legs morphologically normal but fly has difficulty
work of extra veins and thickening present between walking. Penetrance and expressivity good. Via-
L3 and L4. RK3. bility poor. RK2.
rubroad: see rub *Rw: Rough wing
ruby: see rb location: 2-56 [locus from crossing over in triploids
rud: ruddle (Schultz)].
location: 1-3.3. discoverer: Harnly.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- phenotype: Wings notched and veins irregular. An
phenylalanine (CB. 3025). occasional extra antenna. Rw/+/+ triploid female
discoverer: Fahmy, 1953. slightly fertile. Rw/+ female sterile. RK3.
references: 1958, DIS 32: 74. rwg: see hdp*"*^
phenotype: Eye color dull reddish brown. Classifi- ry: rosy
cation best in newly eclosed flies. Good viability location: 3-52.0 [0.3 unit to the right of kar
and fertility. RK2. (Schalet)].
other information: One allele each induced by CB. origin: Spontaneous,
1528, CB. 3026, and X rays. discoverer: Bridges, 38c4.
rudimentary: see r references: Glassman and Mitchell, 1959, Genetics
rugose: see rq 44: 153-62.
rumpled: see rmp Hubby and Forrest, 1960, Genetics 45: 211-24.
nix: rougbex Chovnick, Schalet, Kemaghan, and Talsma, 1962,
location: 1-15.0. Am. Naturalist 96: 281-96.
discoverer: Bridges, 33d24. Chovnick, Schalet, Kemaghan, and Krauss, 1964,
phenotype: Eyes smaller than wild type and uni- Genetics 50: 1245-59.
formly rough. Male sterile. RK2. phenotype: Eye color reddish brown; contains about
cytology: Locus from 5D3 through 6A2 (Demerec, 35 percent normal red pigment and 82 percent
Kaufmann, Fano, Sutton, and Sansome, 1942, normal brown pigment (Nolte, 1955, J. Genet. 53:
Carnegie lost. Wash. Year Book 41: 191). 1—10). Lacks detectable amount of x an thine dehy-
rux* drogenase, like mat (Forrest, Glassman, and
discoverer: Curry, 3711. Mitchell, 1956, Science 124: 725-26; Glassman
phenorype: Eyes small and rough like rux. Both and Mitchell, 1959; Hubby and Forrest, 1960). Ac-
sexes fertile. RK1. cumulates enzyme's substrates, hypo::anthine and
2-amino-4-hydroxypteridirse and lacks its products,
origin: Spontaneous. uric acid and isoxanthopterin (Mitchell, Glassman,
and Hadorn, 1959, Science 129: 268-69). In vitro

and in vivo complementation between ma/ + and ry* origin: Spontaneous.
demonstrated (Glassman, 1952, Proc. Natl. Acad. discoverer: Hubby.
Sci. U.S. 48: 1491-97; Glassman and McLean, references: Hubby and Forrest, 1960, Genetics 45:
1962, Proc. Natl. Acad. Sci. U.S. 48: 1712-18). 211-24.
Pigmentation nonautonomous in ry eye disks trans- Hubby, 1961, DIS 35: 46.
planted into wild-type hosts (Hadorn and Schwink, phenotype: Eye color reddish brown. Produces
1956, Nature 177: 940-41). RK1. traces of uric acid and isoxanthopterin but xanthine
cytology: Placed in region 87D-F, on basis of its in- dehydrogenase activity not demonstrable in ex-
clusion in Df(3R)ry = Df(3R)87D-E;87E-F (Grell, tracts. RK1.
1962, Z, Vererbungslehre 93: 371-77). ry3a
other information: Separable into at least six non- origin: X ray induced in cu kar chromosome,
complementing but recombinationally separable references: Chovnick, Schalet, Kernaghan, and
sites (Chovnick, Schalet, Kernaghan, and Krauss, Krauss, 1964, Genetics 50: 1245-59.
1964). ry recombines with ry26 and ry2 but not Schalet, 1964, DIS 39: 62-64.
with ry33, ry9, ry8, or ry42 and has been inter- phenotype: Eye color like ry. RK1.
preted as an intracistronic rearrangement. other information: To the right of ry23 and ry24 and
to the left of ry26. No crossovers recovered with
rys, ry9, ry4, or rys.
41
25 ry*
5 3a 7 origin: X ray induced in cu kar chromosome.
24 4 6 references: Chovnick, Schalet, Kernaghan, and
23 9 8 42 26 2 Krauss, 1964, Genetics 50: 1245-59.
i I i I 1 Schalet, 1964, DIS 39: 62-64.
phenotype: Eye color like ry. RK1.
other information: Between ry24 and ry26. No cross-
-3 overs recovered with ry9, ry3a, ry, or ry8.
8.86 x 10
ry-
origin: X ray induced in cu kar chromosome.
Map of ry locus references: Chovnick, Schalet, Kernaghan, and
From Chovnick, Schalet, Kernaghan, and Krauss, 1964, Krauss, 1964, Genetics 50: 1245-59.
Genetics 50: 1245-59. Schalet, 1964, DIS 39: 62-64.
phenotype: Eye color like ry. RK1.
other information: To the left of rys. No crossovers
ry2 recovered with ry23, ry24, ry9, or ry3*.
origin: Spontaneous. ry*
discoverer: Hadorn and Schwinck, 55c. origin: X ray induced in cu kar chromosome.
references: 1956, Nature 177: 940-41. references: Chovnick, Schalet, Kernaghan, and
1956, Z. Induktive Abstammungs- Vererbungslehre Krauss, 1964, Genetics 50: 1245-59.
87: 528-53. Schalet, 1964, DIS 39: 62-64.
phenotype: Eye color reddish brown, like ry; color phenotype: Eye color like ry. RK1.
of ocelli and testes approximately normal. Mal- other information: To the right of ry26.
pighian tubes slightly lighter than normal, short f7
and malformed, and contain large pteridine globules origin: X ray induced in cu kar chromosome.
in lumen. Lacks xanthine dehydrogenase, like ry references: Chovnick, Schalet, Kernaghan, and
(Forrest, Glassman, and Mitchell, 1956, Science Krauss, 1964, Genetics 50: 1245-59.
124: 725—26). Pigmentation nonautonomous in Schalet, 1964, DIS 39: 62-64.
transplants of ry2 eye anlage into wild-type hosts phenotype: Eye color like ry. RK1.
and in eyes of ry2 hosts transplanted with wild-type other information: To the right of ry26.
fat bodies, Malpighian tubes, and eye disks. Wild- ry 8
type eye disks transplanted into ry2 hosts form re- origin: X ray induced in cu kar chromosome,
duced amount of red eye pigment. Reciprocal trans- references: Chovnick, Schalet, Kernaghan, and
plants of eye disks or Malpighian tubes between ry2 Krauss, 1964, Genetics 50: 1245-59.
and tattl did not increase drosopterin formation Schalet, 1964, DIS 39: 62-64.
(Schwink, 1960, DIS 34: 105). Survival of ry2 tem- phenotype: Eye color like ry. RK1.
perature sensitive in early pupa; low at 25°C but other information: To the right of ry23, ry24, rys,
normal at 18°. Drosopterin formation at 18°C during and ry9 and to the left of ry26. No crossovers re-
late pmpm and early imago about twice that at 25°. covered with ry3* or ry4.
RK1. -9
ottar information: To the right of ry26 (Chovnick, origin: X ray induced in cu kar chromosome.
Schalet, Keraagbein, and Krauss, 1964, Genetics references: Chovnick, Schalet, Kernaghan, and
50: 1245-59). Krauss, 1964, Genetics 50: 1245-59.
MUTATIONS 211
Schalet, 1964, DIS 39: 62-64. references: 1964, DIS 39: 62-64.
phenotype: Eye color like ry. RK1. phenotype: Eye color like ry. RK1.
other information: Between ry23 and ry&. No cross- *ry22
overs recovered with ry24, rys, ry3a, or ry*. origin: X ray induced.
*ry> 0 discoverer: Schalet.
origin: X ray induced in 1(3)26 Sb Ubx chromosome. references: 1964, DIS 39: 62-64.
discoverer: Schalet. phenotype: Eye color like ry. RK1.
references: 1964, DIS 39: 62-64. ry23
phenotype: Eye color like ry. RK1. origin: X ray induced in cu kar chromosome.
*ryn references: Chovnick, Schalet, Kernaghan, and
origin: X ray induced in 1(3)26 Sb Ubx chromosome. Krauss, 1964, Genetics 50: 1245-59.
discoverer: Schalet. Schalet, 1964, DIS 39: 62-64.
references: 1964, DIS 39: 62-64. phenotype: Eye color like ry. RK1.
phenotype: Eye color like ry. RK1. other information: To the left of ry9. No crossovers
*ry12 recovered with ry2* or ry$.
origin: X ray induced in 1(3)26 Sb Ubx chromosome. r y24
discoverer: Schalet. origin: X ray induced in cu kar chromosome.
references: 1964, DIS 39: 62-64. references: Chovnick, Schalet, Kernaghan, and
phenotype: Eye color like ry. RK1. Krauss, 1964, Genetics 50: 1245-59.
*ry13 Schalet, 1964, DIS 39: 62-64.
origin: X ray induced in 1(3)26 Sb Ubx chromosome, phenotype: Eye color like ry. RK1.
discoverer: Schalet. other information: To the right of ry3a. No cross-
references: 1964, DIS 39: 62-64. overs recovered with ry23, ry5, or ry9.
phenotype: Eye color like ry. RK1. ry 25
*ry14 origin: X ray induced in cu kar chromosome.
origin: X ray induced, references: Chovnick, Schalet, Kernaghan, and
discoverer: Schalet. Krauss, 1964, Genetics 50: 1245-59.
references: 1964, DIS 39: 62—64. Schalet, 1964, DIS 39: 62-64.
phenotype: Eye color like ry. RK1. phenotype: Eye color like ry. RK1.
*ry1S other information: To the right of ry26.
origin: X ray induced, ry26

phenotype: Eye color like ry. RK1. Krauss, 1964, Genetics 50: 1245-59.
*fyl6 Schalet, 1964, DIS 39: 62-64.
origin: X ray induced, phenotype: Eye color like ry. RK1.
discoverer: Schalet. other information: To the right of ry42 and to the
references: 1964, DIS 39: 62—64. left of ry2. Used as a reference point for locating
phenotype: Eye color like ry. RK1. mutants to the right or left portions of the ry
cistron.
ry 17
origin: X ray induced. ry27
discoverer: Schalet, origin: X ray induced in cu kar chromosome.
references: 1964, DIS 39: 62-64. discoverer: Schalet.
phenotype: Eye color like ry. RK1. references: 1964, DIS 39: 62-64.
ry 78 Schalet, Kernaghan, and Chovnick, 1964, Genetics
origin: X ray induced. 50: 1261-68.
discoverer Schalet. phenotype: Homozygous lethal. Eye color of
references: 1964, DIS 39: 62-64. ry2?/ry2 like ry. RK2A.
phenotype: Eye color like ry, RK1. cytology: Association with Df(3R)ry27 (breakpoints
ry 19 unknown) inferred from genetic data.
origin: X ray induced. •^28
references: 1964, DIS 39: 62-64. discoverer: Schalet.
phenotype: Eye color like ry. RK1. references: 1964, DIS 39: 62—64.
ry20 Schalet, Kernaghan, and Chovnick, 1964, Genetics
origin: X ray induced. 50: 1261-68.
discoverer: Schalet. phenotype: Homozygous lethal. Eye color of
references: 1964, DIS 39: 62—64. ry28/ry2 like ry. RK2A.
phenotype: Eye color like ry. RK1. cytology: Association with D§(3R)ry2S (breakpoints
ll unknown) inferred from genetic data.
origin: X ray induced, *ry29
discoverers Schalet. origin: X ray induced in a cu kar chromosome.
discoverer: Schalet. references: 1964, DIS 39: 62-64.

references: 1964, DIS 39: 62-64. Schalet, Kernaghan, and Chovnick, 1964, Genetics
Schalet, Kernaghan, and Chovnick, 1964, Genetics 50: 1261-68.
50: 1261-68. phenotype: Homozygous lethal. Eye color of
3
phenotype: Homozygous lethal. Eye color of ry S/Ty2 like ry. RK2A.
ry29/ry2 like rym RK2A. cytology: Associated with T(l;3)ry3S =
cytology: Association with Dt(3R)ry29 (breakpoints T(l;3)20;87C-E;91B-C (Lindsley).
unknown) inferred from genetic data. U
*ry30 origin: X ray induced in a cu kar chromosome.
origin: X ray induced in a cu kar chromosome. discoverer Schalet.
50: 1261-68. phenotype: Homozygous lethal. Eye color of
phenotype: Homozygous lethal. Eye color of ry36/ry2 like ry. RK2A.
ry30/ry2 like ry. RK2A. cytology: Association with Df(3R)ry36 (breakpoints
cytology: Association with Df(3R)ry30 (breakpoints unknown) inferred from genetic data.
unknown) inferred from genetic data.
*ry3 7
*ry31 origin: X ray induced in a cu kar chromosome.
origin: X ray induced in a cu kar chromosome, discoverer: Schalet.
references: 1964, DIS 39: 62-64. phenotype: Eye color like ry. RKl.
Schalet, Kernaghan, and Chovnick, 1964, Genetics *ry38
50: 1261-68. origin: X ray induced in cu kar chromosome.
phenotype: Homozygous lethal. Eye color of discoverer: Schalet.
ry31/rya like ry. RK2A. references: 1964, DIS 39: 62-64.
cytology: Association with Df(3R)ry31 (breakpoints phenotype: Eye color like ry. RKl.
unknown) inferred from genetic data. ry40
*ry32 origin: X ray induced in cu kar chromosome.
origin: X ray induced in a cu kar chromosome. discoverer: Schalet.
references: 1964, DIS 39: 62-64. phenotype: Eye color like ry. RKl.
Schalet, Kernaghan, and Chovnick, 1964, Genetics ry4l
50: 1261-68. origin: X ray induced in cu kar chromosome.
phenotype: Homozygous lethal. Eye color of references: Chovnick, Schalet, Kernaghan, and
ry3a/ry2 like ry. RK2A. Krauss, 1964, Genetics 50: 1245-59.
cytology: Association with Df(3R)ry32 (breakpoints Schalet, 1964, DIS 39: 62-64.
unknown) inferred from genetic data. phenotype: Eye color like ry. RKl.
*ry33 other information: To the right of ry26
origin: X ray induced in a cu kar chromosome.
Schalet, Kernaghan, and Chovnick, 1964, Genetics Krauss, 1964, Genetics 50: 1245-59.
50: 1261-68. Schalet, 1964, DIS 39: 62-64.
phenotype: Homozygous lethal. Eye color of phenotype: Eye color like ry. RKl.
ry33/ry2 like ry. RK2A. other information: To the left of ry26. Position to
cytology: Association with Df(3R)ry33 (breakpoints the right of ry8 inferred from low rate of recombina-
unknown) inferred from genetic data. tion of ry43 compared to rys with ry26.
*ty34 ry43
origin: X ray induced in a cu kar chromosome. origin: X ray induced in cu kar chromosome.
d i scoverer: Schalet. discoverer: Schalet.
references: 1964, DIS 39: 62-64. references: 1964, DIS 39: 62—64.
Schalet, Kernaghan, and Chovnick, 1964, Genetics phenotype: Eye color like ry. RKl.
50: 1261-68. *ry44
phenotype: Homozygous lethal. Eye color of origin: X ray induced in cu kar chromosome.
ry34/ry2 like ry. RK2A. discoverer: Schalet.
cytology: Association with D£[3R)ry34 (breakpoints references: 1964, DIS 39: 62-64.
unknown) inferred from genetic data. phenotype: Eye color like ry. RKl.
ry3S ry4S
origin: X ray induced in cu kar chromosome. origin: X ray induced in cu kar chromosome.
MUTATIONS 213
references: 1964, DIS 39: 62—64. *rySS

phenotype: Eye color like ry, RKl. origin: X ray induced.
*ry46 discoverer: Kernaghan.
origin: X ray induced in cu kar chromosome, references: Schalet, 1964, DIS 39: 62—64.
discoverer: Schalet. phenotype: Eye color like ry. RKl.
references: 1964, DIS 39: 62-64. ryS6
phenotype: Eye color like ry. RKl. origin: X ray induced.
*ry47 discoverer: Kernaghan.
origin: X ray induced in cu kar chromosome. references: Schalet, 1964, DIS 39: 62-64.
references: 1964, DIS 39: 62-64. ry 57
discoverer: Kernaghan.
ry48 references: Schalet, 1964, DIS 39: 62—64.
origin: X ray induced in cu kar chromosome. phenotype: Eye color like ry. RKl.
references: 1964, DIS 39: 62-64. ,58
ry4 9 discoverer: Kernaghan.
origin: X ray induced in cu kar chromosome, references: Schalet, 1964, DIS 39: 62-64.
rySO discoverer Kernaghan.
references: 1964, DIS 39: 62-64. 60
ry
*rySl discoverer: Kernaghan.
references: 1964, DIS 39: 62-64.
Schalet, Kernaghan, and Chovnick, 1964, Genetics origin: X ray induced.
50: 1261-68. discoverer: Kernaghan.
phenotype: Homozygous lethal. Eye color of references: Schalet, 1964, DIS 39: 62—64.
rySl/ry2 like ry. RK2A. phenotype: Eye color like ry. RKl.
cytology: Association with Df(3R)ry5i (breakpoints ry 62
unknown) inferred from genetic data. origin: X ray induced,
ry52 discoverer: Kernaghan.
origin: X ray induced in cu kar chromosome. references: Schalet, 1964, DIS 39: 62—64.
Schalet, Kernaghan, and Chovnick, 1964, Genetics origin: X ray induced,
50: 1261-68. discoverer: Kernaghan.
phenotype: Homozygous lethal. Eye color of references: Schalet, 1964, DIS 39: 62-64.
rySl/ry* like ry. RK2A. phenotype: Eye color like ry* RKl.
cytology: Association with Df(3R)ryS2 (breakpoints ry64
unknown) inferred from genetic data. origin: X ray induced,
ryS3 discoverer Kernaghan.
origin: X ray induced in cu kar chromosome, references: Schalet, 1964, DIS 39: 62—64.
discoverer Schalet. phenotype: Eye color like ry. RKl.
references: 1964, DIS 39: 62-64. ry65
ryS4 discoverer Kernaghan.
references: 1964, DIS 39: 62-64. *r/66
Schalet, Kernaghan, and Chovnick, 1964, Genetics origin: X ray induced,
5(h 1261-68. discoverer. Kernaghan.
phenotype: Homozygotes almost completely lethal; a references: Schalet, 1964, DIS 39: 62—64.
few homozygous females, which are also pic, Schalet, Kernaghan, and Chovnick, 1964, Genetics
survive. Eye color of ry^4/ry3 like ry. RK2(A). 50: 1261-68.
phenotype: Homozygous lethal. Eye color of phenotype: Homozygous lethal. Eye color of
ry66 /ry3 like ry. RK2A. rylt/ry3 like ry. RK2A.
cytology: Association with Dt(3R)ry66 (breakpoints cytology: Association with Df(3R)ry76 (breakpoints
unknown) inferred from genetic data. unknown) inferred from genetic data.
*ry6 7 * r y 77
origin: X ray induced. origin: X ray induced in kar3 chromosome.
discoverer: Kernaghan. discoverer: Schalet.
references: Schalet, 1964, DIS 39: 62-64. phenotype: Homozygous lethal. Eye color of
phenotype: Eye color like ry. RK1. ry77/ry3 like ry. RK2A.
cytology: Association with Df(3R)ry77 (breakpoints
*ry68 unknown) inferred from genetic data.
origin: X ray induced. *ry78
di scoverer: Kernaghan. origin: X ray induced in kar3 chromosome.
references: Schalet, 1964, DIS 39: 62-64. discoverer: Schalet.
phenotype: Eye color like ry. RK1. phenotype: Homozygous lethal. Eye color of
*ry69 ry78/ry3 like ry. RK2A.
origin: X ray induced, cytology: Association with Df(3R)ry78 (breakpoints
discoverer: Kernaghan. unknown) inferred from genetic data.
references: Schalet, 1964, DIS 39: 62-64. ry«t-F: rosy-el ectrophoretic Fast
phenotype: Eye color like ry. RK1. origin: Naturally occurring allele.
*ry70 discoverer: Yen.
origin: X ray induced. references: Yen and Glassman, 1965, Genetics 52:
discoverer: Kernaghan. 977-81 (fig.).
references: Schalet, 1964, DIS 39: 62-64. 1966, Genetics 54: 369-70.
Schalet, Kernaghan, and Chovnick, 1964, Genetics phenotype: Specifies a xanthine dehydrogenase
50: 1261-68. molecule that migrates relatively rapidly in poly-
phenotype: Homozygous lethal. Eye color of acrylamide gel electrophoresis. Kinetic parameters
ry70/ry3 like ry. RK2A. of enzyme same as those of other xanthine dehy-
cytology: Association with Df(3R)ry70 (breakpoints drogenase isozymes. In hybrids with ryei"s, en-
unknown) inferred from genetic data. zymes with at least three mobilities formed;
*ry7l strongest activity found in a position intermediate
origin: X ray induced, between mobilities of the enzymes specified by the
discoverer: Kernaghan. two parental types. RK3.
references: Schalet, 1964, DIS 39: 62-64. ryl'h rosy-el ectrophoretic Intermediate
phenotype: Eye color like ry. RK1. origin: Naturally occurring allele.
ry 72 discoverer: Yen.
origin: X ray induced. references: Yen and Glassman, 1965, Genetics 52:
discoverer Kernaghan. 977-81 (fig.).
references: Schalet, 1964, DIS 39: 62-64. 1966, Genetics 54: 369-70.
phenotype: Eye color like ry. RK1. phenotype: Specifies a xanthine dehydrogenase
Vy'73 molecule whose mobility is intermediate between
origin: X ray induced. mobility of molecules specified by ryei~F and
discoverer: Kernaghan. ry*t~s. Kinetic parameters same as those for other
references: Schalet, 1964, DIS 39: 62-64. xanthine dehydrogenase isozymes. RK3.
phenotype: Eye color like ry. RK1. ry«l-S: rosy-el ectrophoretic Slow
'ry' 74 origin: Naturally occurring allele.
origin: X ray induced. discoverer: Yen.
discoverer: Schalet. references: Yen and Glassman, 1965, Genetics 52:
phenotype: Homozygous lethal. Eye color of 977-81 (fig.).
ry74/rya like ry. RK2A, 1966, Genetics 54: 369-70.
cytology: Association with Df(3R)ry7* (breakpoints phenotype: Specifies a xanthine dehydrogenase
unknown) inferred from genetic data. molecule that migrates relatively slowly in poly-
*ry75 acrylamide gel electrophoresis. Kinetic parameters
origin: X ray induced in kar3 chromosome. of enzyme same as those of other xanthine dehy-
discoverer: Schalet. drogenase isozymes. In hybrids with ryol"F, en-
phenotype: Homozygous lethal. Eye color of zymes with at least three mobilities formed;
ry75/ry3 like ry. RK2A. strongest activity found in a position on the gel
cytology: Association with Df(3R)iy7S (breakpoints intermediate between the positions of the enzymes
unknown) inferred from genetic data. specified by the two parental types. RK3.
*ry76 rymt-Sl; rosy-el ectrophoretic Slow Intermediate
origin: X ray induced in kar3 chromosome. origin: Naturally occurring allele.
discoverer. Schalet. discoverer. Yen.
MUTATIONS 215
references: Yen and Glassman, 1965, Genetics 52: cytology: Salivary glands normal. Placed in the
977-81 (fig.). 21E1-2 doublet, on basis of its inclusion in the
1966, Genetics 54: 369-70. synthetic deficiency derived by combining Y-
phenotype: Specifies a xanthine dehydrogena se centric portion of T(Y;2)21E = T(Y;2)21D4-E1 and
molecule whose mobility is intermediate between 2-centric portion of T(2;4)astv = T(2;4)21E2-3;101.
mobility of molecules specified by ryel~s and Heterozygosity for deficiencies including 21E1-2
ryel-It Kinetic parameters same as those for other produces S phenotype (Lewis, 1945).
xanthine dehydrogenase isozymes. RK3. other information: A pseudoallele of ast. In cross-
over tests, 5 localizes to the left of ast (Lewis,
ryK: rosy of Kernaghan 1941, Proc. Natl. Acad. Sci. U.S. 27: 31-35;
origin: X ray induced in cu kar chromosome. Lewis, 1945, 1951).
discoverer: Kernaghan. S2
references: Schalet, 1964, DIS 39: 62—64. origin: Spontaneous in In(2L)Cy + Jn(2R)Cy and not
Schalet, Kernaghan, and Chovnick, 1964, Genetics separated from inversions.
50: 1261-68 discoverer: Redfield, 25k.
phenotype: Homozygous lethal. Eye color of references: Stern and Bridges, 1926, Genetics 11:
ryK/ty2 like ry* RK2A. 507-8.
cytology: Association with Df(3R)tyK (breakpoints phenotype: Like S. RK1A.
unknown) inferred from genetic data. SSlb
s: sable discoverer: Meyer, 51b.
discoverer Bridges, 12gl9. phenotype: Eyes of S*-*6/+ rough but not reduced in
references: Morgan and Bridges, 1916, Carnegie size. RK1.
Inst. Wash. Publ. No. 237: 34. SS6f
phenotype: Body color dark with prominent trident. origin: Synthetic.
Classification good at 19°C; overlaps wild type discoverer: Thompson, 56f.
increasingly with higher temperatures. Viability references: Burdick, 1956, DIS 30: 69.
sometimes reduced, s is nonautonomous in gynan- phenotype: Like S. S5**/S lethal. RK1A.
dromorphs containing both s and + tissue (Lewis, cytology: Associated with In(2LR)SS6i-
1955, DIS 29: 134). Tyrosinase formed in adult In(2LR)21 C8-Dl;60Dl-2^22A3-Bl;60B-CR derived
(Horowitz and Fling), s / s 2 easily classified. RK1 from single recombinant between In(2LR)bwV* and
at 19°C. In(2LR)SMl, which is deficient for 21D1-22A3.
S*
discoverer: Bridges, 17e9. *SD": Star-Deficiency

references: Morgan, Bridges, and Sturtevant, 1925, origin: X ray induced,
Bibliog. Genet. 2: 234. discoverer: E. B. Lewis, 1940.
phenotype: Body color less dark than s but trident references: 1945, Genetics 30: 147-51.
more prominent. Expression best at 19°C; overlaps phenotype: Less extreme than S. RK1A.
wild type at 25° and 30°. Viability excellent. RK1 cytology: Associated with Df(2L)Sl = Df(2L)21C3-
at 19°C. 4;22A2-3.
S: Star $Df2
location: 2-1.3 (0.02 unit to the left of ast). origin: X ray induced,
origin: Spontaneous. discoverer: E. B. Lewis, 1940.
discoverer: Bridges, 15bl2. references: 1945, Genetics 30: 147-51.
references: Bridges and Morgan, 1919, Carnegie phenotype: Like 5. RK1A.
Inst. Wash. Publ. No. 278: 259 (fig.). cytology: Associated with Dt(2L)S2 « Df(2L)21C6-
Morgan, Bridges, and Sturtevant, 1925, Bibliog. D1;22A6-Bt.
Genet. 2: 213 (fig.).
Lewis, 1945, Genetics 30: 137-66. origin: X ray induced,
1951, Cold Spring Harbor Symp. Quant. Biol. 16: discoverer: E. B. Lewis, 1940.
159-74 (f%.). references: 1945, Genetics 30: 147—51.
phenotype: Eyes slightly smaller and narrower than phenotype: Less extreme than S. RK1A.
wild type; texture somewhat rough from rounded, cytology: Associated with Df(2L)S3 - Df(2L)21D2-
irregular facets. Arrangement of hairs on surface 3.21F2-22AI.
of eye irregular. S/aat has small rough eyes; *$Df4
S m*t/+ + is like S/+. Enhanced by B(S); partially origin: X ray induced,
suppresses px and net (Bedichek, 1936, DIS 5; 24; discoverer E. B. Lewis, 1940.
Lewis, 1945). Hooozygote dies in late embryonic references: 1945, Genetics 30: 147-51.
stag® (Slvert*@v~DobKb*»sky, 1927, Arch. phenotype: Like S. RK1A.
Eotwickluogsiaech. Organ. 109: 535—48; Sotwien- cytology: Associated with Dl(2L)S4 **Df(2L)21C3-
blfcfc and Hoettoar, 1938, Genetics 23: 169). RK1. 4;22B2-3.

origin: X ray induced. 1932, Genetics 17: 637-59.
discoverer: E. B. Lewis, 1940. 1935, DIS 3: 15.
references: 1945, Genetics 30: 147-51. phenotype: Eye color wine at eclosion; becomes
phenotype: Less extreme than S. RK1A. dark salmon with age. Homozygous lethal. RK1A.
cytology: Associated with Df(2L)S5 = Df(2L)2lC2- cytology: Associated with T(2;3)Sa; breaks proximal
3.-22A3-4. in 2L and 3L.
*sab: straight abdomen
origin: X ray induced in net ho. location: 1-58.9.
discoverer: E. B. Lewis, 1940. origin: Induced by D-l:6-dimethanesulfonyl mannitol
references: 1945, Genetics 30: 147—51. (CB. 2511).
phenotype: Like 5. RK1A. discoverer: Fahmy, 1958.
cytology: Associated with Df(2L)S7 =Df(2L)21C3- references: 1964, DIS 39: 58.
4,-21 F2-22A1. phenotype: Abdomen long, narrow, and straight.
*SK: Star of Krivshenko Bristles somewhat fine. Male viable and fertile.
discoverer: Krivshenko. RK3.
references: 1936, DIS 5: 8. sable: see s
phenotype: Eyes rough like S. Reported to be homo- sable duplication: see su(s)
zygous viable. RK2A. safranin: see sf
cytology: Associated with In(2LR)SK; breakpoints salmon: see £
near ends of 2L and 2R. Salmon: see Sa
*$*-; Star of Lewis *saw: sawtooth
origin: X ray induced. location: 1-0.0 (very close to right of sc).
discoverer: E. B. Lewis, 1940. origin: Ultraviolet induced,
references: 1945, Genetics 30: 147—51. discoverer: Edmonds on, 51 g.
phenotype: Like 5. RK1A, references: 1952, DIS 26: 60.
cytology: Associated with T(2;3)SL = T(2;3)21E2- phenotype: Hairs along wing edge so arranged that
3;81F;88D6-8. edge appears serrated. Wings may warp, especially
S*: Star of duller in female. Fertility and viability excellent. Clas-
origin: X ray induced in In(2L)Cy +In(2R)Cy. sification originally easy, but stocks apparently
discoverer: Muller, 1928. accumulate modifiers so that they now appear
references: Painter and Muller, 1929, J. Heredity nearly wild type. RK2.
20: 287-98. other information: Not separated from sc in two
phenotype: Eyes like S, but perhaps more variable. crossovers between ac and sc or in 60 crossovers
RK1A. between sc and pn. Not covered by Dp(l;2)sci9 =
cytology: Associated with T(2;3)SM = T(2;3)21E2- Dp(l;2)lBl'2;lB4-7;25-26. Locus must be slightly
3;79D2-E1. Superimposed on In(2L)Cy = to the right of sc
In(2L)22Dl-2;33F5-34Al +ln(2R)Cy = In(2R)42A2'
3;58A4-B1; separable from the latter. origin: Ultraviolet induced.
Sr: see ast discoverer: Edmonds on, 51 f.
*S"; Star of Whittinghitt references: 1952, DIS 26: 61.
origin: X ray induced in Cy. phenotype: More extreme than saw. Wing margins as
discoverer: Whlttinghill, 47b. in saw, but wings strongly warped up or down; thin
phenotype: Like S but somewhat more extreme. texture especially in female. Viability reduced.
RK1A. Fly often becomes stuck in food owing to warped
other information: Inseparable from In(2L)Cy. wings. Fertility good; classification easy. RK2.
$X: Star from X irradiation sb: see ml
origin: X ray induced simultaneously with astx.
discoverer: E. B. Lewis, Sb: Stubble
references: 1945, Genetics 30: 157. location: 3-58.2.
phenotype: Eyes of Sx asr*/+ + slightly smaller origin; Spontaneous.
than S/+; L2 occasionally interrupted distally; discoverer. Bridges, 23d21.
resembles S +/+ astx, except for L2 abnormality. references: Dobzhansky, 1930, Z. Induktive
RK1. Abatamnmngs- Vererbungslehre 54: 427—57 (fig.).
cytology: Salivary chromosomes appear normal. phenotype: Bristles of Sb/+ less than one-half
other information: astx but not Sx has been re- normal length, and somewhat thicker than wild
covered alone from Sx ast^. type. HooK*zygous lethal. Sh/sbd2 more extreme
S-i: see e(S) than S&/+. sbdP Sb behaves as a recessive sbd
*Sa: Salmon allele but is homozygous lethal. Classifiable in
location: 2- or 3- (rearrangement). single dose in triploids. Developmental studies by
origin: X my induced. Lees and Waddington [l943, Proc. Roy. Soc.
discoverer: Van Atta, 30kl. (London), Ser. B 131: 87-110 (f%.)] show that
MUTATIONS 217
trichogen is shifted to lie more or less on the level phenotype: Sbv'/+ has mixture of wild-type and Sb
of the tormogen. RK1. bristles. In X/X/Y female and X/Y/Y male,
cytology: Salivary chromosomes normal (Morgan, bristles nearly all Sb. lnX/0 male, bristles usu-
Bridges, and Schultz, 1937, Carnegie Inst- Wash. ally all wild type. Sbv/Sb and homozygous Sbv
Year Book 36: 301). Placed in 89B4-5, probably in are lethal. RK1A.
89B4 by Lewis (1951, Cold Spring Harbor Symp. cytology: Associated with T(2;3)Sbv = T(2;3)41A-
Quant. Biol. 16: 159—74). These probably corre- C;88;89B. Superimposed on In(3R)Mo - In(3R)93D;
spond to 89B9-10 on Bridges's revised map. 98F2-6.
other information: Sb is pseudoallelic to and lies
0.01 to 0.03 unit to the right of sbd?. Deficiency
for the Sb locus produces no dominant phenotype
(Lewis, 4951).
sbd: stubbloid
From Dobzhonsky, 1930, Z. Indukttve Abstammungs-
Vererbungslehre 54: 427-57.
sbd: stubbloid
location: 3-58.2.
discoverer: Sturtevant, 1926.
synonym: S6 r ; Stubble-recessive.
references: Stern, 1929, Biol. Zentr. 49: 261—90.
Dobzhansky, 1930, Z. Induktive Abstammungs-
Vererbungslehre 54: 427-57 (fig.).
phenotype: Bristles short but usually slightly longer
than in Sb/+. One or both wings often shortened
and crumpled at base. Tibia and femur often
shortened, thickened, and bowed. Viability some-
what low. RK2.
cytology: Placed in region 89B4-5 by Lewis (1951,
Sb: Stubble Cold Spring Harbor Symp. Quant. Biol. 16: 159-
Edith M. Wallace, unpublished. 74). This probably corresponds to 89B9-10 on
Bridges's revised map.
other information: Pseudoallelic to Sb and lies to
origin: Spontaneous. the left of it (Lewis, 1951).
discoverer: Merriam, 63b.
phenotype: Bristles somewhat shorter and thicker origin: Spontaneous.
than Sb. Wings and legs normal. Homozygous discoverer: Harnly, 271.
lethal. Sb63b/Sb viable and fertile; more extreme synonym: So 1 " 2 .
than either heterozygote. RK1. phenotype: Most bristles about three-fourths normal
other information: Allelisra inferred from failure to length although some, for example, posterior post-
recover recombinants among 100 progeny of alars, are shorter. Less extreme than sbd. sbd^/Sb
has shorter bristles than homozygous sod 2 or S6/+.
S6r: see sbd sbd2 Sb/+ + has wild type bristles (Lewis, 1951,
SbSpi: Stubble-Spike Cold Spring Harbor Symp. Quant. Biol. 16: 159—
origin: X ray induced. 74). RK1.
discoverer: Moore, 31dl5.
references: 1935, DIS 3: 27. origin: X ray induced,
phenotype: Bristles of SbsP1/+ about two-thirds discoverer: E. B. Lewis.
normal length. Wings and legs normal. Bristles of references: 1948, DIS 22: 72-73.
homozygote one-fourth normal length. Wings re- cytology: Associated with Dt(3R)abdl°S
duced, crumpled, or blistered. Legs often short Dt(3R)88 F.89B4-5.
and bowed. S^P^Sb viability about 30 percent
wild type. Bristles and wings shorter than homozy- origin: X ray induced.
gous SbSpl. RK1. discoverer: E. B. Lewis.
Stubble-Variegated cytology. Associated with T(2;3)mbd1(>6
origin: X ray induced in In(3R)Mo, Sb ar. TC2;3)22B;89B.
discoverer: E. B. Lewis, 1948. sbdh stubbloid-letho!
references: 1956, DIS 30: 76-77. origin: X ray induced.
discoverer: E. B. Lewis. present [Lees and Waddington, 1942, DIS 16: 70-
references: 1949, DIS 23: 92. 70a; 1943, Proc. Roy. Soc. (London), Ser. B 131:
pbenotype: sbd!/sbd is sbd; sbdl/Sb is lethal. 87—HO]. Suppressed by su(Hw) and su(Hw)2. RK1.
RK2A. cytology: Placed in 1B3, on basis of its exclusion
cytology: Associated with T(2;3)Me = T(2;3)48C1- from Df(l)260-2 = Df(l)lB2-3, and its inclusion
2;59D2-3;80-81 + In(3L)63C;72El-2 + in the inverted section of In(l)sc8 = In(l)lB2-
In(3LR)69E;9lC + In(3R)89B;97D. 3.-20B-D1 but not of In(l)sc4 = In(l)lB3-4;19F-
sbr: small bristle 20C1 (Muller and Prokofyeva, 1935, Proc. Natl.
location: 1-33.4. Acad. Sci, U.S. 21: 16-26; Demerec, Kaufmann,
origin: Spontaneous. Fano, Sutton, and Sansome, 1942, Carnegie Inst.
discoverer: Curry. Wash. Year Book 41: 191; Sutton, 1943, Genetics
phenotype: Bristles small; one or more missing, par- 28: 210-17).
ticularly the posts cute liars. RK2. SC2
*sbs: stubs origin: X ray induced.
location: 1-0.9. discoverer: Dubinin, 1928.
origin: Induced by ethyl methanesulfonate (CB. references: 1929, Biol. Zentr. 49: 328-39 (fig.).
1528). Serebrovsky and Dubinin, 1930, J. Heredity 21:
discoverer: Fahmy, 1956. 259-65 (fig.).
references: 1959, DIS 33: 90. phenotype: Almost all bristles and hairs missing
phenotype: Wing abnormalities vary from extreme re- from sternites and tergites. Most scutellars and
duction in size to partial incision of margin with some humerals absent. Most bristles on head and
L2 and L3 closer together. Eyes small and thorax present, but the bare abdomen provides easy
slightly rough. Male viable and fertile. Female classification. Abdomen tends to be swollen.
sterile. RK2. Wings poorly expanded. Viability low. RK2.
*sbt: shorter bristles other information: Reported to be a transposition of
location: 1-32.8 (no crossovers with ras among 669). tip of X to X heterochromatin (Dubinin, 1929). The
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- s c 2 studied by Sturtevant was not a transposition
phenylalanine (CB. 3007). and mapped as a point mutation at the left end of
discoverer. Fahmy, 1954. theX.
references: 1959, DIS 33: 90. *sc3
phenotype: Bristles slightly short and thin. Wings origin: X ray induced (simultaneously with ac 2 ),
slightly divergent. Male late hatching. Viability discoverer: Dubinin, 1928.
and fertility greatly reduced. RK3. references: 1929, Biol. Zentr. 49: 328-39 (fig.)-
*sby: small body Serebrovsky and Dubinin, 1930, J. Heredity 21:
location: 1-60.8. 259-65 (fig.)-
origin: Gamma ray induced. phenotype: Most bristles affected, principally ven-
discoverer: Fahmy, 1958. trals, orbitals, verticals, postverticals, ocellars,
synonym: sby-61. humerals, presuturals, notopleurals, supra-alars,
references: 1964, DIS 39: 58. postalars, sternopleurals, abdominals, and anterior
phenotype: Extremely small, lightly pigmented fly. dorsocentrals; scutellars and postdorsocentrals
Viability and fertility reduced. RK.3. usually present. Viability of male low; female vir-
cytology: Placed in salivary region 18A4 to 18B8, tually lethal. RK2.
on basis of its inclusion within deficiency re- cytology: Salivary chromosomes appear normal
sulting from recombining left end of In(l)y4 - (Morgan, Bridges, and Schultz, 1935, Carnegie
ln(l)lA8'Bl;18A3-4 with right end of In{l)sc!> = Inst. Wash. Year Book 34: 290).
In(l)lB2-3;18B8-9 (Norton and Valencia, 1965, DIS sc3-»
40: 40), origin: Spontaneous derivative of sc3.
sby-6t: see sby discoverer: Sturtevant.
sby-62: see srb references: 1935, DIS 3: 15.
sc: scute phenotype: Partial reversion from sc3. RK2.
location: 1-0.0.
origin: Spontaneous. sc3B: scute-3 of Bridges
discoverer: Bridges, 16a22. discoverer: Bridges, 26d26.
references: Morgan, Bridges, and Sturtevant, 1925, phenotype: Like sc, but does not affect anterior
Bibliog. Genet. 2: 235, 211 (fig.), notopleurals. RK1.
phenotype: Causes loss or marked reduction in sc*
number of scutellar, coxal, oceilar, first and origin: X ray induced in y.
second orbital, anterior notopleural, postvertical, discoverer: Agol, 1928.
tergital, and sternal bristles. Bristle patterns for references: 1929, Zh. Eksperira. Biol. 5: 86-101
•c through sc** tabulated by Dubinin (1933, J. (fig.).
Genet. 27: 446). Bristle sockets missing; bristle Serebrovsky and Dubinin, 1930, J. Heredity 21:
cell* absent 19 hr after pupation, when normally 259-65 (fig.).
MUTATIONS 219
Agol, 1931, Genetics 16: 254-66. 1963 Verhandl. Deut. Zool. Ges., Zool. Anz.
Muller and Raffel, 1940, Genetics 25: 541-83. Suppl. 26: 87-92). RK2A.
phenotype: Extreme scute. Bristles of head, except cytology: Associated with In(l)sc8 -In(l)lB2-
anterior verticals, absent. Only posterior noto- 3;20B-Dl.
pleurals and alars remain on sides; abdominals, sc«c.o. X: see Df(l)scS
ventrals, coxals, and scutellars also missing. scDENc.o. X: see Df(l)scs
Slight variegation for Hw. RK1A. sc*
cytology: Associated with In(l)sc4 =In(l)lB3- origin: X ray induced.
4;19F-2OC1. discoverer: Levit, 1929.
SCS references: 1930, Arch. Entwicklungsmech. Organ.
origin: X ray induced in y. 122: 770-83.
discoverer: Gaissinovitsch, 1928. phenotype: Like sc, but scutellars always absent.
references: 1930, Zh. Eksperim. Biol. 6: 15—24. Abdomen swollen and wings poorly expanded like
Serebrovsky and Dubinin, 1930, J. Heredity 21: s C 2. RK2A.
259-65 (fig.). cytology: Associated with In(l)sc9 -ln(l)lB2-
phenotype: Sternital and scutellar bristles reduced 3;18B8-9.
in number; others rarely affected. sc5/sc6 is prac- sc1 °: see ac3
tically wild type. RK1.
sc« origin: X ray induced in In(l)ac3.
origin: X ray induced. discoverer: Sturtevant, 1930.
discoverer: Serebrovsky, 29a21. references: 1935, DIS 3: 15.
references: 1930, Arch. Entwicklungsmech. Organ. 1936, Genetics 21: 444-66.
122: 88-104. phenotype: Like sc3 but more extreme. Viability
Serebrovsky and Dubinin, 1930, J. Heredity 21: low. RK2A.
259-65 (fig.)« cytology: Associated with Dl(l)sc10-1 -Df(l)lBl-
phenotype: Slight sc; removes coxals, ocellars, first 2;1B2-14, which is superimposed on In(l)ac3 =
and second orbitals, postverticals, and anterior ln(l)lB2-3;lB13-Cl and is therefore deficient for
notopleurals. Scutellars and sternitals not af- only band 1B2 (Schultz).
fected. RK1. : see ac3
other information: No inversion. *sc?2
discoverer: Shapiro, 1929.
origin: X ray induced in wa. references: 1930, Zh. Eksperim. Biol. 6: 347-64.
discoverer: Dubinin, 1929. phenotype: First and second orbitals reduced or ab-
references: 1930, Zh. Eksperim. Biol. 6: 300—24. sent; other head bristles, posterior scutellars, and
Serebrovsky and Dubinin, 1930, J. Heredity 21: coxals also affected. Also shows achaete effect.
259-65 (fig.). Viability of homozygous female low. RK2.
Dubinin, 1933, J. Genet. 27: 443-64. *5C13
phenotype: Like sc, but anterior notopleurals not af- origin: X-ray-induced derivative of sc (induced
fected. s c 7 tends to suppress expression of h simultaneously with ac*).
(Steinberg, 1942, DIS 16: 68). RK1A. discoverer: Dubinin, 1929.
cytology: Associated with In(l)sc7 = ln(l)lB4- references: 1930, Zh. Eksperim. Biol. 6: 300-24.
6;SD3-6. 1932, J. Genet. 26: 37-58.
other information: w° separable from sc7 by ex- 1933, J. Genet. 27: 443-64.
change in triploid female. phenotype: Like sc, but scutellars invariably absent
and ocellars, postverticals, and first and second
origin: X ray induced. orbitals less frequent. Anterior and posterior dor-
discoverer: Sidorov, 1929. socentrals also absent, as are thoracic hairs be-
references: 1931, Zh. Eksperim. Biol. 7: 28—40. cause of ac*. Viability low. RK2.
Noujdin, 1935, Zool. Zh. (Moscow) 14: 317-52. *SC15
Sidorov, 1936, Biol. Zh. (Moscow) 5: 3-26. origin: X ray induced,
phenotype: Slight sc; supra-alars, sternopleurals, or discoverer: Muller.
other bristles sometimes affected. Extra bristles synonym: scutex.
may be present. Shows Hw effect and may be rec- references: Patterson and Muller, 1930, Genetics
ognized in heterozygote, homozygote, or male by 15: 495-577 (fig.).
presence of one or more hairs on anterior meso- Dubinin, 1933, J. Genet. 27: 443-64 (fig.),
pleural region. The Hw effect interacts strongly phenotype: Originally a lie lie to sc and sernilethal in
with h to produce extremely hairy wings (Steinberg, male. Subsequently allelic to y, ac, and sc and
1942, DIS 16: 68). scs/0 male nearly lethal; sur- male lethal. Lethal form exaggerates other ac and
vivors show variegation for y and ac; lethality sup- ac alleies in heterozygote. RK2A.
pressed by a Y chromosome, partially suppressed cytology: Presumably associated with Di(l)®c*s;
by parts of the Y (Hess, 1962, DIS 36: 74-75; breakpoints unknown.
other information: Apparently, y + and ac + were in- *sc260.J6

serted elsewhere in the genome as in sc19 or origin: X ray induced.
s c ^ , and became separated from the left end of X, discoverer: Sutton, 1938.
and were lost. references: 1943, Genetics 28: 210-17.
phenotype: sc260'i6/sc overlaps wild type. Lethal
origin: X ray induced. homozygous and hemizygous. RK2.
discoverer: League. cytology: Salivary chromosomes normal.
references: Muller, 1935, Genetica 17: 237-52. other information: y not affected.
phenotype: Scute liar bristles absent and sternitals * s c 260-7 7
reduced. RK1A. origin: X ray induced.
cytology: Associated with T(l;2)sc*9 = T(1;2)1B1- discoverer: Sutton, 39d.
2;1B4'7,25-26. references: 1943, Genetics 28: 210—17.
SC29 phenotype: Male and homozygous female viable and
discoverer: Agol, 1930. fertile. RK2A.
phenotype: Similar to sc7. Viable and fertile. cytology: Associated with T(l;2)sc260-17 =
RK2A. T(1;2)1B2-3;31C.
cytology: Associated with In(l)sc29 = *sc260.J8
In(l)lB;13A2-5 (Raffel). origin: X ray induced.
other information: Genetically, left break of discoverer: Sutton, 39d.
In(l)sc^9 is to the right of l(l)sc; the right break references: 1943, Genetics 28: 210—17.
is between g and / (Muller). phenotype: Male sterile. RK2A.
$c49c cytology: Associated with T(l;2;3)sc^60-18 -
origin: Induced by P 3 2 simultaneously with Hw49c. T(1;2)1A6".B1;41D-E + T(l;3)7A2-Bl;80C
discoverer: E. B. Lewis, 1965. * S C 260-20
other information: Overlooked at the time Hw* ?c origin: X ray induced.
was described. Possibly of subsequent sponta- discoverer: Sutton, 39e.
neous origin. references: 1943, Genetics 28: 210—17.
phenotype: Male and homozygous female viable and
origin: Spontaneous in raa v; arose simultaneously fertile. RK2A.
with su(s)S2<:, cytology: Associated with T(l;3)sc260-20 =
discoverer: Green, 52c. T(l;3)lA8-Bl;61Al-2.
references: 1952, DIS 26: 63. sc260.22
phenotype: Not described. RK1A (ranked by Green). origin: X ray induced.
other information: Association with In(l)scSic discoverer: Sutton, 39f.
(breakpoints unknown) inferred, since sc52c has references: 1943, Genetics 28: 210—17.
been inseparable from raa v. phenotype: Both sexes viable and fertile. RK2A.
cytology: Associated with In(l)sc^60"^2 =ln(l)lB2-
origin: X-ray-induced derivative of sc6. 3;lE2-3.
discoverer Goldat. *sc260.23
references: 1936, Biol, Zh. (Moscow) 5: 803—12. origin: X ray induced.
cytology: Associated with In(l)ac90 =ln(l)lB4- discoverer: Sutton, 1939.
7;1D2-E1. references: 1943, Genetics 28: 210—17.
phenotype: Both sexes viable. RK3A.
origin: X-ray-induced derivative of sc*. cytology: Associated with T(l;?)sc^0-23 -
discoverer: Goldat.
references: 1936, Biol. Zh. (Moscow) 5: 803-12. sc260-25
cytology: Associated with T(l;2)aci15 » T(1;2)1A6- origin: X ray induced.
B1;25F. discoverer: Sutton, 39k.
SC26C14 references: 1940, Genetics 25: 628—35.
origin: X ray induced. phenotype: Variegates for y, ac and probably 1(1 )J1,
discoverer: Sutton, 39b. but not svr, more extreme than s c ^ . Homozygous
references: 1943, Genetics 28: 210-17. lethal. RK2A.
phenotype: Both sexes viable and fertile. RK2A. cytology: Associated with In(lLR)Bc^°-25 =
cytology: Associated with Io(l}sc260-14 = in(i)|B2- In(lLR)lB2-3.
3;11D3S, *sc26Q-26
SC26Q-JS
origin: X ray induced, discoverer: Sutton, 391.
discoverer: Demerec, 381. references: 1943, Genetics 28: 210—17.
references: Sutton, 1943, Genetics 28: 210—17. phenotype: Viability reduced in male. Male fertile.
phenotype: Male sterile. Viability reduced. RK2A. RK2A.
cytology: Associated with T(l;3)sc26&-15 = cytology: Associated with TXl;2)uc260-36 »
T(1;3)1B4-5;71C-D. T(l;2)lB4-5;41F2~3;58B2-3 + ln(2LR)27D2-3;41A.
MUTATIONS 221
*sc260-2 7 references: Muller, 1932, Proc. Intern. Congr.

origin: X ray induced. Genet., 6th. Vol. 1: 225.
discoverer: Sutton, 391. Muller, Prokofyeva, and Raffel, 1935, Nature 135:
references: 1943, Genetics 28: 210—17. 253-55.
phenotype: Male viable but sterile. RK2A. cytology: Associated with InfiyscJ1 =In(l)lA4-
cytology: Associated with T(l;2)sc260'27 = 5;lB4-5.
T(1;2)1A8-B1 ;15E;19F;33-34;57B-C.
*sc260-29 origin: X ray induced.
origin: X ray induced. discoverer: Jacobs-Muller.
discoverer: Sutton, 40a. references: Muller, 1932, Proc. Intern. Congr.
references: 1943, Genetics 28: 210-17. Genet., 6th. Vol. 1: 225.
phenotype: Male viable but sterile. RK2A. 1934, DIS 2: 60.
cytology: Associated with T(l;2;3)sc260-29 = phenotype: Scute and achete effects. RK3A.
T(1;2;3)1A6-B1;22A-B;34A-B;75C-E. other information: Associated with T(l;3)scJ4 =
*sc^: scute of Ago/ T(1;3)1B;3A3-C2;61A.
discoverer: Agol. *scK: scute of Krivshenko
references: 1936, DIS 5: 7. discoverer: Krivshenko.
phenotype: Similar to sc. Viability low. RK2A. references: Agol, 1936, DIS 5: 7.
other information: Associated with In(l)scA; break- phenotype: Similar to sc, but semilethal in male and
points unknown. lethal in homozygous female. RK2A.
*scB 1: scute of Brande cytology: Associated with T(l;3)scK; breakpoints
origin: X ray induced in In(l)y* = In(l)lA8- unknown.
Bl;18A3-4. SCK3
discoverer: Brande, 37g. origin: X ray induced.
phenotype: Similar to sc. Viability Good. RK2A. discoverer: Krivshenko, 53j29.
scD1: scute of Dobzhansky references: 1959, DIS 33: 95-96.
origin: X ray induced simultaneously with a y. cytology: Associated with T(l;3)scK3 = T(1;3)1B2-
discoverer: Dobzhansky, 1930. 3;61Al-2.
references: 1935, DIS 3: 16. S C L3; scute of Levy
Morgan, Bridges, and Schultz, 1935, Carnegie Inst. discoverer Levy, 1932.
Wash. Year Book 34: 290. phenotype: In addition to scute, it has a spoonlike
phenotype: Weak sc. RK2. wing caused by a mutation to the right of sc.
cytology: Salivary chromosomes apparently normal Viable. RK2.
(Schultz). 5CL8
SCO2 discoverer: Levy, 1932.
origin: Spontaneous in y. references: Muller, Raffel, Gershenson, and
discoverer: Dobzhansky, 1931. Prokofyeva-Belgovskaya, 1937, Genetics 22: 87—
references: 1935, DIS 3: 16. 93.
phenotype: Slight sc. RK2. Muller and Raffel, 1938, Genetics 23: 160.
scFah: scute of Fahmy Raffel and Muller, 1940, Genetics 25: 541-83.
origin: Induced by DL-p-NN-di(2-chloroethyl)amino- phenotype: Similar to sc4 but more extreme. Homo-
phenylalanine (CB. 3007). zygous female sterile. RK2A.
discoverer: Fahmy, 1954. cytology: Associated with In(l)scL8 =In(l)lB3~
references: 1958, DIS 32: 74. 4;20B~C.
phenotype: Bristles, principally orbitals, verticals, * J C P I ; scute of Pan shin
postverticals, and ocellars missing. Scutellars and discoverer: Panshin, 1934.
postdorsocentrals left nearly intact. Male viable phenotype: Like sc. RK2A.
and fertile; female homozygous lethal. scFah/sc cytology: Associated with T(l;2;3)scP1; break-
has only occasional absence of postvertical or points unknown.
ocellar bristles. RK2A. scs1: scute of Sinitskaya
cytology: Associated with D^l)scFah =Df(l)lA8- discoverer: Sinitskaya, 34c.
Bl;lB2-3. references: Muller, 1935, DIS 3: 50.
scH: scute of Hackett Muller and Raffel, 1938, Genetics 23: 160.
origin: Gamma ray induced. Raffel and Muller, 1940, Genetics 25: 541-83.
discoverer Hackett, 46a. Crew and Lamy, 1940, J. Genet. 39: 273—83.
references: Muller and Valencia, 1947, DIS 21: 70. phenotype: Rather extreme «c allele; slight Hw ef-
phenotype: Similar to sc but more extreme. RK2A. fect; hairs often removed from abdomen and wings.
cytology: Associated with T(1;4)BC» « T(1;4)1B4- Homozygous female sterile and low in viability.
C3; 101 -102. Male fertile and fairly viable. RK2A.
SC JJ : scute of Jacobs-Muller cytolo-gy: Associated with In(l)8csi * ln{l)lB3-
origin: X ray induced simultaneously with l(l)Jl. 4;2®B-D1. Warn found in combination with In(l)S *
discoverer, Jacobs-Muller. In(l)6Al-3;lQFlQ-llAl.
*Scar: Scarred
discoverer: Sinitskaya, 1934. location: 2- or 3- (rearrangement).
phenotype: Similar to sc?. RK1A. origin: X ray induced.
cytology: Associated with T(l;2)scs2 = T(1;2)1B4- discoverer: Yu, 48h.
7;6QC-E. references: 1949, DIS 23: 65.
*scSo: scute of Sytko phenotype: Eyes elliptical with indented, glassy
discoverer: Sytko. posterior margin. Wings spread at 45° from body
references: Agol, 1936, DIS 5: 7. from body axis. Enhanced at 28°C. Homozygous
phenotype: Like sc; viability of homozygous female lethal. RK1A.
low. RK2. cytology: Associated with T(2;3)Scar =
s c v ' : scute of Valencia T(2;3)27E;95A + In(3)91F;96A.
origin: Gamma ray induced. scarlet: see st
discoverer: J. I. Valencia, 46h23. scarp: see scrp
synonym: scvi, Inp (Inp signifies a pericentric in- Scarred: see Scar
version). *sch: slender chaetae
references: Muller and Valencia, 1947, DIS 21: 69- location: 1-21.1.
70. origin: Induced by D-p-NN-di-(2-chlorethyl)amino-
phenotype: Extreme scute and achaete. Viability phenylalanine (CB. 3026).
low. RK2A. discoverer: Fahmy, 1955.
cytology: Associated with In(lLR)scVl = references: 1959, DIS 33: 90.
In(lLR)lA8-C3. phenotype: Bristles thin and slightly shortened.
scV2 Eyes slightly smaller and brownish. Body small.
origin: Gamma ray induced. RK2.
discoverer: J. I. Valencia, 46h23. *Scn: Scutenick
references: Muller and Valencia, 1947, DIS 21: 70. location: 4- [included in Df(4)M].
phenotype: Both achaete and scute variably af- discoverer: Padoa, 1931.
fected. Some tendency for extra or twin bristles. references: Bridges, 1935, Biol. Zh. (Moscow) 4:
Abdominal bristles markedly fewer both dorsally 401-20.
and ventrally. Male and homozygous female viable Padoa, 1938, Monit. Zool. Ital. 49: 279-84.
and fertile. RK2A. phenotype: Scutellum shortened, with nick at poste-
cytology: Associated with In(l)scV2 = In(l)lB2- rior edge; scutellar bristles missing. Ocelli re-
3;20B-F. duced, with disturbed hairs and bristles. One or
both eyes often small or absent. All characters
Sc: Scotched eye overlap wild type. Eye effect is strongest at 19°C
location: 1-4.5 (about 4 or 5). but other effects weaker. Scutellum effect best at
origin: X ray induced. 28° but eyes normal. Homozygous lethal. RK2.
discoverer. Muller. cytology: Placed in salivary chromosome region
references: 1946, DIS 20: 67. 101E through 102B16, on basis of its inclusion in
phenotype: Ommatidia disarranged near posterior Df(4)M = Df(4)101E-F;102B6-17.
margin of eye. Resembles spaCat. Good viability
and fertility in heterozygous female. Male lethal. Sco: Scutoid
RK2. location: 2-51.0.
Sd: see Scp origin: X ray induced.
sc-Dp: see Dp(l;f)100 discoverer: Krivshenko, 56115.
sc-Inh-3: see Sufsc) references: 1959, DIS 33: 96.
sea: scabrous 1960, DIS 34: 55.
location: 2-66.7. phenotype: Scutellar bristles usually absent; one or
ori g i n: Spontaneous. both postscutellar bristles sometimes present but
discoverer: Ives, 34j2. are shorter and thinner than normal. Ocellar and
references: 1935, DIS 4: 10. humeral bristles often absent. Homozygous lethal.
phenotype: Eyes large and rough. Ocellar bristles RK1.
85 percent absent at 25°C and 10 percent absent at cytology: Salivary chromosomes apparently normal.
18°. Postverticals occasionally missing. Bristle Placed in region between 34E5 and 35D1, on basis
effect more extreme in male at 21° and in female at of its inclusion in Df(2L)64j = Df(2L)34E5-
28°. Most bristles subject to twinning. May be F1;35C3-D1 (E. H. Grell).
extra rows of acrostichal hairs. RK1. Scoop: see Scp
cytology: Placed in region 49D1-3, on basis of its scooped: see scp
inclusion in Df(2R)v&* - Dt(2R)49C2-Dl;50A2-3 but scooped thickvein: see set
not inDfC2R)vgB = Df(2R)49D3-4;50A2-3 (Morgan, Scotched eye: see Sc
Bridges, and Schultz, 1938, Carnegie Last. Wash. scp; scooped
Year Book 37: 205). location: 1-19.3.
scalloped: see sd discoverer Muller, 1926.
MUTATIONS 223
phenotype: Wings turn up slightly; classification origin: Spontaneous.

fairly reliable. RK2. discoverer: Hannah, 53b.
cytology: Placed between 6A3-4 and 6F10-11 references: Hannah and Stromnaes, 1955, DIS 29:
(Demerec, Kaufmann, Fano, Sutton, and Sansome, 121-23.
1942, Carnegie Inst. Wash. Year Book 41: 191). Hannah, 1958, Genetics 43: 878-905 (fig.).
*Scp: Scoop phenotype: Sex combs may be present on all six
location: 3- (not located). legs of male. At least one extra sex comb present
origin: Induced by 2-chloroethyl methanesulfonate in 75—90 percent of males. Third pair of legs less"
(CB. 1506). often affected than second. Scx/Pc more extreme
discoverer: Reddi. than Scx/+; male usually has large sex comb on all
synonym: Sc (preoccupied). six legs. Lethal homozygous and when heterozy-
references: 1963, DIS 37: 53. gous with Antp49 and Antpso. Expression of Scx/+
phenotype: Wing size reduced; proximal one-third of enhanced in males that are also heterozygous for
wing compressed laterally; distal two-thirds spoon- bx, bxd, and Ubx. Furthermore, with bx and bxd,
like. Three furrows run length of wing and surface the enhancement is greater if mutants are in cou-
is wrinkled. Abdomen cylindrical and untapered pling rather than repulsion, but compounds with
posteriorly. Pigmented abdominal bands darkened. Ubx show no phase difference (Hannah-Alava,
Excellent viability and fertility. RK3. 1964, Z. Vererbungslehre 95: 1-9). RK2.
*scr: scruH
location: 1-22.0.
discoverer: Neel, 41b22.
1942, Genetics 27: 532.
phenotype: Hairs and bristles missing or doubled,
and deranged. Eyes small and rough. Scute Hum
more convex than wild type. Wing margins, espe-
cially posterior, often incised. Wings occasionally
blistered. All characters variable; a few flies ap-
pear normal. RK3.
cytology: Salivary chromosomes appear normal.
scrp; scarp
location: 2-74 (to the left of c; not an allele of L). Sex: Extra sex comb
origin: Spontaneous. Third legs of Sex moles
discoverer: Hansen and Gardner, 1960. From Hannah, 1958, Genetics 43: 878-905.
1962, Genetics 47: 587-98 (fig.). sd: scalloped
phenotype: Ventral one-third of eye flattened and location: 1-51.5.
separated from dorsal two-thirds by a furrow. Pen- origin: X ray induced.
etrance 80 percent at 30°C; at 25°, eyes are wild discoverer: Gruneberg, 28j20.
type. Temperature-effective period from forty- references: 1929, Biol. Zentr. 49: 680-94.
second to sixty-eighth hour of development. RK3. 1934, DIS 2: 9.
scruff: see scr phenotype: Wing margins scalloped and veins thick-
*sct: scooped thickvein ened. Eyes slightly roughened. Does not overlap
location: 1-16.0. wild type. Waddington (1940, J. Genet. 41: 7 5 -
origin: Induced by methyl methanesulfonate (CB. 139) concluded that sd reduces size of prospective
1540). wing area during larval period and shifts plane
discoverer: Fahmy, 1956. along which wing area is folded out from imaginal
references: 1960, DIS 34: 49. bud. Scalloping visible in prepupal wing and
phenotype: Wings short and scooped; inner margin scorable in unexpended wing. RK1.
frequently incised in several places; veins thick-
ened. Eyes darker and slightly altered in shape.
Abdominal tergites slightly ridged. Male sterile;
viability about 40 percent normal. RK2.
scute: see sc
scute Inhibitor on chromosome 3: see Su(sc)
Scutenick: see Sen
scutex: see sc1 s
Scutoid: see 5co
Sex; Extra sex comb
location: 3-47 <0.3 unit to the right of Pc and to the sd: scalloped
left of p}. Edith M. Wallace, unpublished.
cytology: Placed in salivary chromosome region SD: Segregation Distorter

13B2-F17 on basis of its inclusion in Dp(l;f)A12 = location: 2-55 (near the heterochromatic-euchromatic
Dp(l;i)lB-C;13Bl-5 but not in the proximal part of junction).
the X derived from T(1;4)A4 = T(1;4)13F6- origin: Naturally occurring allele recovered near
14A1;1O2F (inferred from Patterson, 1938, Am. Madison, Wisconsin. The same or similar alleles
Naturalist 72: 193—206, also frontispiece of Texas found in populations in Baja California (Mange,
Univ. Publ. 4032). 1961, Am. Naturalist 95: 87-96), Kentucky, and
sd: see sprd Illinois (Greenberg, 1962, DIS 36: 70).
sd* discoverer: Hiraizumi.
origin: X ray induced. references: Sandier, Hiraizumi, and Sandier, 1959,
discoverer: Panshin, 33g7. Genetics 44: 233—50.
references: 1935, DIS 3: 28. phenotype: Female and homozygous male normal.
phenotype: More extreme than sd. Wings small and The majority of functional sperm of heterozygous
scalloped. Like vg at high temperatures. Crossing male, often 95 percent, carry the SD-bearing
over inhibited. RK2A. second chromosome. Meiosis and sperm develop-
cytology: Associated with In(l)sd2; breakpoints un- ment in SD/+ male are without visible abnormality;
known. it has been postulated that SD acts by directing its
,«/35 homolog into a normally nonfunctional half of the
origin: Spontaneous. sperm (Peacock and Erikson, 1965, Genetics 51:
discoverer: Hollander, 1935. 313-28). RK3.
references: 1937, DIS 8: 8. cytology: Base of 2R may be normal (Lewis, 1962,
phenotype: Like sd. RK2. DES 36: 87). All naturally occurring SD chromo-
other information: Allelism inferred from phenotype. somes contain inversions in the right arm. The in-
version present varies and is not required for SD
origin: X ray induced. activity.
discoverer: Clark, 56j. other information: SD-bearing chromosomes in nature
references: Andrew, 1959, DIS 33: 82. have St-SD: Stabilizer of SD located at the tip of
phenotype: More extreme than sd. Expression en- 2R, in the absence of which SD action is more var-
hanced by high temperature. Visible in prepupal iable (Sandier and Hiraizumi, 1960, Genetics 45:
wing buds. Interacts with Bx and in. RK1. 1269—87). There are in addition a great variety of
cytology: No gross chromosomal abnormality. modifiers of SJD activity in the genome (Sandier and
Hiraizumi, 1959, Proc. Natl. Acad. Sci. U.S. 45:
sd58d 1414-22; Sandier and Rosenfeld, 1962, Can. J.
origin: Gamma ray induced. Genet. Cytol. 4: 453—57). The SD locus is complex
discoverer: Ives, 58dl4. and consists of at least a distorter element and an
references: 1961, DIS 35: 46. element that renders chromosome 2 immune to SD
phenotype: Wings reduced to vestiges, like vg. Hal- action (Sandier and Hiraizumi, 1960, Genetics 45:
teres and bristles also like vg. ad58d/sd has 1671-89).
strap-shaped wing. RK2A. sdx: spreadex
cytology: Associated with In(l)sdS8di breakpoints location: 1- (rearrangement).
unknown. origin: X ray induced.
**</«; scalloped-sterile discoverer: Muller.
origin: X ray induced. synonym: spx (preoccupied).
discoverer Muller. references: 1965, DIS 40: 35.
references: 1946, DIS 20: 67-68. phenotype: Wings spread widely apart and often di-
phenotype; Wings divergent and slightly nicked. rected somewhat downward, as in Dichaete. Ab-
Male sterile. RK2. domen of female tends to be narrow and shrunken.
other information: Allelism inferred from position Fertility sufficient for maintaining homozygous
and phenotype. No evidence of chromosome rear- stock. RK2A.
rangement. cytology: Associated with In(l)sdx; breakpoints un-
known.
sd*P: scalloptd-spatula se; sepia
origin: X ray induced inIn(l)BcslLBC*R+dl-49. location: 3-26.0.
discoverer: R. M. Valencia, 1959. discoverer: E. M. Wallace, 13elO.
synonym: sp. references: Bridges and Morgan, 1923, Carnegie
references: 1959, DIS 33: 99. Inst. Wash. Publ. No. 327: 86 (fig.).
1965, DIS 40: 37. Sturtevant and Beadle, 1939, An Introduction to
phenotype: Wings cut at tips and along both margins. Genetics, Saunders, p. 64 (fig.).
md*P +/+ Bxr give slight nicking of wings. RK1A. phenotype: Eye color brown at eclosion, darkening
cytology: No gross rearrangement in addition to to sepia and becoming black with ag@. Pigmenta-
In(l}tscSiLfiC8R4<lt-49, but possibly a local dis- tion of ocelli normal. Chromatographically, me
turbance in pairing. eyes characterized by having no red pigment and
MUTATIONS 225
an accumulation of the yellow pigment, sepiapterin

(Hadorn and Mitchell, 1951, Proc. Natl. Acad. Sci.
U.S. 37: 650—65); other pteridines present in
greater-than-normal amounts. s e / + can be distin-
guished from +/+ in that it has more isoxanthop-
terin and other pale pteridines; the red drosopterins
are at wild-type level, so that se appears com-
pletely recessive on ordinary visual examination
(Ziegler-Gunder and Hadorn, 1958, Z. Vererbungs-
lehre 89: 235—45). Structure of the sepiapterin is
2-amino-4-oxo-6-lactyl-3,4,7,8-tetrahydropteridine Ser: Serrate
(Forrest and Nawa, 1962, Nature 196: 372-73). Edith M. Wallace, unpublished.
Eye color autonomous in se eye disks transplanted
into wild-type hosts (Beadle and Ephrussi, 1936, sexcombless: see sx
Genetics 21: 230). RK1. sf: safranin
location: 2-71.5.
discoverer Hungerford, 51 j. discoverer: Bridges 16a6.
references: Redfield, 1952, DIS 26: 68. references: Morgan, Bridges, and Sturtevant, 1925,
phenotype: Like se. RK1. Bibliog. Genet. 2: 235.
5U phenotype: Eye color soft dark brown. More easily
origin: Spontaneous, classified in male and in aged fly. Larval Mal-
discoverer: Clark, 51k. pighian tubes pale yellow; classifiable (Brehme
references: 1952, DIS 26: 60. and Demerec, 1942, Growth 6: 351—56). RK2.
phenotype: Like se. RK1. sf2
origin: Spontaneous, discoverer: Spencer, 25k.
discoverer: Andrew, 58k. synonym: bronze.
phenotype: Like se. RK1. 1935, Am. Naturalist 69: 223-38.
1937, DIS 7: 21.
origin: Spontaneous. phenotype: Like sf but possibly less extreme; 47
discoverer: Clancy, 61c. percent normal red and 98 percent normal brown
references: 1964, DIS 39: 65. pigment (Nolte, 1955, Genetics 53: 1-10). Eye
phenotype: Like se. RK1. color autonomous in larval optic disks transplanted
se-like 62: see pni into wild-type host (Beadle and Ephrussi, 1936,
sed: see Hnr3 Genetics 21: 230). Larval Malpighian tubes bright
Segregation Distorter: see SD yellow like wild type (Beadle, 1937, Genetics 22:
*semi-f: semi forked 587-611). RK2.
location: 3- (not located). sP
discoverer: Lancefiled, 18b. discoverer: Ives, 39c.
references: 1918, Am. Naturalist 52: 462—64. references: Curry, 1939, DIS 12: 45.
phenotype: Homozygotes that are also heterozygous phenotype: Like sf. RK2.
for f have slightly forked bristles. RK3.
sep: separated origin: From heat-treated larvae.
location: 3- (rearrangement). discoverer. Ives, 32e28.
discoverer; Muller. synonym: dark eye (1934, DIS 1: 33).
phenotype: Most of posterior cross vein absent, one- references: Plough and Ives, 1935, Genetics 20:
third usually remaining attached to vein L5. RK2A. 42-69.
cytology: Associated with ln(3LR)®ep = phenotype: Like sf. RK2.
In(3LR)65E;85E (Lewis, 1951, DIS 25: 108-9). *sf-3: safranin in chromosome 3
sepia: see se location: 3- (not located).
sepiaoid: see HnT3 discoverer: Bridges, 15al5.
Sen Serrate references: Bridges and Morgan, 1923, Carnegie
location: 3-92.5 (to the right of Pf). Inst. Wash. Publ. No. 327: 126.
origin: Spontaneous. phenotype: Eye color dull brown. RK3.
discoverer: Spencer, 3517. *sfc; stiff chaetae
references: Curry, 1939, DIS 12: 46. location: 1-3.2.
phenotype: Wings notched at tip; deepest notch at origin: Induced by D-p-NN-di-(2-chloroetnyl)aminO"
second posterior cell. In tripioids, one dose of Ser phenylalanine (CB. 3026).
overlaps wild type. Homozygous lethal. RK1. discoverer: Fahmy, 1955.
references: 1958, DIS 32: 74. references: 1949, DIS 23: 61-62.
phenotype: Bristles short and stiff; occasionally phenotype: Like Sh. RK1.
one missing. Fertility and viability good. RK2.
other information: One allele induced by CB. 1592. origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
*sg: shortened wing phenylalanine (CB. 3025).
location: 3- (left arm). discoverer: Fahmy, 1954.
origin: Spontaneous. synonym: Shw: Shaker downheld.
discoverer: Herskowitz, 47118. references: 1959, DIS 33: 90.
references: 1949, DIS 23: 57. phenotype: Fly quivers when etherized. Wings fre-
phenotype: Wings abnormal at base; veins inter- quently droop at sides. Thorax often dented, par-
rupted, missing, or thickened. Many flies have ticularly near anterior border. Homozygote viable
short, rounded wings that curve upward slightly. and fertile. RK1.
RK3. other information: One allele each induced by X rays
*sge: shifted genitals and CB. 1592. Two alleles induced by CB. 1540.
location: 1-48.4. shaven: see sv
origin: Induced by 2-chloroethyl methanesulfonate *shb: shortened bristles
(CB. 1506). location: 1-39.0.
discoverer: Fahmy, 1956. origin: Induced by S-2-chloroethylcysteine (CB.
references: 1958, DIS 32: 74. 1592).
phenotype: Male genitalia and anal plates rotated to discoverer: Fahmy, 1957.
various degrees (up to 90°). Wings slightly diver- references: 1959, DIS 33: 90.
gent and drooping, occasionally one outheld. Eyes phenotype: Bristles slightly short and thin. Wings
slightly dark. Male sterile. Viability about 70 per- broad, often convex or concave. Fly somewhat
cent normal. RK2. large. Male fertile; viability about 50 percent wild
*sh: short winged type. Female sterile. RK3.
location: 3-56. shd: see spl
origin: Spontaneous, *she: sherry
discoverer: Bridges, 23d3. location: 3-0.
synonym: short wing. origin: Spontaneous.
references: Morgan, Bridges, and Sturtevant, 1925, discoverer: Kaliss, 36al3.
Bibliog. Genet. 2: 235. references: 1937, DIS 8: 9.
1935, DIS 3: 16. phenotype: Eye color sherry. Sterile inter se but
phenotype: Wings small; similar to dy. RK2. both sexes crossfertile. RK3.
*sh-5: short-5 *shf: shifted
origin: Spontaneous. discoverer: Bridges, 13a.
discoverer: Spencer, 26j. references: Morgan and Bridges, 1916, Carnegie
references: 1934, DIS 1: 35. Inst. Wash. Publ. No. 237: 63.
1935, Am. Naturalist 69: 223-38. phenotype: Vein L3 fails to reach wing margin and
phenotype: Wing veins L5 and L3 short; do not is shifted toward L4. Anterior crossvein usually
reach wing margin. Expression variable; overlaps lacking. Wings divergent. Postscutellar bristles
wild type. RK3. small and erect. Body small. Viability 60 percent
Sh: Shaker of wild type. Female often sterile. RK2.
location: 1-57.7. cytology: Placed between 6A3 and 6F11 based on
origin: X ray induced. deficiency analysis using shf2 (Demerec,
discoverer: Catsch. Kaufmann, Fano, Sutton, and Sansome, 1942,
references: 1944, Z. Induktive Abstammungs- Carnegie Inst. Wash. Year Book 41: 191).
Vererbungslehre 82: 64-66.
phenotype: Causes spasmodic tremor of legs and origin: X ray induced.
abdomen in moderately etherized male and homozy- discoverer: Oliver, 29j29.
gous female; very little effect in deeply anaesthe- references: 1935, DIS 3: 28.
tized fly. Heterozygous female shows reduced 1935, DIS 4: 10.
effect, with shaking confined to forelegs. Expres- phenotype: Veins closer together than in wild type.
sion and viability excellent. RK1. L3 and L4 tend to fuse near anterior crossvein;
Sh2 anterior crossvein shortened, knotted, or absent.
origin; X ray induced. Phenotypic effect visible in prepupal wing bud, the
discoverer: Novitski, 48k. two longitudinal veins diverging at a smaller than
references: 1949, DIS 23: 61-62. normal angle [Waddington, 1940, J. Genet. 41: 75—
phenotype: Like Sh but male lethal. RK2. 139 (fig.)j. Eyes sometimes slightly rough. Scu-
Sh* tellar bristles often absent. Scutellum short. Wings
origin: X ray induced. narrow and often warped downward. Fertility and
discoverer: Novitski, 49b. viability good. RK2.
MUTATIONS 227
short-bristte: see ml
origin: Spontaneous. shortened wing: see sg
discoverer: Curry, 37d26. shortened bristles: see shb
phenotype: Like shf2 but more extreme. Viability shortened veins: see svs
about 70 percent wild type. Frequently infertile. shorter bristles: see sbt
RK2. shorter legs: see shl
shovel: see sho
*shp: shrimp
location: 1-47.5.
phenotype: Small fly. Eclosion delayed. Male via-
bility about 30 percent wild type. Both sexes fer-
tile. RK3.
shfh shifted-3 shr: shrunken
From Bridges and Brehme, 1944, Carnegie Inst. Wash. location: 2-2.3.
Publ. No. 552: 173. discoverer: Bridges.
phenotype: Body small and dark. Viability and fer-
s/if ov : shifted-oval tility good. Overlaps wild type unless combined
origin: Induced by P 3 2 . with abb, where mutual enhancement occurs. RK3.
discoverer: Bateman, 1950. cytology: Placed between 22A3 and 22B1, on basis
references: 1950, DIS 24: 55. of its inclusion in Df(2L)S2 = Df(2L)21C6-
phenotype: Eyes rough and narrow. First basal wing Dl;22A6~Bl but not in Df(2L)S5 = Di(2L)21C2-
cell absent because L3 and L4 veins close. Wings 3;22A3-4 (Lewis, 1945, Genetics 30: 137-66).
narrow and pointed. Viability and fertility low. shrimp: see shp
RK2. shrunken: see shr
other information: On basis of phenotype and posi- shrunken-3: see wz
tion, could be an allele of either ov or shf or both; *sht: short tarsi
not tested. location: 1-20.9.
shifted genitals: see sge origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
*shl: shorter legs phenylalanine (CB. 3007).
location: 1-36.3. discoverer: Fahmy, 1953.
origin: Induced by 2-fluoroethyl methanesulfonate references: 1959, DIS 33: 90.
(CB. 1522). phenotype: Legs extremely short; reduction in
discoverer: Fahmy, 1957. length most pronounced in metatarsal and tarsal
references: 1959, DIS 33: 90. regions. Some tarsi fused; others absent. Bristles
phenotype: Small fly with short legs. Male viability thin and short. Adult short lived. RK3.
and fertility low. RK3. shv: short vein
shm: short macros location: 2-3.8 (between ast and ho).
origin: Induced by trie thy lenemelamine (CB. 1246). discoverer: Pope, 1947.
discoverer. Fahmy, 1953. references: Lewis, 1947, DIS 21: 69.
references: 1959, DIS 33: 90. phenotype: Veins L2 and L4 do not reach wing
phenotype: Bristles short and stiff. Eclosion de- margin. RK1.
layed. Male sterile and viability reduced. RK2. shV: see avs
*sho: shovel Shw: seeS/i*
location: 2- (not located). *S/: Ski
origin: Spontaneous in In(2L)t. location: 2-36.
discoverer: GoodSmith, 49k. discoverer Clausen, 1511.
references: Ives, 1952, DIS 26: 65. references: Clausen and Collins, 1922, Genetics 7:
phenotype: Wings short and rounded. Viability good. 385-426.
RK2A. Bridges and Morgan, 1923, Carnegie Inst. Wash.
short bristle: see stb Publ. No. 327: 149 (fig.).
short macros: see shm phenotype: Homozygous or heterozygous Si com-
short tarsi: see sht bined with homozygous si-3 produces wings with
short vein: see shv turned up tips. Double homozygote has also a
short wing: see sw crimped costal vein. Other genotypes wild type.
short wing: see sh RK3.
short winged: see sh *si-3: ski-3
short-5: see s/i-5 location: 3-46.5.
discoverer: Clausen, 1511. phenotype: Dark streak extends down middle of

references: Clausen and Collins, 1922, Genetics 7: thorax from neck to tip of scutellum. Wings may
385-426. diverge and droop. Overlaps wild type. Enhanced
Bridges and Morgan, 1923, Carnegie Inst. Wash. by b or e s . Homozygous lethal. RK2.
Publ. No. 327: 149. cytology: Salivary chromosomes apparently normal
phenotype: si-3/si-3 fly has upturned wingtips when (Bridges).
homozygous or heterozygous for Si, otherwise Ski: see Si
normal. RK3. ski-3: see si-3
side wings: see s/w Skilike: see Si I
*Sit: Skilike si: small wing
discoverer: Goldschmidt. origin: Spontaneous.
references: 1947, J. Exptl. Zool. 104: 216. discoverer: Bridges, 15121.
phenotype: Wings turned up at tips. Semidominant. phenotype: Wings about 80 percent normal length,
Poor viability. RK3. straight edged, and blunt tipped. Crossveins rather
other information: Not an allele of Si. close. Eyes large and slightly rough. RK2.
silver: see svr Sl2
silver tips: see stp origin: X ray induced.
sine ocu/is: see so discoverer: Dobzhansky, 31b3.
singed: see sn references: Sivertzev-Dobzhansky and Dobzhansky,
*siw: side wings 1933, Genetics 18: 173-92.
location: 1-58.5. phenotype: Similar to si but possibly more extreme.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- RK2.
phenylalanine (CB. 3025). *sl34
discoverer: Fahmy, 1955. origin: Pound among progeny of cold-treated male.
references: 1958, DIS 32: 74. discoverer: Gottschewski, 1934.
phenotype: Wings rotated on long axis so that inner phenotype: Wings like si, but eyes normal. RK2.
margin is higher than costal margin. Male sterile; *SI: Splotched
viability about 50 percent wild type. RK2. location: 1-56.9 (to the right of f).
phenotype: Wing hairs disarranged in small patches.
Male infertile. Viability excellent. RK1.
sla: slimma
location: 1-48.6.
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
phenotype: Fly slim with very narrow abdomen.
Body length normal. Eclosion delayed slightly.
Wings curve slightly, sla/sib, and sla/sld wild
type. Male fertile and viable. Female sterile;
viability about 50 percent wild type. RK3.
other information: Two alleles each induced by CB.
3007 and CB. 3025.
*slb: slim body
location: 1-45.3.
origin: Induced by ethyl methanesulfonate (CB.
1528).
Sk: Streak references: 1958, DIS 32: 74.
From Bridges and Morgan, 1919, Carnegie Inst. Wash. phenotype: Body narrow but of normal length.
Pubi. No. 278: 216. stb/ala and alb/aid wild type. Viability and fer-
tility good in both sexes. RK3.
Sk: Streak sic: slim chaetae
location: 2-16.0. location: 1-3.6.
origin: Spontaneous, origin: Induced by L-p~NN-di-(2-chloroethyl)amino-
discoverer: Bridges, 12k27. phenylalanine (CB. 3025).
references: Bridges and Morgan, 1919, Carnegie discoverer: Fahmy, 1954.
Inst. Wash. Publ. No. 278: 222 (fig.). references: 1959, DIS 33: 90-91.
MUTATIONS 229
phenotype: Bristles thin and short. Inner wing origin: Induced by 2-chloroethyl methanesulfonate
margins occasionally incised. Both sexes viable (CB. 1506).
and fertile. RK1. discoverer: Fahmy, 1956.
sld: slender references: 1958, DIS 32: 75.
location: 1-50.1. phenotype: Wings usually slightly upheld or spread.
origin: Induced by p-NN-di-(2-chloroethyl)amino- Viability and fertility good. RK3.
phenylethylamine (CB. 3034). sm: smooth
discoverer. Fahmy, 1957. location: 2-91.5.
phenotype: Fly rather small and slim with narrow discoverer: Bridges, 35cl4.
abdomen, sld/sla and sld/slb wild type. Male fer- phenotype: Abdomen partially denuded of bristles
tile but shows delayed eclosion and reduced via- and shrunken. Wings usually warped and semi-
bility. Female very inviable. RK3. erect. Acrostichal hairs disarranged. Tendency for
other information: One allele induced by CB. 3025. erect postcutellars. Male genitalia often dis-
turbed. Anal protuberance of female bent down.
*sldPfa: slender-pointed abdomen Viability 30 percent wild type. Both sexes entirely
origin: Induced by 2-chloroethyl methanesulfonate sterile. RK2.
(CB. 1506). sm: see smk
discoverer: Fahmy, 1956. sma: smaller
synonym: pta. location: 1-29.9.
references: 1959, DIS 33: 88. origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
phenotype: Fly small, with narrowed abdomen and phenylalanine (CB. 3025).
slightly altered eye and wing shape. Male sterile; discoverer: Fahmy, 1953.
viability about 25 percent wild type. RK3. references: 1958, DIS 32: 75.
slender chaetae: see sc/i phenotype: Body small. Eye color frequently dark.
slight: see sit Viability and fertility good. RK2.
slim: see slm other information: One allele each induced by CB.
slim body: see sib 1528, CB. 1540, CB. 2511, CB. 3007, CB. 3025,
slim bristle: see smb CB. 3026, CB. 3034. Two alleles induced by CB.
slim chaetae: see sic 1414.
slimma: see sla small: see sml
slimmer abdomen: see sin small body: see sby
slm: slim small body 62: see srb
location: 1-33.7. small bristle: see sbr
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- small eye: see oss
phenylalanine (CB. 3025). small narrow: see smn
discoverer: Fahmy, 1955. small pallid: see smp
references: 1958, DIS 32: 75. small round', see srd
phenotype: Small fly with narrow abdomen. Viability small thin: see sth
and fertility good. RK3. small thorax: see smt
other information: One allele induced by CB. 1506. small tvmoroid: see stu
*sln: slimmer abdomen small wing: see si
location: 1-53.5. smaller: see sma
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- smaller body: see srb
phenylalanine (CB. 3025). smaller eye: see sme
discoverer: Fahmy, 1953. smaller thinner: see smh
references: 1959, DIS 33: 91. sma I hid: see smd
phenotype: Rather small fly with narrow abdomen. *smb: slim bristle
Occasionally, wings slightly upheld and eyes location: 1-23.1.
small or misshapen. Male infertile; viability about origin: Induced by ethyl methanesulfonate (CB.
15 percent wild type. Female sterile. RK3. 1528).
slope wing: see s/w discoverer Fahmy, 1956.
sit: slight references: 1959, DIS 33: 91.
location: 2-106.3. phenotype: Bristles thin and rather short. Male
origin: Spontaneous. viable and fertile; female sterile. RK2.
discoverer. Curry, 39b20. other information: One allele induced by CB. 1540.
references: 1939, DIS 12: 45. smd: small old
phenotype: Fly small. Bristles short and thin. En- location: 1-61.1.
hances px. Viability and fertility good. RK3. origin: Induced by DL-p-NN-di-(2-chloeoethyI)amino-
slV: see avr phenylalanine (CB. 3007).
*slw: slope wing discoverer Fahmy, 1954.
phenotype: Rather small body. Eyes frequently *smp: small pallid

dark. Viability and fertility good. RK2. location: 1-25.6.
cyfology: Placed in salivary chromosome region origin: X ray induced.
18A4-18B8, on basis of its inclusion within the discoverer: Fahmy, 1954.
deficiency resulting from recombining left end of references: 1959, DIS 33: 91.
In(l)y4 - In(l)lA8-Bl;18A3-4 with right end of phenotype: Fly quite small and lightly pigmented.
In(l)sc9 =In(l)lB2-3;18B8-9 (Norton and Valencia, Bristles slightly thin. Occasional eye misshapen.
1965, DIS 40: 40). Male viable and fertile. Female sterile. RK2.
other information: One allele each induced by CB. *s/nf: small thorax
1414, CB. 1540, CB. 1592, and CB. 3007. Two location: 1-51.9.
alleles each induced by CB. 1506 and CB. 1528. origin*. Induced by L-p-NN-di-(2-chloroethyl)amino-
Seven alleles induced by CB. 3025 and 10 by X phenylalanine (CB. 3025).
rays. discoverer: Fahmy, 1953.
*sme: smaller eye references: 1958, DIS 32: 75.
location: 1-68.9. phenotype: Thorax and head small. Wings corre-
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- spondingly short but of normal width and frequently
phenylalanine (CB. 3025). wavy. Both sexes fertile. Viability about 50 per-
discoverer. Fahmy, 1955. cent wild type. RK2.
phenotype: Small fly with small, round, and slightly
dark eyes. Wings occasionally diverge. Male
sterile; viability about 50 percent wild type. RK2.
*smh: smaller thinner
location: 1-1.5.
origin: Induced by methyl methanesulfonate (CB.
1540).
discoverer Fahmy, 1956.
phenotype: Rather small fly with thin bristles. Both
sexes viable and fertile. RK2.
*smk: smoky
location: 2-58.6.
discoverer: Edmonds on and Meyer, 49d.
synonym: sm (preoccupied).
phenotype: Body color dark, especially along sides
of thorax. Similar to e* but somewhat lighter. At
27°C, female sterile and male fertile; at 17°, both
sexes fertile. Viability and classification good.
RK2. sn: singed
*sml: small From Mohr, 1922, Z. Intuktive Abstammungs-
location: 1-25. Vererbungslehre 28: 1-22.
origin: Induced by P33.
discoverer: Bateman, 1950. sn: singed
phenotype: Body small; wings short; eyes small, origin: Spontaneous.
rough, and bulging. Thoracic hairs irregular. Eclo- discoverer: Mohr, 18j5.
sion delayed. 10 percent normal viability. RK3. references: 1922, Z. Induktive Abstammungs-
Vererbungslehre 28: 1—22 (fig.).
*smn: small narrow Bender, I960, Genetics 45: 867-83 (fig.).
location: 1-45.7. phenotype: Bristles twisted and shortened. Hairs
origin: Induced by 2-chloroethyl methanesulfonate wavy. Female sterile. Bender (1960) finds that, in
(CB. 1506). ovaries of sterile sn female, vitellogenesis is re-
discoverer Fahmy, 1955. tarded and eggs never develop beyond stage 13.
references: 1959, DIS 33: 91. Mohr (1922) reported that eggs laid are short and
phenotype: Fly weak and inviable; usually dies have flattened filaments, sn heterozygous with
within 48 hr of eelosion. Wings frequently upheld fertile alleles is fertile; sn heterozygous with
slightly. Abdomen narrow. RK3. sterile alleles of sn is sterile. RK1.
smoky, see smk cytology: Demerec and Sutton place locus between
smooth: see sm 7C4-5 and 8C1-2 (Demerec, Kaufmann, Fano,
MUTATIONS 231
Sutton, and Sansome, 1942, Carnegie Inst. Wash. discoverer: Skinner, 42cl8.
Year Book 41: 191). Hannah-Alava places it in references: Ives, 1943, DIS 17: 50.
7D1-2. phenotype: Like sns. Female sterile. RKl.
other information: The singed locus is divided into s n 73o7
three recombinationally different sites (Ives and origin: X ray induced in R(l)2.
Noyes, 1951, Anat. Record 111: 565; Hexter, 1955, discoverer: Hannah, 1947.
Proc. Natl. Acad. Sci. U.S. 41: 921-25; 1957, references: Valencia, 1966, DIS 41: 58.
Genetics 42: 376). sn occupies right site. Some of cytology: Associated with T(l;3)sn13ttl =
the double mutants synthesized by Hexter and T(1;3)6C;7C9-D1;79D2-E1.
studied by Bender (1960).
* s n 27-70
50k origin: Induced by mustard gas.
36a 5 discoverer. Sobels, 57j.
3 I references: 1958, DIS 32: 85.
1 * s n 27-49
origin: Induced by mustard gas.
Map of the sn locus discoverer: Sobels and Jansen, 57j.
Drawn from Hexter, 1957, Genetics 42: 376. references: Sobles, 1958, DIS 32: 85.
*sn29-7
origin: Spontaneous. discoverer: Sobels and Schouten, 571.
discoverer: Bridges, 1912. references: Sobels, 1958, DIS 32: 85.
Bibliog. Genet. 2: 235. origin: X ray induced.
Bender, 1960, Genetics 45: 867-83 (fig.), discoverer: Patterson.
phenotype: Bristles wavy, twisted at ends. A weak references: 1934, DIS 2: 59.
allele of sn. Hairs wild type. Female fertile. RK2. phenotype: Like sn. Female sterile. RKl.
other information: Occupies middle pseudoallelic sn***
site (Hexter, 1955, Proc. Natl. Acad. Sci. U.S. 41: discoverer. Duncan, 34e20.
921-25). references: 1935, DIS 4: 10.
sn* phenotype: Bristles show slight sn effect. Hairs
origin: Spontaneous. kinked. Female fertile. RKl.
discoverer Mohr, 22fll. sn36o
references: 1923, Hereditas 4: 142-60 (fig.). origin: Spontaneous.

Bender, 1960, Genetics 45: 867-83 (fig.). discoverer: Spencer, 36a21.
phenotype: Bristles and hairs curved and twisted as references: Bender, 1960, Genetics 45: 867—83
in sn. Female entirely fertile. RK1. (fig-)-
other information: Occupies left pseudoallelic site phenotype: Bristles gnarled in a fairly extreme
(Hexter, 1955, Proc. Natl. Acad. Sci. U.S. 41: manner. Hairs wild type. sn36a is only allele to
921-25). cause pronounced reduction in replication of
sn4 oocyte nurse cell DNA [King and Burnett, 1957,
origin: Spontaneous. Growth 21: 263—80 (fig.)]. sn36a also causes more
discoverer: Bridges, 30a26. extreme retardation of vitellogenesis than other
references: Bender, 1960, Genetics 45: 867—83 female-sterile sn alleles (Bender, 1960). Female
(fig-), sterile. sn3â sn4 homozygote has nearly normal
phenotype: Bristles kinked at ends as in sn2. Hairs bristles and is sterile. RKl.
wild type. Female fertile. sn36a sn4 homozygote other information: Occupies left pseudoallelic site
has nearly normal bristles and is sterile. RK2. (Hexter).
other information: Occupies middle pseudoallelic Sp37b
site (Hexter, 1955, Proc. Natl. Acad. Sci. U.S. 41: origin: Spontaneous,
921-25). discoverer Poulson, 37b.
snS references: 1938, DIS 10: 55.
origin: Spontaneous. 1939, DIS 12: 49.
discoverer: Bridges, 30b5. phenotype: Like sn. Female sterile. RKl.
references: Bender, 1960, Genetics 45: 867—83.
phenotype: Bristles and hairs kinked. Expression origin: Spontaneous.
intermediate between an and sn4. Female sterile. discoverer: Buzzati-Traverso, 39fcl9.
RKl. references: 1940, DIS 13: 49.
other information: Occupies right pseudoallelic site phenotype: Like sn. Female sterile. RKl.
(Hexter). *sn41l
*snss: singed-5 of Skinner origin: Spontaneous.
origin: Spontaneous. discoverer: Oliver, 41i25.
references: 1942, DIS 16: 53. discoverer Muller.

phenotype: Like sn. Female sterile. RK1. references: Bender, 1960, Genetics 45: 867—83.
sn'9BbS
origin: X ray induced. origin: X ray induced in In(l)scs* LscSR+dl-49.
discoverer: Belgovsky. discoverer: Muller, Valencia, and Valencia, 1946-53.
references: 1946, DIS 20: 63. references: Valencia, 1966, DIS 41: 58.
phenotype: Weak sn; hairs unaffected. Female fer- cytology: Associated with T(l;3)snI9Bbs =
tile. RK1. T(l;3)3Cl-2;7C9-10;72A-B.
other information: Crossing over unaffected.
*5n48h *snK: singed of Krivshenko
origin: X ray induced. discoverer Krivshenko.
discoverer Lindsley, 48hll. references: Agol, 1936, DIS 5: 7.
references: 1949, DIS 23: 60. phenotype: Like sn. Female sterile. RK1.
phenotype: Like sn4. Female fertile. RK1.
*sn49h snX2: singed from X irradiation
origin: Induced by P32. origin: X ray induced.
discoverer: R. C. King, 49h. discoverer Muller.
references: Poulson and King, 1949, DIS 23: 63. references: Bender, 1960, Genetics 45: 867—83.
phenotype: Like sn. Female sterile. RK1. snb: sunburst
snS0k location: 3-34 or 47 [6.5 units from D (3-40.4)].
discoverer: Ives. discoverer: Dobzhansky.
references: Ives andNoyes, 1951, Anat. Record phenotype: Eye color soft maroon with seven flecks.
111: 565. Overlaps wild type. Classification best in fly at
Bender, 1960, Genetics 45: 867-83. least one day old. Larval Malpighian tubes some-
phenotype: Kinky hairs and gnarled bristles. Fe- what lighter than normal (Brehme and Demerec,
male sterile. RK1. 1942, Growth 6: 351-56). RK3.
other information: Occupies right pseudoallelic site. so: sine oculis
locotion: 2-57.1.
origin: Spontaneous. discoverer: Milani, 1939.
discoverer Hillman, 55a. references: 1941, DIS 14: 52.
references: 1957, DIS 31: 82. Buzzati-Tra verso, 1946, DIS 20: 63.
phenotype: Bristles and hairs affected, but not so Milani, 1946, Boll. Soc. Ital. Biol. Sper. 23: 111 —
extreme as sn3. Female fertile. RK1. 13.
1951, DIS 25: 79.
origin: Spontaneous. 1951, Rend. 1st. Lombardo Sci. Lettere, Ser. 3, 84:
discoverer: Kadel. 143-54.
references: 1957, DIS 31: 83. phenotype: Ocelli always absent. Eyes usually re-
phenotype: Like sn. Female sterile. RK1. duced to small groups of ommatidia. More extreme
at elevated temperatures; lethal at 30°C. so eye
origin: Gamma ray induced, disks transplanted into wild-type host develop
discoverer: Mickey, 61k. autonomously as do wild-type disks in so host
references: 1963, DIS 38: 31. (Castiglioni, 1950, DIS 24: 79). RK2.
phenotype: Like sn, RK2. SO*
origin: Spontaneous derivative of so.
origin: Gamma ray induced. discoverer. Milani, 1939.
discoverer: Mickey, 61k. references: 1946, Boll. Soc. Ital. Biol. Sper. 22:
references: 1963, DIS 38: 31. 1025-28.
phenotype: Like sn3. RK1. 1949, Sci. Genet. 3: 106-112.
sn63a phenotype: Less extreme than so. Ocelli absent.
origin: Found among progeny of male treated with Eyes usually normal, but sometimes reduced or de-
radio frequency. formed. Homozygous expression not affected by
discoverer: Mickey, 63a. temperature. At 20°-23°C, so2/so eyes like
references: 1963, DIS 38: 29. &o2/®o2. At 27° so2/BO may resemble so/so; eyes
phenotype: Like sn. RK1. range from normal to greatly reduced or deformed.
sn63h RK2.
origin: Found among progeny of male treated with scum somors
radio frequency. location: 1-40.8.
discoverer: Mickey, 63b 19. origin: Induced by DL-p-KN-di-(2-chloa-oethyl>rnino-
reference*: 1963, DIS 38: 29. phenvlalaniae (CB. 3007).
sne discoverer: Fahmy, 1953.
MUTATIONS 233
phenotype: Pigmentation of body and e y e s dark and discoverer: M. (Mann) Lesley.

dull. Wings occasionally divergent or blistered. synonym: Br: Bristled.
Good viability and fertility. RK2. references: 1923, Genetics 8: 27—36.
other information: One allele induced by CB. 1414. phenotype: Sternopleural bristles increased in
sp: speck number. At 19°C wild type; at 25° overlaps wild
location: 2-107.0. type; at 28—30°no overlap. Apparently does not
origin: Spontaneous. affect sternopleural bristles on metathoracic seg-
discoverer: Morgan, 10c. ment converted by bx to a tnesothoracic segment
synonym: ol-2: olive-2. (Waddington, 1939, Growth Suppl. 1 pp. 37—44).
references: Bridges and Morgan, 1919, Carnegie Homozygous lethal. RK2.
Inst. Wash. Publ. No. 278: 128 (fig.). cytology: Salivary chromosomes apparently normal
Morgan, Bridges, and Sturtevant, 1925, Bibliog. (Morgan, Bridges, and Schultz, 1937, Carnegie
Genet. 2: 211 (fig.), 236. Inst. Wash. Year Book 36: 301).
phenotype: Axils of wings have black specks. Body cytology: Placed in salivary chromosome region
color dark. In pupa, region of anal papilla is dark 27C1 to 28C1 (E. H. Grell).
(Waddington). RK1.
cytology: Placed in 60B13-60C5, on basis of its
inclusion in the 2RPXD element of T(l;2)Bld =
T(1;2)1C3-4;6OB12-13 and Df(2R)Px = Df(2R)60B8-
10;60Dl-2 but not in Df(2R)Px* =Df(2R)60C5~
6;60D9-10 [Bridges, 1937, Cytologia (Tokyo), Fujii
Jub. Vol 2: 745-55].
Sp: Sternopleural
sp~w: see WSP

spa: sparkling
location: 4- [probably most distal visible locus on
chromosome 4 (Abrahamson, Herskowitz, and
Muller, 1956, Genetics 41: 410-19)].
discoverer L. V. Morgan, 34k6.
sp: speck references: 1941, DIS 14: 52.
From Bridges and Morgan, 1919, Carnegie Inst. Wash. 1947, Genetics 32: 200-19.
Publ. No. 278: 129. phenotype: Byes rough in varying degrees and some-
what bulging. Affected by genetic modifiers. More
sp: see extreme at 17°-19°C than at 22°-25°. Heterochro-
SP2 matin and sex affect expression so that X/0 >
discoverer: Bridges, 25f. X/X > X/Y > X/X/Y; also enhanced by M(2)S2*0.
phenotype: Darker speck and body color than sp. spa haplo-4's have an exaggerated phenotype.
Tyrosinase formed in adults (Horowitz). RK1. RK2.
$pS6 7: speck of Shuman cytology: Placed in 102D-F, on basis of the ab-
origin: Spontaneous. sence of spa + from the 2LD4P element of T(2;4)b =
discoverer Shuman, 61c. T(2;4)25E;102C15-Dl (E. B. Lewis). Observa-
references: Meyer, 1963, DIS 37: 51. tions on its further location conflict. Fahmy re-
phenotype: Similar to sp. RK1. stricts its location to 102D, on basis of its inclu-
spu: speck from ultraviolet sion in D%4)M63* •= D%4)101E;102D13-El, whereas
origin: Ultraviolet induced. Hochman places it between 102E2 and 102F10, on
discoverer: Meyer, 52d. basis of its inclusion in Df(4)U « Di(4)102E2-
references: 1955, DIS 29: 74. 10;W2F2-10.
phenotype: Weak allele. spu/@p not difficult to
classify, but spa/sp2 overlaps wild type. RK2. sparkling-Cataract
Sp: Sternopleural origin: X ray induced,
location: 2-22.0. discoverer. BeLgovsky, 1936.
origin: Spontaneous. syrtonym: Cat.

Morgan, 1941, DIS 14: 52, discoverer: Bridges, 30dl5.
phenotype: Posterior third or half of eye of heterozy- phenotype: Wings short and broad; pointed at tip,
gote rough; facets irregular and fused. Homozygous and warped at base. Effect on wing shape arises
lethal. Stocks vary in expression, presumably be- from excessive contraction of epithelium from in-
cause of genetic modifiers. X/X and X/0 flies that flated stage onward (Waddington, 1940, J. Genet.
are spaCat/spa show the bulging eyes and rough- 41: 75—139). Overlaps wild type in existing stock.
ening of spa and the posterior fused facets of RK3.
spaCat; X/X/Y and X/Y flies have only the cytology: Placed in salivary chromosome region
spaCat phenotype. spaCat/spaPo1 has fusion of 27C1 to 28C1 (E. H. Grell).
facets over entire surface of eye and roughness in spdfv: spade-flag
posterior region of eye. spaCat/4-sim is wild type. origin: Spontaneous.
RK2. discoverer: Doane, 60fl4.
spa*(lxh sparkling-enhancer of lozenge synonym: fg.
discoverer: H. A. Bender, 65b23. 1961, DIS 35: 45-46.
phenotype: Homozygote wild type in absence of lz; phenotype: Wings about two-thirds the length and
eyes strongly roughened in presence of heterozy- three-fourths the width of wild type; held tentlike
gous Iz3, lz^G, or lz-D. Slight eye roughening when over abdomen. Alulae absent or vestigial; proximal
both spae(Ix) and a lz allele are heterozygous. posterior wing margins often irregular with tend-
spae(lz)/spaPot and spae(tx)/spaP6S have very ency to fold under about vein L4. Venation usually
rough eyes but normal tarsal claws and sperma- normal with occasional blistering. spd{6/ spd has
thecae. RK3. phenotype varying from slight shortening of wings
spaP &'; sparkling-poliert typ e to a shape midway between the two homozygotes.
origin: Spontaneous. Excellent viability and fertility. RK1.
discoverer: Sturtevant, 1961. spe: see lzs
phenotype: Eyes small, rough, and glazed. More specific dilutor: see dil
extreme than spaPot or spaP65. Nonpigmented speck: see sp
tarsal claws. RK1. spectacled: see lzs
spaP65 spermatheca: see spt
origin: Spontaneous. spineless: see s$
discoverer: H. A. Bender, 65J11. spiny legs: see sple
phenotype: Eyes somewhat reduced in size, rough, spl: split
and partially glazed. More extreme than spaP°l but location: 1-3.0.
less so than apaP6*. Tarsal claws unpigmented origin: X ray induced.
and possibly reduced; reminiscent of certain loz- discoverer Dubinin.
enge mutants. Pulvilli and accessory female repro- synonym: shd; fa3 (1934, DIS 1: 10).
ductive structures appear normal. Heterozygote references: Serebrovsky and Dubinin, 1930, J.
with spaP°l and spaP61 has affected tarsal claws Heredity 21: 259-65.
as well as rough eyes. Heterozygote with spa has Agol, 1931, Genetics 16: 262.
slightly roughened eyes at 25°C but markedly phenotype: Eyes rough and small. Many bristles
roughened eyes at 18°; female somewhat more ex- doubled; sometimes missing. Bristle effect caused
treme than male. Viability and fertility good. RK1. by an extra division of initial bristle-forming cell
spaP°': sparkling-poliert [Lees and Waddington, 1943; Proc. Roy. Soc.
origin: Spontaneous. (London), Ser. B 131: 87-110 (fig.)]. Few bristles
discoverer: Hadorn, 51a. (but not their sockets) regularly removed from pos-
synonym: pol. terior border of tergites in spl/+ heterozygotes
references: Rickenbacher, 1953, DIS 27: 59. (Welshons). spl in heterozygotes with other reces-
1954, Z. Induktive Abstammungs- Vererbungslehre sive members of the iV pseudoallelic series is vir-
86: 61-68 (fig.). tually normal, but it is spl when heterozygous with
phenotype; Eyes rather small; surface smooth and N. RK1.
glassy. During second day of pupal life, retinula cytology: Placed in band 3C7, on basis of interac-
cells withdraw from other cells of eye disk. tion with N. Salivary chromosomes normal
spaPol/spaCet has extreme phenotype; spaPol/apa (Welshons).
slightly more extreme than spa {Sturtevant, 1961, other information: A member of the pseudoallelic
DIS 35: 47). Homozygote has excellent viability series at the Notch locus; located between NNic
and fertility. RK1. and N36*-1 03 (Welshons and Von Halle, 1962,
spade: see spd Genetics 47: 743-59).
sparkling: see spa *spl2
spastic: see sps origin: Spontaneous.
spd: spade discoverer: Gottschewski, 1935.
location: 2-21.9 [to the left of Sp (E\ H. Grell)]. phenotype: Like sp!, but eyes smaller. RK1.
MUTATIONS 235
splay wing: see sp/w spread wings: see spr

sple: spiny legs spreadex: see sdx
location: 2-54 (5.5 units to the right of b). sps: spastic
discoverer: Goldschmidt. origin: Ultraviolet induced,
references: 1945, Univ. Calif. (Berkeley) Publ. discoverer: Edmondson and Meyer, 49d.
Zool. 49: 503-4, 521. references: 1951, DIS 25: 73.
phenotype: Hairs on legs irregular, giving a spiny phenotype: Pupal and postpupal lethal. Fly that
appearance. RK3. emerges from pupal case unable to walk or fly.
split: see spl Spastic contraction and jerking of leg and wing
split thorax: see spx muscles. Fly becomes overturned and stuck; sur-
Splotched: see SI vives less than 24 hr; sterile. Muscles so relaxed
sp/w; splay wing in etherized fly that mutant indistinguishable from
location: 1-58.6. normal fly. RK3.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- spt: spermatheca
references: 1958, DIS 32: 75. discoverer: Hadorn, 43e.
phenotype: Wings shortened and usually slightly references: Hadorn and Graber, 1944, Rev. Suisse
divergent. Eyes small and occasionally rough and Zool. 51: 418-23.
deformed. Body size reduced slightly. Emergence Graber, 1949, Z. Induktive Abstammungs-
delayed. Male sterile; viability about 10 percent Vererbungslehre 83: 106-35 (fig.).
wild type. RK3. phenotype: At 28°C female has two spennathecae,
other information: One allele induced by CB. 1246. but ducts partly fused; at 25° only one enlarged
*spot: spot spermatheca on one duct; at 18° a duct with three
location: 3- (not located). branches, each bearing a spermatheca. Tempera-
discoverer Hersh, 34hl5. ture-sensitive period in third larval instar. Female
references: 1935, DIS 4: 14. fertility not greatly affected. RK3.
phenotype: Dark spot appears below eye on posterior spt: see stt^
margin of head. Expression variable. RK3. *spw: spur wing
spotted white: see wsP location: 3- (right arm).
spotty: see stt origin: Spontaneous,
spotty-tergum: see stt2 discoverer Wallbrunn.
spr: spread wings references: 1942, DIS 16: 54.
location: 3- [right arm associated with In(3R)P\. phenotype: Wings vary from normal to large fan-
origin: Spontaneous. shaped structures with extra veins; often a spur-
discoverer: Bridges, 36cl6. shaped lobe from costal margin. Penetrance better
phenotype: Wings held out at wide angle. Both in female and in old cultures. RK3.
sexes sterile. RK3A.
*Spr: Spread spx: split thorax
location: 3- (rearrangement). location: 1-22.6.
discoverer: Oliver, 32k21. discoverer Fahmy, 1956.
references: 1935, DIS 4: 15. references: 1959, DIS 33: 91-92.
phenotype: Wings held outstretched perpendicular to phenotype: In extreme manifestation, thorax split
body axis; drooping in older fly. Homozygous into two segments by longitudinal furrow; abdom-
lethal. Heterozygote viability somewhat low. Fe- inal tergites also split along mid-dorsal line. Eyes
male fertile; male quite infertile. RK2A. deformed. In least abnormal fly, always a hairless
cytology: Associated with In(3L)Spr, breakpoints stripe along the dorsal midline of thorax. Wings
unknown. often slightly divergent. Occasionally one or both
*sprd: spread palpi abnormal in position or structure. Viability
location: 3-65. and fertility rather low in male, very low in female.
origin: Spontaneous in In(3R)C. RK3.
discoverer Dexter, 13k. other information: One allele each induced by CB.
synonym: sd (preoccupied), 2511 and CB. 3007. Two alleles induced by CB.
references: Bridges and Morgan, 1923, Carnegie 1528.
Inst. Wash. Publ. No. 327: 105. spx: see sdx
phenotype: Wings spread at right angles to body. *sq: square
RK2A. location: 2-8.4.
other information: Probably separable from In(3R)C = discoverer: Bridges, 17hl7.
ln(3R)92Dl-El; 100F2-3. phenotype: Wings truncated with squarish or oblique
Spread: see Spr tip. Overlaps wild type. Viability erratic. RK3.
*Sq: Squat *sr300.24

location: 2-38. origin: X ray induced.
origin: Spontaneous. discoverer: Alexander.
discoverer: Bridges, 15k29. references: Ward and Alexander, 1957, Genetics 42:
references: Bridges and Morgan, 1919, Carnegie 42-54.
Inst. Wash. Publ. No. 278: 283-84 (fig.). phenotype: Homozygous lethal. RK2A.
phenotype: Wings short, broad, blunt, arched, and cytology: Associated with Df(3R)sr300.24 =
less transparent than normal. Thorax and head Df(3R)90C2-4;91A2-5.
short and broad. Legs short and weak. Overlaps *sr300.101
wild type. Homozygous lethal. RK3. origin: X ray induced.
sr: stripe discoverer: Alexander.
location: 3-62.0. references: Ward and Alexander, 1957, Genetics 42:
discoverer: Bridges, 22b6. 42-54.
references: Bridges and Morgan, 1923, Carnegie phenotype: Homozygous lethal. RK2A.
Inst. Wash. Publ. No. 327: 244. cytology: Associated with Df(3R)sr300.101 -
phenotype: Trident pattern on thorax replaced by Df(3R)90D2-4;91A6-8.
broad dark median stripe; color intensified by e/+ *sr300.240
and e/e. Midthorax flattened; some hairs turn origin: X ray induced.
toward midline; colorless bubbles in midthorax. discoverer: Alexander.
Wings tend to droop or be raised. RK1. references: Ward and Alexander, 1957, Genetics 42:
cytology: Placed between 90D2 and 90F7, on basis 42-54.
of its inclusion in both Dt(3R)srlO0.394 = phenotype: Homozygous lethal. RK2A.
Df(3R)90C2-7;90F3-7 and Df(3R)sr300.101 - cytology: Associated with Tp(3)sr3 00.240 -
Df(3R)90D2-4;9lA6-8 (Ward and Alexander, 1957, Tp(3)75C;89E;92A.
Genetics 42: 42-54). *srb: smaller body
*sr3 2 location: 1-62.0.
origin: X ray induced. origin: Induced by S-mustard (CB. 1735).
discoverer Alexander. discoverer: Fahmy, 1960.
references: 1960, Genetics 45: 1019-22. synonym: sby-62: small body 62.
phenotype: Homozygous lethal. RK2A. references: 1964, DIS 39: 58.
cytology: Associated with In(3R)sr3-2 = In(3R)90Dl- phenotype: Body size slightly reduced. Bristles
B1;93B-E. finer. Both sexes viable. Female fertility low.
*sr4.2 RK3.
origin: X ray induced. cytology: Not included in deficiency for 18A4-18B8
discoverer Alexander. formed by combining left end of In(l)y4 = ln(l)lA8-
references: 1960, Genetics 45: 1019-22. Bl;18A3-4 with right end of In(l)sc9 = In(l)lB2-
phenotype: Homozygous lethal. RK2A. 3;18B8-9, although sby (1-60.8) is (Norton).
cytology: Associated with T(2;3)sr*-2 = *srd: small round
T(2;3)30C;90C-96. location: 1-0.6.
*srJ00.23 origin: Induced by 2-chloroethyl methanesulfonate
origin: X ray induced. (CB. 1506).
discoverer Alexander. discoverer: Fahmy, 1955.
references: Ward and Alexander, 1957, Genetics 42: references: 1959, DIS 33: 92.
42-54. phenotype: Fly small with slightly dark, rounder,
cytology: Associated with Tfir;3)sri00.33 _ small eyes. One or both postvertical bristles fre-
T(Y;3)90E2-3. quently missing. Both sexes viable and fertile.
* $ r 100.372 RK3.
origin: X ray induced. ss; spineless
discoverer Alexander. location: 3-58.5.
references: Ward and Alexander, 1957, Genetics 42: discoverer Bridges, 14a3.
42-54. references: Bridges and Morgan, 1923, Carnegie
phenotype: Homozygous lethal. RK2A. Inst. Wash. Publ. No. 327: 109 (fig.).
cytology: Associated with 7X2;3)8r^0.312 m Morgan, Bridges, and Sturtevant, 1925, Bibliog.
7X2,3)40-41 ;9OD2-E1. Genet. 2: 211 (fig.), 236.
*sr100.394 phenotype: Bristles only a little larger than hairs;
origin: X ray induced. dorsocentrals least reduced. Postscutellars erect.
discoverer: Alexander. No effect on legs or aristae. Growth of bristles
references: Ward and Alexander, 1957, Genetics 42: slows during development [Lees and Waddington,
42-54. 1943, Proc. Roy. Soc. (London), Ser. B 131: 8 7 -
phenotype: Homozygous lethal. RK2A. l i o j . RK1.
cytology: Associated with DS[3R)mt1(>0-394 m cytology: Locus placed in 89C1-2 (Lewis, 1963,
Di[3R)9OC2-7;9OF3-7. Am. Zoologist 3: 33-56).
MUTATIONS 237
other information: A compound locus (Hexter). affecting aristae, e.g., th and al, but are affected
by those operating on tarsi, e.g., fj, d, app, and ey
origin: Spontaneous. (Waddington, 1939, Growth, Suppl. 1, pp. 37-44;
discoverer Poulson, 37b. Braun, 1940). RK1.
references: Poulson and King, 1948, DIS 22: 55. other information: To the left of ssa40a (Hexter).
phenotype: Similar to ss but with some differences. ssa40a
Vertical bristles, particularly posterior verticals, origin: Spontaneous.
smaller than in ss; some scutellars shorter and discoverer: Buzzati-Traverso, 40a2.
have square tips; occasionally some scutellars references: 1940, DIS 13: 49.
missing. Viability good. RK1. phenotype: Antennae and aristae tarsuslike. Legs
always show four tarsal joints fused and swollen.
In male, sex combs enlarged and sometimes present
on the second pair of legs. Bristles practically
wild type. RK1.
other information: To the right of s s a (Hexter).
discoverer. Neel, 4H30.
phenotype: Like ss*. RK2.
origin: Spontaneous derivative of ssa*0a.

discoverer: Buzzati-Traverso, 44al7.
phenotype: Antennae and aristae tarsuslike but
without claws. Male legs normal. Less extreme
than ssa*Oa. Bristles practically normal. RK1.
discoverer: Meyer, 52g.
phenotype: Extreme allele. Bristles and hairs re-
duced so much that male sometimes lacks sex
55°; spinel ess-aristapedia combs. Aristae leglike, with claws. Homozygote
From Bridges and Brehme, 1944, Carnegie Inst. Wash. weak; male sterile; female only slightly fertile.
Publ. No. 552: 179. RK2.
*ssa53*
ss°: spineless-aristapedia origin: Spontaneous.
origin: Spontaneous. discoverer: Pitemick, 1953.
discoverer Balkaschina, 1926. phenotype: Aristae tarsuslike with fused, distorted
references: 1929, Arch. Entwicklungsmech. Organ. joints and terminal claws. Bristles reduced to ves-
115: 448-63 (fig.). tiges, and hairs short. Wings spread and drooping.
phenotype: Antennae and aristae tarsus like; size Tarsal joints of legs fused, swollen, and distorted.
approaches normal tarsus; two claws at tip. Third Viability low; fly sterile. ssaS3e/ssa like ssa.
joints of antennae like parts of a tarsal row but saa53e/ssiao53 iias fleshy proximal segments of
with broad, flat, plate like lobes below. Bristles aristae. RK2.
like those of a medium to slight Minute. Frequent SS<»63c
extra dorsocentral bristles. Development of tarsi origin: Spontaneous.
in place of aristae enhanced by low temperature discoverer: Merriam and Pitemick, 63c.
(Villee, 1943, Genetics 28: 94). Antennal disks phenotype: Aristae tarsuslike, with terminal claws.
from ssa larvae give rise to leglike structures Tarsal joints of legs sometimes swollen or fused.
when transplanted into wild-type hosts; when disks Bristles shorter than in ssa. ssa£3c/ssa like ss*.
are pretreated with colchicine the developing struc- RK1.
tures more aristalike (Vogt, 1947, Experientia 3: ssaB: spineless-aristapedia of Bridges
156—59). Disks from wild-type larvae also develop origin: Spontaneous.
autonomously when transplanted into &&a hosts discoverer. Bridges, 38all.
(Braun, 1940, Genetics 25: 143-49). Similar re- phenotype: Bristles of female like a slight Minute,
sults observed in mosaics resulting from X-ray-in- especially postscutellars. At 25°C, aristae incon-
duced somatic exchange (Roberts, 1964, Genetics spicuously thickened at base; plumed or threadlike
49: 593—98). •«*/«* has normal aristae but for rest of extent. At 14°, ssaB enhanced and re-
bristles like ss" or slightly smaller. Regions of sembles smm (Villee, 1943, Genetics 28: 94). Legs
aristae converted into tarsi not affected by mutants frequently have lumps at second joint of tarsi;
more pronounced in male and result in doubling of cytology: Placed between 73A2 and 73B1, on basis
sex combs, which are strung along first and second of its inclusion in Df(3L)st* 00.62 = Df(3L)73A2-
fused joints. Eyes a little flattened. Except at 3.-73A10-B1 (Ward and Alexander, 1957, Genetics
low temperatures, all characters slight and may 42: 42-54).
overlap wild type. ssaB/ss has slight Minute
phenotype but wild-type legs and aristae. origin: Ultraviolet induced.
aS
ssaBfssaSp like ss P, with large tarsal aristae. discoverer: Meyer, 54i.
*ssaF: spineless-aristapedia of von Finck phenotype: Like st. RK1.
origin: Spontaneous derivative of s s a . *sfT00.62
discoverer, von Finck, 1937. origin: X ray induced.
references: 1942, Biol. Zentr. 62: 379—400. discoverer: Alexander.
Vogt, 1946, Bioi. Zentr. 65: 238-54. references: Ward and Alexander, 1957, Genetics 42:
phenotype: Bristles normal at all temperatures. 42-54.
Arista leglike at 18°C, leglike at base at 25°, phenotype: Homozygous lethal. RK2A.
normal at 29°. Temperature-sensitive period during cytology: Associated with Df(3L)stloo-62 =
third larval instar. Dominant to more extreme Dt(3L)73A 2-3; 73A10-B1.
alleles and recessive to less extreme. RK1. *5fl00.126
* ss a5p; spineless-aristapedia of Spencer origin: X ray induced.
origin: Spontaneous. d i sc o verer: Alexander.
discoverer: Spencer, 36dl5. references: Ward and Alexander, 1957, Genetics 42:
synonym: arp-1. 42-54.
references: 1937, DIS 7: 5. phenotype: Like st. RK2A.
phenotype: Aristae transformed into nearly normal cytology: Associated with T(Y;3)st100-126 =
tarsi with claws. Third joint of antenna cylindrical T(Y;3)73A2-3.
rather than platelike; hence, antenna is longer and *sf7OO.I77
more leglike than in s s a . Thorax humpy; legs origin: X ray induced.
weakened and misshapen. Bristles practically wild discoverer: Alexander.
type. Viability and fertility good. ssaSP/ssa has references: Ward and Alexander, 1957, Genetics 42:
good arista 1 legs. RK2. 42-54.
ss-A; see AntpR phenotype: Homozygous lethal. RK2A.
ssAT: see AntpLC cytology: Associated with Df(3L)st100-171 =
* s s /so51 ; spineless-isoallele Df(3L)72E4-5; 74 C2-3.
origin: Spontaneous. *Sf100.200
discoverer. Pitemick, 1953. origin: X ray induced.
phenotype: Homozygote is wild type. ssiaoS3/ssa, discoverer Alexander.
ssiao53/sett63ct an( j 8Slao53/ssa53t> faye thick- references: Ward and Alexander, 1957, Genetics 42:
ened proximal segments of aristae, like ssaB. 42-54.
RK3. phenotype: Homozygous lethal. RK2A.
ssv; spineless-variegated cytology: Associated with Df(3L.)stl0O.2OO =
origin: X ray induced, Df(3L)72E4-5; 73A10-B1.
discoverer E, B. Lewis. * s f700.359
phenotype: Variegates for spineless character but origin: X ray induced.
completely mutant for aristapedia. Male sterile. discoverer: Alexander.
RK2A. references: Ward and Alexander, 1957, Genetics 42:
cytology: Associated with T(l;3)ssv *= 42-54.
T(l;3)20;89B;100F. phenotype: Like st. Homozygous lethal. RK2A.
st: scarlet cytology: Associated with TC2;3)st'°0.359 -
location: 3-44.0. T(2;3)21 C3-5;73A2-3;98F2-4.
origin: Spontaneous. st*P: scar let-spotted
discoverer. Richards, 16kl8. origin: Spontaneous.
references; 1918, Biol. Bull. 35: 199-206. discoverer: Bridges, 36bl9.
Bridges and Morgan, 1923, Carnegie Inst. Wash. phenotype: Eyes scarlet with facets and groups of
Publ. No. 327: 172 (fig.). facets that appear wild type. Darkening spreads in
phenotype: Eyes bright vermilion, darkening with old fly. Not a variegated position effect. stBP/st
age. Ocelli colorless, even in old fly; a reliable like &t*P. Larval Malpighian tubules pale yellow
trait for classifying st me. Eyes of bw, st white. and classifiable (Brehme and Demerec, 1942,
Eye color autonomous in larval optic disks trans- Growth 6: 351-56). RK2.
planted into wild-type hosts (Beadle and Ephrussi, cytology: Salivary chromosomes appear normal.
1936, Genetics 21: 230). Larval Malpighian tubes *St: Stumpy
pale yellow (Beadle, 1937, Geoetics 22: 587—611). location: 1-55.5.
MUTATIONS 239
discoverer: Muller, 2612. references: 1958, DIS 32: 75.

references: 1935, DIS 3: 30. phenotype: Short, thin bristles. Viability and fer-
phenotype: Wings and abdomen short. Bristles tility good. RK2.
Minute. Eyes rough. Male lethal. RK2. std: staroid
St-SD: Stabilizer of Segregation Distorter location: 2-56.5.
location: 2- (close to and probably distal to bw). origin: Spontaneous.
origin: Naturally occurring allele. discoverer: E. M. Wallace, 31c26.
discoverer: Sandier and Hiraizumi. phenotype: Eyes small, oval, and very rough.
references: 1960, Genetics 45: 1269—87. Bristles short. Wings slender, dusky, and warped;
phenotype: Decreases variability of transmission marginal veins irregular; gap in L4; L5 short.
ratio of SD-bearing second chromosome among Body dwarfed. Thorax has dark streak. Male
SD/+ males. RK3. sterile. Female semisterile. Viability variable. At
other information: Present on SD-bearing chromo- 19°C, eye character remains but other abnormali-
somes recovered from natural populations. ties disappear. RK2.
Sternopleural: see Sp
*sth: small thin
location: 1-3.7.
(CB. 1506).
phenotype: Fly small, with short thin bristles. Eyes
frequently deformed and rough. Wing shape and
position slightly atypical. Male ecloses late but is
viable and fertile. Female sterile. RK3.
stiff chaetae: see sfc
*sto: stocky
location: 1-29.8.
phenotype: Fly short and stocky. Wings short but
normal in width. Eyes large and pear shaped.
Bristles slightly shorter than normal. Male sterile;
viability about 50 percent normal. RK2.
sta: stubarista other information: One allele induced by CB. 1528.
From Bridges and Brehme, 1944, Carnegie Inst. Wash. *sfo'Pw: stocky-tapered wings
Publ. No. 552: 180. origin: Induced by DL~p-NN-di-(2-chloroethyl)amino-
sta: stubarista discoverer: Fahmy, 1954.
location: 1-0.3. synonym: tpw.
origin: X ray induced. references: 1958, DIS 32: 76-77.
discoverer: Oliver, 32122. phenotype: Wings slightly shortened and broadened,
references: 1935, DIS 4: 15. with tip pointed at L3 rather than being smoothly
phenotype: Third joints of antennae short, blunt, rounded. Eyes small and oval. Slightly dusky
free of hairs, and yellowish. Aristae bases thick- thorax. Both sexes viable; female rather infertile.
ened, axes sometimes short, and branches irreg- RK2.
ular. All bristles and hairs extremely short and *stp: silver tips
sparse. Eyes rotated on head slightly so that the location: 1-46.1.
long axis is vertical. RK2A. origin: Induced by 2-chloroethyl methanesulfonate
cytology: Placed in region between 1D3 and 2B, on (CB. 1506).
basis of its association with T(l;3)sta = discoverer: Fahmy, 1956.
T(1;3)1D3-E1;2A;89B21-C4. references: 1959, DIS 33: 92.
sta1*: see crm phenotype: Fly slightly smaller than normal.
Stabilizer of Segregation Distorter: see St-SD Bristles thin, weak, and most are unplgmented;
Star: see 5 hairs unaffected. Male sterile; viability low. RK3.
staroid: see std
*stb: short bristle *Stp-l: Strapped in chromosome 1
location: 1-14.6. location: 1-50.6 (not allelic with sd).
origin: Induced by L-p-NN-di-(2-chtoToethyl)amino- origin: Spontaneous.
phenylalanine (CB. 3025). discoverer: Hannah.
discoverer. Fahray, 1955. references: 1950, Genetics 35: 669.
phenotype: Expression limited to male. About 15 particularly on fourth tergite. Tergites occa-
percent of Stp-1; Stp-2/+ males show some scal- sionally ridged or broken. Bristles long and
loping of wing margins. Most Stp-1; Stp-2/Stp-2 straggly. Male sterile; viability about 30 percent
males have some degree of scalloping; varying normal. RK2.
from a small nick to vestigal-like wings. Modified other information: Allelism inferred from similarity
by both genetic and environmental factors. Without in phenotype and genetic location at 34.1.
Stp-2, Stp-1 has no effect. RK3. *stu: small tumoroid
*Stp-2: Strapped in chromosome 2 location: 1-20.4.
location: 2- (right arm between c and sp). origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
origin: Spontaneous. phenylalanine (CB. 3025).
discoverer: Hannah, discoverer: Fahmy, 1954.
references: 1950, Genetics 35: 669. references: 1959, DIS 33: 92.
phenotype: 15 percent of Stp-1; Stp-2/+ and most phenotype: Fly small; frequently has small mela-
Stp-1; Stp-2/Stp-2 males show incising of wing notic pseudoturnors. Viability 5 percent wild type.
margin. Stp-2/Stp-2/+ and Stp-2/Stp-2/Stp-2 inter- Male fertile. RK3.
sexes show scalloping in the presence or absence stubarista; see sta
of Stp-1. RK3. stubarista-P3^: see crm
*Sfr: Stretched wings Stubble: see Sb
location: 2-67. Stubble~recessiv&: see sbd
discoverer: Tanaka, 34al2. stubbloid: see sbd
references: 1937, DIS 8: 11. Stubby: see Sy
phenotype: Wings divergent. Homozygous lethal. Stubby-30: see
RK2. Sttibby-31119: see
straight abdomen: see sab stubs: see sbs
Strapped: see Stp Stumpy: see St
straw: see stw stw: straw
strawberry: see swb location: 2-55.1 [0.03 unit to the right of rl (Tano,
Streak: see Sk 1966, Japan. J. Genet. 41: 299-308); between rl
streaked stern i: see sts and apbtt (Sturtevant, 1949, DIS 23: 98)].
streakex: see stx discoverer: Bridges, 17fll.
Stretched wings: see Str references: Morgan, Bridges, and Sturtevant, 1925,
stripe: see sr Bibliog. Genet. 2: 237.
sts: streaked sterni phenotype: Hair color yellowish, especially on legs.
location: 1-60.3. Bristles pale at tips. Heterozygous deficiency for
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- stw produces paling of body color. RK2.
phenylalanine (CB. 3007). cytology: Placed in 41B or C, on basis of pale body
discoverer: Fahmy, 1954. color of heterozygotes for the deficiency from 41B3
references: 1959, DIS 33: 92. through 42A2 formed by combining left end of
phenotype: Small fly with light body color. Brown In(2R)Cy = ln(2R)42A2-3;58A4-Bl with right end of
areas on abdominal stemites often form two longi- In(2R)bwVDel =In(2R)41B2-Cl;59E2-4 and inclu-
tudinal lines. Eclosion delayed. Viability and fer- sion of stw in several cytologically invisible defi-
tility low. RK3. ciencies at base of 2R, e.g., Df(2R)M-S2 (Schultz).
*sff: spotty stw*
location: 1-34.3. discoverer Bridges, 21 g.
origin: Induced by p-NN-di-(2-chloroethyl)aniino- synonym: swy,
phenylethylamine (CB. 3034). references: Morgan, Bridges, and Sturtevant, 1925,
discoverer: Fahmy, 1955. Bibliog. Genet. 2: 239.
references: 1959, DIS 33: 92. phenotype: Hairs pale yellow; bristles brownish
phenotype: Fly small. Wings slightly deformed. with yellow tips. Wings pale yellow and somewhat
Small dark spots on anterior abdominal segments. thin and warped. Slightly more extreme than sfw.
In extreme cases, tergites broken and abnormally Larval mouth parts straw colored at basal prongs
rejoined and hairs deranged. Eyes rather small. and classifiable with difficulty in third-instar
Male sterile; viability about 50 percent wild type. larvae (Bretone, 1941, Proc. Natl. Acad. Sci. U.S.
RK2. 27: 254-61). RK2.
stw3
origin: Induced by 2-chloroethyl methanesulfonate origin: X ray induced,
(CB. 1506). discoverer. Serebrovsky, 1930.
discoverer: Fahmy, 1956. phenotype: Hairs, bristles, wings, and wing veins
synonym: apt: spotty-tergum. straw yellow. Body yellowish with pronounced
references: 1959, DIS 33: 91. dark trident. Tyrosinase formed in adult
phenotype: Fly small; wings wrinkled or pleated. (Horowitz). Wings thin and buckled. Hairs on wing
Darkly pigmented spots dispersed over abdomen, cells incompletely chitinized (Waddkifton, 1941,
MUTATIONS 241
Proc. Zool. Soc, Ser. A 111: 173-80). Puparium references: 1927, Proc. Intern. Congr. Genet., 5th.
noticeably lighter than wild type. Larval mouth Vol. 2: 1193-1200.
parts straw colored at basal prongs; classifiable in phenotype: Suppresses b so that body color is only
living larva (Brehme, 1941, Proc. Natl. Acad. Sci. slightly darker than wild type. No dominant effect.
U.S. 27: 254-61). RK2. Egg hatch in homozygous crosses about 30 per-
other information: Waddington found that irradiation cent, apparently owing to effect on male. RK2.
of stw$ homozygote 2 days before eclosion pro- cytology: Locus placed between 1B4 and 1C4 on
duces reverse mutations that appear as single basis of not being carried on the 2RDXP element
wild-type wing hairs (1940, Nature 146: 335). of T(l;2)Bld = T(1;2)1C3-4;6OB12-13 and being
present on Df(l)260-l =Dt(l)lB4-6.
discoverer: Mather, 37k30. other information: Gives frequent reversion to
phenotype: Body pale yellow. Legs almost color- normal allele.
less. Wings colorless, thin, and fragile. Black *su(B): suppressor of Bar
areas of abdomen still black, but heavily sprinkled location: 2-94.
with pale spots. Larval mouth parts normal origin: Spontaneous.
(Brehme, 1941, Proc. Natl. Acad. Sci. U.S. 27: discoverer: Steinberg, 361.
254-61). RK2. synonym: m(B): modifier of Bar.
origin: Ultraviolet induced. 1937, DIS 8: 11.
discoverer Meyer, 51d. 1939, DIS 12: 49.
references: Meyer and Edmondson, 1951, DIS 25: 1940, Collecting Net 15: 173.
73. 1941, Genetics 26: 325-46, 440-51.
phenotype: Semilethal or associated with a closely phenotype: When homozygous, increases number of
linked semilethal. RK2. eye facets from about 75 to 220 in B male and to
140 in B/B female. Affects all B effects but not
origin: Ultraviolet induced. ey2 or wild type. RK2.
discoverer: Meyer, 51e. *su(B)2: suppressor of Bar in chromosome 2
references: Meyer and Edmondson, 1951, DIS 25: location: 2-46 or -60 (7 units from Tft).
73. origin: Spontaneous.
phenotype: Like stw. Viability low. RK2. discoverer Gans.
phenotype: su(B)2/su(B)2 causes B/+ female to
origin: Ultraviolet induced. appear wild type. RK2.
discoverer: Meyer, 53f. *su(B)4: suppressor of Bar in chromosome 4
phenotype: Bristles yellowish. Wing color pale, but origin: Spontaneous.
often overlaps wild type. Eclosion delayed. Poor discoverer: Brehme, 39k.
viability. RK2. synonym: tn(B)4: modifier of Bar in chromosome 4.
*stwD: straw-Dominant references: 1942, DIS 16: 47.
origin: Spontaneous, phenotype: Facet number in eyes of B male in-
discoverer: Kiil, 38k28. creased, approaching that of B/+ female. Effect
references: Mossige, 1939, DIS 12: 47. increases with age of culture. B/B and B/+ fe-
phenotype: Body and bristles of homozygote light male not affected. RK3.
yellow; wings thin, buckled, and curled. In hetero- Sufbw^1): Suppressor of brown-Variegated
zygote, wings less abnormal; body and bristles location: 2-105.2.
wild type. stwD/stw3 like stw^. stw^/M(2)S2 has origin: Spontaneous.
exaggerated stw phenotype. RK1. discoverer: Kadel, 59bl7.
*stx: streakex synonym: Su-Pm: Suppressor of Plum.
location: 1- (rearrangement). references: 1959, DIS 33: 95.
origin: X ray induced. phenotype: Su(bwvi)/bwvi has wild-type eye color
discoverer: Muller, 26k30. with peppering of dark spots instead of the more or
references: 1935, DIS 3: 30. less uniform brown of bwvl/+« Effect on various-
phenotype: Dark streak down dorsal midline of bwv chromosomes varies from none fen- some to
thorax. Semilethal. RK3A. complete suppression for others. Homozygous
cytology: Associated with ln(l)stx; in the left end viable. RK2.
but breakpoints unknown. cytology: No gross aberration (Lindsley).
su-: see su( ) other information: Su(bwvi) may be a tandem dupli-
Su-: see Su( ) cation. Homozygous Su(bwVl) female produces 0.3
su^-pr: see In(3R)su(pr) percent reversions associated with crossing over
*su(b): suppressor of black in a manner analogous to reversions of 6.
location: 1-0.1. su(Cbx): suppressor of Contrabithorax
discoverer: Plough, 23j28. origin: Spontaneous-
discoverer: E. B. Lewis. references: 1937, DIS 8: 11, 13.

reference*: 1955, Am. Naturalist 89: 73-89. 1938, Genetics 23: 305.
phenotype: Almost completely suppresses Cbx; 1942, DIS 16: 70.
wings made virtually normal, and segmental trans- phenotype: f su(f) has nearly wild-type bristles; in
formations strongly reduced. RK2. about half the flies, some bristles slightly
shortened or twisted at tips. Autonomous in gynan-
Su(Cy): Suppressor of Curly dromorph. f alleles may be divided into suppres-
location: 2- (not located). sible and insuppressible. Among the suppressible
origin: Spontaneous in In(2LR)bwvl. are f, f4, and fs; among the insuppressible are f3
discoverer: Thompson, 61 e. and f3N (Green, 1955, Proc. Natl. Acad. Sci. U.S.
references: 1963, DIS 38: 28. 41: 375—79). su(f) also interacts with wa to make
phenotype: Su(Cy)/Cy has wild-type wings. RK3. the eye of wa su(f) nearly white (Green, 1959,
other information: Separable from In(2LR)bwvl, Heredity 13: 303-15). su(f)/Df(l^nal has Minute-
*su(dx): suppressor of deltex like bristles; eyes rough and ocelli reduced or ab-
location: 1-5. sent, as are ocellar and other head bristles; acros-
discoverer: Bridges, 35c26. tichal rows irregular. Excessive melanization,
synonym: su^-dx: suppressor in X chromosome of especially on head; some crippling of legs. Is a
deltex. fertile female (E. H. Grell). su(f)/In(l)sc*Lsc8R
phenotype: Reduces phenotype of and imparts male and su(f)/0 are normal (Von Halle). RK2.
fertility to dx 8 '. RK2. cytology: Salivary chromosomes appear normal. Lo-
cated near heterochromatic-euchromatic junction as
Su(dx): Suppressor of deltex judged by the fact that su(f) + is carried by certain
location: 2- (not located). free X duplications, e.g., Dp(l;f)3, Dp(l;f)12,
origin: Spontaneous. Dp(l;f)52, Dp(l;f)167 (Lindsley and Sandier, 1958,
discoverer: Bridges, 31a3. Genetics 43: 547—63) and by BSY (Zimmering,
references: Morgan, Bridges, and Schultz, 1931, 1959, DIS 33: 175-76).
Carnegie Inst. Wash. Year Book 30: 410. *Su(f): Suppressor of forked
phenotype: Su(dx)/+ reduces dxst to a slight but location: 2-74.
recognizable, fully fertile phenotype. Su(dx)/Su(dx) origin: X ray induced.
converts dxst to nearly wild type. RK3. discoverer. Dobzhansky, 1931.
Su(dx)2 synonym: Su®-f: Suppressor of forked of
origin: Spontaneous. Dobzhansky*
discoverer: Bridges, 31fl. phenotype: Heterozygous Su(f) reduces expression
references: Morgan, Bridges, and Schultz, 1931, of fz bristles blunt and wavy. Female fertility low.
Carnegie Inst. Wash. Year Book 30: 410. Homozygous lethal. RK3(A).
phenotype: Less effective than Su(dx) as a sup- other information: Crossing over probably reduced.
pressor of dx. RK3. Su(H): Suppressor of Hairless
other information: Found in dx stock, as was Su(dx), location: 2-50.5.
along with ed. Su(dx)2 may s imply be ed Su(dx), or origin: Spontaneous.
it may be of independent origin. Allelism inferred discoverer. Plunkett, 24i.
from phenotype alone. references: Nash, 1965, Genet. Res. 6: 175—89.
Su(er): Suppressor of erupt phenotype: Su(H)/+ is wild type, with L5 occa-
location: 2- (near en). sionally shortened. Su(H)/+; H/+ has nearly
origin: Present in many stocks. normal bristles but shortened L4 and L5. Does not
discoverer Glass, 1941. suppress lethality oiH/H. H2, a stronger allele,
references: 1944, Genetics 29: 436-46. not suppressed so much as H. Homozygous lethal.
1957, Science 126: 683-89. RK3.
phenotype: Only effect is suppression of er. Semi- cytology: Placed in region between 34E5 and 35D1
dominant. Exposure to 1000 r of X rays from on the basis of its lethality in combination with
shortly after fertilization [8 min, according to Df(2L)64j ~ Df(2L)34E5-Fl;35C3-Dl (E. H. Grell).
Glass (1957) but not until 10 hr, according to *su(Hw): suppressor of Hairy wing
Hildreth] to middle of second larval instar inhibits location: 3-54.8.
Su(®r), and er is then manifested in about 98 per- origin: Spontaneous.
cent of flies. Tryptophan fed to larvae has a sim- discoverer. Bridges, 23e4.
ilar effect. Some related compounds have a lesser references: 1932, Proc. Intern. Congr. Genet., 6th.
effect; kynurenine and indole acetic acid have Vol. 2: 12-14.
little or no effect. RK3. phenotype: Hw male and Hw/+ female made wild
su(f): suppressor of forked type; Hw/Hw female has only a trace of Hw pheno-
location: 1-65.9 (to the right of mml and left of bb). type. Also suppresses «c and ct6 completely and
origin: X ray induced. to a lesser extent /and B. Body rather squat.
discoverer. Vhittiaghill, 37g4. Wings slightly spread and warped. Female sterile;
synonym: au^-f. male fertile. Viability good. RK2.
MUTATIONS 243
other information: L. V. Morgan, Bridges, and T. H. phenotype: Completely suppresses pr, but fly is
Morgan discovered suppressors that were probably dilapidated and poorly viable. Both sexes sterile.
allelic (if not the same allele) to Su(Hw). These Enhances Hw. RK3(A).
mutations all lost. cytology: Association with In(3R)su(pr) (breakpoints
su(Hw)2 unknown) inferred from crossover reduction in 3R.
origin: Spontaneous in bx3 chromosome. su(pr)&: suppressor of purple of Bridges
discoverer: E. B. Lewis, 1948. discoverer: Bridges, 29al3.
references: 1949, DIS 23: 59-60. references: 1932, Z. Induktive Abstammungs-
phenotype: Resembles description of su(Hw). Al- Vererbungslehre 60: 207—18.
most completely suppresses Hw; sc1, scDi, and Schultz and Bridges, 1932, Am. Naturalist 66:
3 5 6 a 10
SCZ)2 b u t n o t S C 2 ( sc" "^, sc , sc , sc?, s c , sc , 323-34.
Si S2 6 n 3
sc , or,sc ; dm; ct but not ct ; lz but not lz , phenotype: Eye color of pr; su(pr)B is wild type.
lz34k, lz36, tz37hf lz48ti lz6t OT iza. Bx2 b u t n o t Eyes large and bulging. Wing venation irregular;
Bx, Bx3, BxJ, Bxr, or Bx^9k; t)X3> bx34°, a n d body color pale; low viability; late hatching, and
bxd but not bx, Cbx, pbx, or Ubx; ci but not cis7&, short lived. Male entirely sterile; female partially
ciD, or ciw. Partially suppresses B and f and the sterile. su(pr)B/su(pr) suppresses pr, viability and
yellow wing color of y2. Does not suppress y, y2S, fertility high. RK3.
y34ci yv2, ac, ac3, svr, svrPoi, su(s)2, su(s)s, tw, *su(s): suppressor of sable
br, kz, pn, pn2, su(wB), w, wa, w<*2, w*3, wbt, location: 1-0.
WX yyCO yyl discoverer: Bridges, 1915.
2
spl, cho, cho , ec, peb, rb, bo, ex, cv, vs, cm, sn, synonym: Originally called sable duplication.
sn2, sn3, sn4, sn34e, sn36*, oc, gg2, t, t2, t3, t4, references: 1919, Anat. Record, 15: 357-58.
amx, ras, ras2, ras3, v, v36f, vot, m, dy, tw, wy, Schultz and Bridges, 1932, Am. Naturalist 66:
wy2, &> ty> pl, yb, un, if3, cs53, f36a, sy,. car, to, 323-34.
net, a/, ex, S, shv, ho, E(S), Cy, ft, dp, dpov^, pi, phenotype: With su(s), s is nearly wild type. su(s)/+
Sp, b, el, rd", pu, hk, pr, Bl, Alu, It, tl, stws, ap, with s/s is as dark as s/s or nearly so. Also sup-
apblt, pk, ltd, dil2, en, en, sea, vg, eg, L (three presses v (probably only one allele tested). No
alleles), gp, c, fj, sm, a, px, bw, bw2b, bwD, pd, record of testing with pr or sp. RK2.
mr, or, sp, bs2, R, Ly,. D, cp, in, pP, ry? sbd2, Sb, su(s)2
Sb/sbd2, ss*, ssa/ss, DrL, H, Pr, ca, gvl, discoverer: Bridges, 1915.
sv^®, svn, ey, ey2, ey4, ey36e, eyD, M(2)173, references: 1919, Anat. Record 15: 357—58.
M(2)38b, M(2)l2, M(2)p, M(2)S1, M(2)S23, Bonnier, 1926, Hereditas 7: 229-32.
M(2)mS6, M(2)S7, M(2)S29, andM(2)mS13. Does not Schultz and Bridges, 1932, Am. Naturalist 66:
suppress variegation of w+, 7V+, or bw+. Ovaries 323-34.
rudimentary; female sterile. RK2. phenotype: Suppresses s, v, sp, and pr. Shows
su(lz3*): suppressor of /ozenge-34 allele specificity at v locus; suppresses v and v2
location: 3- (not located). but not v361, v**«, v51a, v^lbr or V51c (Green,
origin: Spontaneous. 1952, Proc. Natl. Acad. Sci. U.S. 38: 300-5).
discoverer: H. A. Bender. Allele specificity at s, sp, and pr not tested. Re-
references: Bender and Green, 1960, Genetics 45: stores wild-type eye color, reduces nonprotein
1563-66. tryptophan accumulation (Shapard, 1960, Genetics
phenotype: lz34; su(lz34) eyes are larger, less 45: 359—76) and partially restores tryptophan pyr-
rough, and more normal in color than lz34 alone. rolase activity (Baglioni, 1960, Heredity 15: 87—
Female distinctly more fertile with su(lz34) but 96; Kaufmann, 1962, Genetics 47: 807-17) in sup-
still lacks parovaria and spermathecae. RK2. pressible v mutants. Heterozygous su(s)2 has
su(pd): suppressor of purpleoid slightly suppressive action on v (Shapard, 1960;
location: 3- (not located). Baglioni, I960). RK2.
origin: Spontaneous. su(s)3
discoverer Bridges, 22h. origin: X ray induced.
phenotype: Normal by itself. Changes pd eye color discoverer: Schultz, 33a2.
to wild type. RK2. phenotype: Suppresses s, v, and sp; pr not tested.
Su(Pm): see su(bwvi) RK2.
*su(pr): suppressor of purple Su(s)S0!
location: 3-95.5 [measured for su(pr)B]. origin: X ray induced.
origin: Spontaneous. discoverer: Green.
discoverer Stern, 27c2. synonym: mtSQ!6-v: mippresmar-5016 of vermilion.
synonym: aus-pr. references: 1951, DIS 25: 70.
references: 1929, Z. Induktive Abstammungs- phenotype: Suppreses v, not tested for suppression
Vererbungslehre 52: 373—89. of «, sp, or pr. RK.2.
Schultz and Bridges, 1932, Am. Naturalist 66:
323-34. origin: Spontaneous,
Stern, 1934, DIS 1: 35. discoverer: Green.
synonym: suslS6-v. references: 1950, DIS 24: 59.

references: 1952, DIS 26: 63. phenotype: Homozygous or heterozygous Su(ss)2
phenotype: Like su(s)2 in suppression of v. Not causes ss to have long bristles that are only
tested for suppression of s, sp, or pr. RK2. slightly thin, like a mild Minute; however, the pos-
*Su(s)S2c terior scutellars remain greatly reduced as in un-
origin: Spontaneous; simultaneously with $c$2c. suppressed ss. RK2.
discoverer: Green, Su(ss)3
synonym: suS3c-v. origin: Spontaneous.
references: 1952, DIS 26: 63. discoverer: Hexter, 1950.
phenotype: Suppresses v. Not tested for suppres- references: 1953, DIS 27: 55-56.
sion of other loci. RK2(A). phenotype: ss Su(ss)3 homozygote wild type for all
other information: May be inversion since crossing bristles; ss Su(ss)3/ss + intermediate between ss
over between su(a)S2c and v virtually eliminated. and wild type, ss Su(ss)3/ss bx Su(ss)2 is wild
su(s)s: suppressor of sable of Stern type. RK2.
origin: Spontaneous, su(t): suppressor of tan
discoverer: Stern, 33jl9. location: 3-26.
synonym: su^^-v pr. origin: Spontaneous.
references: 1936, DIS 5: 8. discoverer Bridges, 22k2.
1937, DIS 7: 20, 21. phenotype: Converts t to wild type. RK3.
phenotype: Suppresses v and pr. RK2. su(tu-bw): suppressor of tumor with brown
other information: No record of test with s or sp, but location: 3- (not located but probably in 3L).
said to be allelic to su(s). origin: Naturally occurring allele.
Su(S): Suppressor of Star discoverer Glass, 1941.
location: 2-3; based on cytological location between references: Glass and Plaine, 1952, Proc. Natl.
shr (2-2.3) and ho (2-4.0). Acad. Sci. U.S. 38: 697-705.
origin: Synthetic. Glass, 1954, DIS 28: 74.
discoverer: Curry, 37b. Burnet and Sang, 1964, Genetics 49: 223-35, 599-
references: Morgan, Bridges, and Schultz, 1937, 610.
Carnegie lust. Wash. Year Book 36: 301. phenotype: Reduces incidence of melanotic masses
Lewis, 1945, Genetics 30: 154. in tu-bw homozygote from 85—100 percent in
phenotype: Su(S)/S and Su(S)/+ wild type. RK2A. su(tu-bw)/+ to 5—10 percent in su(tu-bw) homozy-
cytology: Associated with the deficiency for 22D1 gote. Suboptimal ratios of pentose nucleotides,
to 22E1 or the deficiency for 33F to 34A9, or both, cholesterol deficiency, or excess L-tryptophan in
derived by combining the left end of In(2L)Cy = the larval diet, as well as X irradiation of embryos,
ln(2L)22Dl-2;33F5-34Al and the right end of increase incidence of melanotic masses in tu-bw;
Ia(2L)t = In(2L)22D3~El;34A8-9. According to su(tu-bw) homozygote. Glass and colleagues at-
Lewis (1945), the region between 22D1 and 22E1 tribute this to an effect on su(tu-bw), whereas
is more likely responsible. Burnet and Sang believe the reaction controlled by
*Su(sc): Suppressor of scute tu-bw is affected. Does not suppress tu-48
location: 3-59. (Burnett, 1966, DIS 41: 161). RK3.
discoverer Payne, Su(var): Suppressor of variegation
synonym: sc-Inh-3: acute Inhibitor cm chromosome 3; location: 3-41.3.
Bxt-mct-3. origin: Spontaneous.
references: 1921, Genetics 5: 501—42. discoverer Spofford, 61c.
Bridges and Morgan, 1923, Carnegie Inst. Wash. synonym: Su-V,
PubL No. 327: 158. references: 1962, Genetics 47: 986—87.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. 1965, DIS 40: 36.
Genet. 2: 225, 235. phenotype: Reduces variegated mutant effect (some-
phenotype: Tends to restore bristles removed by sc times completely) of w, rst, is, sp/, nd, and dm in
in Su(®c)/+ heterozygotes. RK3. Dp(l;3)N364~S*' Also reduces w variegation of
*Su(ss): Suppressor of spineless In(l)wm4, mt variegation of ln(l)rst3, and sc varie-
location: 3-61 (between toe and sr), gation of In(l)ac8. Enhances sc variegation of
origin: Spontaneous, ht(l}Mc4 and y variegation of ln(l)y3P. Semidomi-
discoverer: Bridges, 22gl5. nant; heterozygote less suppressed than homozy-
references: Morgan, Bridges, and Sttsrtevant, 1925, gote. Shows maternal effect; Stt(var)/+ offspring of
EMblioc. Geoet. 2; 236. Sut(var)/Su(var) more normal than Su(var)/+ off-
phenotype: SU(M)/+ converts »m/»a to wild type spring of Su(var)/+ mothers. Homozygote fertility
•accept for reduced and erect posterior scutellars. slightly reduced. Viability excellent. RK2.
Bonocygtms lethal. RK2. su(ve): suppressor of veinlet
Sw(ssP location: 3- - 0 . 1 (0.1 unit to the left of m).
origin: Spontaneous, origin: Spontaneous.
discoverer: E. B. Lewis, 1947. discoverer Curry, 37a.
MUTATIONS 245
phenotype: At 19°C, suppression of ve is complete references: Morgan, Bridges, and Sturtevant, 1925,
except tip of L2 occasionally missing. At 25° sup- Bibliog. Genet. 2: 235 (fig.).
pression only partial, with some overlap into range Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20.
of unsuppressed ve. At 19°, su(ve)/+ partially sup- phenotype: Bristles reduced, somewhat variably.
presses ve. RK2. Trichogen irregularly displaced and usually partly
cytology: Not included in Dt(3L)D = Df(3L)61E2- converted to socket (Lees and Waddington, 1942,
Fl;62A4-6 from T(Y;2;3)D; therefore, probably lo- DIS 16: 70). sv/sv/sv triplo-4 nearly normal, sv
cated in 61A-E. haplo-4 extreme shaven (Schultz, 1935, Am. Natu-
su(wa): suppressor of white-apricot ralist 69: 30—54). Expression depends on tempera-
location: 1-0.1 (placed at 0.05 by Green). ture: excellent at 19°C, overlaps wild type at 25°,
origin: X ray induced. and entirely wild type at 30°. RK2.
discoverer: Schultz, 1941. cytology: Placed in region between 1O2E2 and
phenotype: Darkens eye color of w° to brownish. 102F10, on basis of its inclusion in Dt(4)ll =
Does not affect waî w*3t vva^, or any other w Df(4)102E2-10; 102F2-10.
allele tested (Green, 1959, Heredity 13: 303-15). SV*: see sv"
RK2(A).
cytology: Placed in region ID or E, on basis of its discoverer: Ives, 35al8.
inclusion in Dp(l;f)112 = Dp(l;f)lE4-Fl; 19-20 but references: 1935, DIS 4: 11.
not in Dp(l;f)3 = Dp(l;f)lD; 19-20 (Gersh). May phenotype: Resembles svn more than sv. Bristles
have small duplication in region 1D-E (Schultz). frequently reduced to stumps. RK2.
*su(w")2
discoverer: Schultz, 1944.
phenotype: Like su(wa). RK2(A).
cytology: May have small inverted section in region
1D-E (Schultz).
su(w")G: suppressor of white-apricot of Green
origin: Spontaneous in In(l)sc8, y31d w*.
discoverer: Green.
phenotype: Like su(wa). RK2A.
*Su(y3P): Suppressor of yellow-3 of Patterson
location: 3-90.
discoverer: Parker, 48h.
synonym: su-y31e.
phenotype: Su(y3P)/+ suppresses y3P to about
normal color, except that wings remain yellowish.
y3P; Su(y3P)/Su(y3P) is darker than wild type but
wings remain yellow. May be suppression of varie-
gation since extra Y chromosomes also suppress S V c/«. shaven-depilate
y3P. No effect on y, y2, y2S, y3d, y4, y35a, or Edith M. Wallace, unpublished.
ytd. Homozygote has low viability and fertility;
occasionally, wings held out from body. RK2. s W *: shaven- dep i I ate
sunburst: see snb discoverer. E. M. Wallace, 37a24.
*sup: superwith phenotype: More extreme than sv". Thorax denuded
location: 3- (not located). over large areas. Both sexes sterile. RK2.
discoverer. Morgan, 10k.
references: Bridges and Morgan, 1923, Carnegie s v : shaven-naked
Inst. Wash. Publ. No. 327: 35. discoverer. Mohr, 31 j ! 3 .
phenotype: Trident pattern on thorax dark. RK.3. synonym: sv 2 .
Super-Bar: see 5^3i references: 1933, Hereditas 17: 317-22 (fig.).
superwith: see sup phenotype: Extremely short bristles. Viability ex-
suppressor: see su( ) cellent. Trichogen irregularly displaced, becoming
Suppressor: see Suf ) more or less converted into tormogen [Lees and
*sv: shaven Waddington, 1943, Proc. Roy. Soc. (London), Ser.
location: 4-3.0 [in dipla-4 txiploids (Sturtevant, B, 131: 87-110 (fig.)]. In triplo-4 sWsv*Vsv",
1951, Proc. Natl. Acad. Sci. U.S. 37: 405-7)]. the phenotype is more normal than in diplo-4. RK1.
origin: Spontaneous. other information: Selective advantage for triplo-4 in
discoverer. Bridges, 20kl4. stocks of tsv*> results in accumulation.
svr: silver phenotype: Like svrPoi but somewhat more extreme.

discoverer: Bridges, 23g23. *svrP°lmH: silver-pointed lanceolate
synonym: slv. origin: Spontaneous derivative of svrPoi.
references: Morgan, Bridges, and Sturtevant, 1925, discoverer Goldschmidt.
Bibliog. Genet. 2: 235. references: 1947, J. Exptl. Zool. 104: 197-222.
Morgan, 1940, DIS 13: 51. phenotype: Wings resemble //; some are truncated.
phenotype: Color of legs, wings, veins, and integu- Semidominant in female; expression poor in male.
ment pale and silvery. Bristles and trident pattern RK2.
on thorax dark. Tyrosinase formed in adult cytology: Salivary chromosomes appear normal
(Horowitz). Wings of all males and some females (Hannah).
pointed. Viability fair. Larval mouth parts normal *svrP°i'M: silver-pointed soft
in color. RK2. origin: Spontaneous.
cytology: Locus placed at 1B5-6 (Demerec, discoverer: Goldschmidt.
Kaufman, Fano, Sutton, and San some, 1942, references: 1945, Univ. Calif. (Berkeley) Publ.
Carnegie Inst. Wash. Year Book 41: 191). Zool. 49: 291-550.
svrP°h silver-pointed phenotype: Nearly wild type except when combined
origin: Spontaneous. with a ba ; then, wings narrowed and pointed at tip.
discoverer: Goldschmidt, 1934. Heterozygous abB also gives narrow wings.
synonym: poi. RK3(A).
references: 1944, DIS 18: 42. cytology: Salivary chromosomes appear to have a
1945, Univ. Calif. (Berkeley) Publ. Zool. 49: 291- two- to four-band inversion of 1E1-4 (Goldschmidt
550. and Hannah, 1944, Proc. Natl. Acad. Sci. U.S. 30:
phenotype: Body color pale. Wings pointed and oc- 299-301).
casionally soft or spread. Suppressed by y. Sup- *svrP°'-*': silver-pointed singed
presses sp and partially suppresses s. RK2(A). origin: Spontaneous derivative of svrPoi.
cytology: Salivary chromosomes show abnormality at discoverer Goldschmidt.
1E3-4 (Goldschmidt and Hannah, 1944, Proc. Natl. references: 1945, Univ. Calif. (Berkeley) Publ.
Acad. Sci. U.S. 30: 299-301). Zool. 49: 291-550.
*svrP° '-*»': silver-pointed blistered phenotype: Indistinguishable from svrPoi. RK2.
origin: Spontaneous. *$vrP°'"*«J; silver-pointed square
discoverer: Goldschmidt, 1934. origin: Spontaneous,
references: 1945, Univ. Calif. (Berkeley) Publ. discoverer. Goldschmidt.
Zool. 49: 291-550. references: 1945, Univ. Calif. (Berkeley) Publ.
phenotype: Barely distinguishable from svrPot; best Zool. 49: 291-550.
recognized in svrPoi~bt; aba, which has long, other information: Presumably arose in svrPol-bl but
pointed wings with blistered area. RK2. not clearly different.
*svrpo'-C<»: silver-pointed from Canton stock *svrPot: silver-Pointed Dominant
discoverer: Goldschmidt. discoverer: Goldschmidt.
references: 1947, J. Exptl. Zool. 104: 197-222. references: 1947, J. Exptl. Zool. 104: 197-222.
phenotype: Almost like svrPoi. RK2. phenotype: Resembles svr, does not suppress sp.
cytology: Salivary chromosomes appear normal Homozygote viable. Wings slightly truncated. RK2.
(Hannah). cytology: Salivary chromosomes appear normal
*svrpot'di*h: silver-pointed dishevelled (Hannah).
origin: Spontaneous. *svrp°l'*: silver-Pointed semidominant
discoverer: Goldschmidt. origin: Spontaneous.
reference*: 1945, Univ. Calif. (Berkeley) Publ. discoverer: Goldschmidt.
Zool. 49: 291-550. references: 1947, J. Exptl. Zool. 104: 197-222.
phenotype: Body color pale with reduction of dark phenotype: Pointed wings with good expression.
bands at posterior edge of abdominal tergites. RK2.
Hairs on tergites few and irregular. In extreme cytology: Salivary chromosomes appear normal
cases, all hairs irregular. Enhanced by abm. Sup- (Hannah).
presses *p. RK3(A). *svs: shortened veins
cytology: Abnormality of 1E3-4 (Goldschmidt and location: 1-24.6.
Hannah, 1944, Proc. Natl. Acad. Sci. U.S. 30: origin: Induced by ethyl methanesulfonate (CB.
299-301). 1528).
*SVTJ»O/-JI; silver-pointed heat discoverer: Fahmy, 1956.
origin: Recovered among F2 of heat-treated larvae. synonym: &hv (preoccupied),
discoverer. Goldschmidt. references: 1959, DIS 33: 90.
references: 1945, Uaiv. Calif. (Berkeley) Publ. phenotype: Wings highly abnormal, varying from
Zool. 4t: 291-550. small stubs to almost full size with inner margin
MUTATIONS 247
cut away. Vein L4 often shortened and posterior origin: Induced by 2-chloroethyl methanesulfonate
crossvein absent. Eyes small and deformed. Male (CB. 1506).
fertile; viability about 50 percent wild type. Fe- discoverer: Fahmy, 1956.
male sterile. RK2. references: 1959, DIS 33: 92.
phenotype: Body color slightly dark; darkened scu-
tellum particularly noticeable. Eyes brownish
(best detected immediately after eclosion) and oc-
casionally misshapen, swy/s is wild type. Via-
bility about 50 percent wild type. Both sexes
fertile. RK2.
sx: sexcombless
references: Mukherjee, 1965, Genetics 51: 285—304
(fig.)-
phenotype: Number of teeth in primary sex comb re-
duced from the normal 10 to 1. Bristles interme-
sw: short wing diate between normal bristles and sex-comb teeth
From Eker, 1935, J. Genet. 30: 357-68. also appear in sex-comb area. Bristle pattern of sx
male basitarsus feminized in other respects. sx/+
sw: short wing reduces the mean number of sex-comb teeth in
location: 1-64.0. tra/tra female from 11.37 to 3.7. Sex-comb devel-
discoverer: Eker, 32al2. opment autonomous in mosaic from either chromo-
references: 1935, J. Genet. 30: 357—68 (fig.). some loss or somatic crossing over in tra/ tra fe-
1939, J. Genet. 38: 201-27. male (Mukerjee and Stem, 1965, Z. Vererbungslehre
phenotype: Above 23°C, most flies have spread and 96: 36—48). Reduces number of teeth in secondary
incised wings with irregular veins; eyes reduced sex comb of en/en male and in primary sex comb of
and roughened. Male expression more extreme than eyD/+ male. Male sterile owing to imperfect devel-
female. Above 27.5°, viability low; above 31°, sw opment of internal duct system; testes often remain
is lethal. At 17°, most flies are wild type; at 14°, unattached to ducts, and are therefore ellipsoidal,
all are wild type. RK2 at 28°C. but contain fully developed sperm (Stern, 1941, J.
swY: see stw2 Exptl. Zool. 87: 113—58). External genitalia also
*swo: swollen antenna greatly modified. Size, shape, and arrangement of
location: 1-1.3. teeth on c las per varies; occasionally more than
origin: Induced by 2-chloroethyl methanesulfonate one penial apparatus (Mukherjee). RK2A.
(CB. 1506). cytology: Associated with In(l)sx = In(l)llD4-
discoverer: Fahmy, 1956. 6;llE2-6;14B8-9;lSE2-4 (Mukherjee, 1963, DIS 38:
references: 1959, DIS 33: 92. 62).
phenotype: Antennae swollen and deformed. Aristae sy: see oss
abnormal. Eyes slightly rough, pear-shaped, and *Sy: Stubby
browner than normal. Body dark. Wings often up- location: 1- or 2- (rearrangement).
held and frequently incised on the inner margins. discoverer: Ives, 34j31.
Small extra sex combs on second tarsal segments phenotype: Bristles short and thick, especially
of forelegs of most males. Male emerges late, is humerals and notopleurals. Male sterile. RK2.
rather inviable, but is fertile. Female sterile. cytology: Associated with T(l;2)Sy; breakpoints
RK2. unknown, but break in X is genetically at the right
other information: One allele induced by CB. 1528. end.
swarthy: see swy Sy3o: see
swb: strawberry Sy3Hi9:
location: 1-2.2. *syn: syndrome
origin: Induced by L-p-NN-di-(2-chloroethyl)amino~ location: 3-14.7.
phenylalanine (CB. 3025). origin: Gamma ray induced.
discoverer: Fahmy, 1958. discoverer: Wallbrunn, 61i21.
phenotype: Eyes large and rough, with glazed sur- phenotype: Eyes of male translucent brown, of fe-
face; color bright fed but patchy. Inner wing male slightly darker than normal. Wings of male
margins often incised. Male viable and fertile; held at right angle to body, of female held out at
female fertility reduced. RK2. about 45°. Viability low. Both sexes sterile. RK2.
swollen antenna: see swa t: ton
*swy: swarthy location: 1-27.5.
location: 1-42.5. discoverer: Bridges, 14gl6.
248 GENETIC VARIATIONS OP DROSOPHILA MELANOGASTER
references: Morgan, Bridges, and Sturtevant, 1925, references: 1953, DIS 27: 56.
Bibliog. Genet. 2: 237. phenotype: Expression ranges from small black
phenotype: Body color more tan than wild type. spots on distal end of femora or proximal end of
Easiest to identify by light tan antennae; male tibiae to cases in which the tibiae, femora, and
easier than female. Not positively phototropic. bases of coxae are encapsulated in a dark,
(McEwen, 1918, J. Exptl. Zool. 25: 49-106). brownish black, glossy covering. Legs weak.
Tyrosinase formed in adult (Horowitz). Larval Some overlap wild type. Viability reduced. RK2.
mouth parts lighter than normal at basal prongs; other information: Possibly an allele of me (1-29.0).
classifiable with difficulty in larva (Brehme, 1941, Tarnished: see bwv3
Proc. Natl. Acad. Sci. U.S. 27: 254-61). RK2 in tarry: see tar
male. tarsi irregular: see ti
cytology: Locus placed in region 8C3 through 8C17 *taw: tawny
(Demerec, Kaufmann, Fano, Sutton, and Sansome, location: 1-41.1.
1942, Carnegie Inst. Wash. Year Book 41: 191), on origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
basis of its inclusion in Df(l)t2S2-l = Df(l)8C2- phenylalanine (CB. 3026).
3;8C14-D1. discoverer: Fahmy, 1955.
f2 references: 1958, DIS 32: 75-76.
discoverer: Bridges, 19d5. phenotype: Head and thorax slightly dark; abdomen
references: Morgan, Bridges, and Sturtevant, 1925, pale. Wings usually scooped or tips curved. Fe-
Bibliog. Genet. 2: 237. male tergites often narrow, serrated, or broken. Vi-
phenotype: Body color not so pale as t, but antennae ability and fertility good. RK3.
color same. Light sensitive. Larval mouth parts taxi: see tx
lighter than normal at basal prongs; classifiable *tb: tiny bristle
with difficulty in larva (Brehme, 1941, Proc. Natl. location: 1-35.8.
Acad. Sci. U.S. 27: 254-61). RK2 in male. discoverer. Bridges, 16a4.
f* references: 1919, J. Gen. Physiol. 1: 645—56.
discoverer: Bridges, 31ell. phenotype: All bristles short and fine; wings some-
phenotype: Lighter than t; tan spot on abdomen. what short. Female fertility low. RK2.
Basal prongs of larval mouth parts lighter than tb: see tbr
normal; classifiable with difficulty in dissected tb-53: see me
larva (Brehme, 1941, Proc. Natl. Acad. Sci. U.S. *tbd: tiny bristleoid
27: 254-61). RK2 in male. location: 1-25.
i* origin: Spontaneous.
discoverer: Bridges, 33c 14. discoverer: Curry, 37g23.
phenotype: Weak f. RK3. phenotype: Bristles short and thin, like a medium
f282-1 Minute. Fly somewhat smaller than wild type.
origin: X ray induced. Good viability and fertility. RK2.
discoverer. Demerec, 34c. cytology: Locus between 7C5 and 8C1 (Demerec,
phenotype: Lethal in male; ceil lethal. RK2A. Kaufmann, Fano, Sutton, and Sansome, 1942,
cytology: Associated with Dtfiyt26^1 =D%1)8C2- Carnegie Inst. Wash. Year Book 41: 191).
3;8C14-D1 (Sutton). Further restricted to 7E1 through 8C2, on the basis
T*: see dp°lv2 of its genetic location to the right of oc which is
for: tapered excluded from Df(l)sn =Dt(l)7B2-3;7D22-El
location: 2-56.6. (Hinton and Welshons, 1955, DIS 29: 125-26).
origin: Ultraviolet induced. tbr: tracheae broken
discoverer: Edmonds on and Meyer, 49c. location: 3- (not located),
phenotype: Wings narrow and pointed; somewhat discoverer: Slatis.
longer than normal. Veins close together. Viability synonym: tb (preoccupied),
good. Female fertility low; male sterile. RK2. references: 1959, Genetics 44: 536.
t&: see ter phenotype: Main tracheal trunks of larva have inter-
Tac: see Pmâc ruptions. Penetrance 17 percent at 16°C, 5 percent
tarn mo: see fmo at 25°. Does not seem to affect viability. RK3.
tan: see t tc: tiny chaetae
tapered: see fa location: 1-51.6.
tar: tarry origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
location: 1-27.3 (0.4 unit from lz, probably to the phenylalanine (CB. 3007).
left), discoverer: Fahmy, 1954.
origin: Found among progeny of deuteron-irradiated references: 1958, DIS 32: 76.
male. phenotype: Bristles extremely short and fine. Eclo-
discoverer: Hildretfa, 51 i. sion delayed. Viability and fertility good. RKl.
synonym: te (preoccupied). other information: One allele induced by CB. 3007.
MUTATIONS 249
*tdd: tiddler completely so by Gla. Enhanced by D, ey, eyD,

location: 1-0.0 (0/871 crossovers with sc). Me, and YL. RK3.
origin: Induced by ethyl methanesulfonate (CB. *tet-b: tetraltera-b
1528). location: Multifactorial.
references: 1958, DIS 32: 76. discoverer: Goldschmidt, 1950.
phenotype: Body small. Viability and fertility good. synonym: tetBd.
RK3. references: 1952, J. Exptl. Zool. 119: 405-60 (fig.).
*te: tenerchaetae 1953, J. Exptl. Zool. 123: 79-114.
location: 1-5.6. phenotype: Wings reduced; not so halterelike as tet;
origin: Induced by triethylenemelamine (CB. 1246). more frequently leglike, with three joints. RK3.
discoverer: Fahmy, 1952. tetrapter: see ttr
references: 1958, DIS 32: 76. *tf: trefoil
phenotype: Bristles short and fine. Eyes dark and location: 2-55 (between 50 and 60).
glistening. Wings frequently small, deformed in discoverer: Morgan, 13k.
various ways. Eclosion delayed. Male viability, references: Bridges and Morgan, 1919, Carnegie
but not fertility, good. Female infertile. RK3. Inst. Wash. Publ. No. 278: 244 (fig.).
Tegula: see Tg phenotype: Scutellum darkened. Base of trident
telegraph: see tg pattern and back of head have extra areas of dark
telescope: see ts pigmentation. Classification uncertain. RK3.
*ten: tenuis chaetae *tft: tufts
location: 1-43.9. location: 2-102 (between px and bw).
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- origin: Gamma ray induced.
phenylalanine (CB. 3026). discoverer. R. M. Valencia, 1959.
discoverer: Fahmy, 1953. references: 1959, DIS 33: 99-100.
references: 1958, DIS 32: 76. phenotype: Sternopleural bristles form a dense tuft.
phenotype: Bristles short and thin. Body small. Ex- Fully penetrant at 20°C, poorly so at 25°. RK2.
pression more extreme in female. Eclosion slightly Tft: Tuft
delayed. Viability, fertility good. RK3. location: 2-53.2.
tenerchaetae: see te origin: X ray induced.
tent: see tnt discoverer Ritterhoff, 52f25.
tenuis chaetae: see fen references: 1952, DIS 26: 68-69.
*ter: terraced phenotype: In heterozygote, number of scuteilar,
location: 2-36. postalar, and dorsocentral bristles increased; scu-
origin: Spontaneous. tellars increased about fivefold, other bristles to a
discoverer: Bridges, 29cl2. lesser extent. Scutellum shortened; furrow between
phenotype: Eyes have horizontal seam; often a tuft it and the mesonotum absent. Bristles present
of bristles at anterior end of seam; lower half of dorsal to the halteres, at junction of thorax and
eyes depressed and small. Variable, overlaps wild abdomen. Small to moderate amounts of fluid tend
type about 20 percent. Occasionally reverts to to remain between the epithelial layers of the
wild type. RK3. wing. Viability and fertility excellent. In homo-
tet: tetraltera zygote, number of bristles increased, like hetero-
location: 3-48.5. zygote, but shorter. Scutellum quite small. Via-
origin: Spontaneous. bility and fertility low. RK1.
discoverer: Goldschmidt, 341. *tg: telegraph
references: 1940, Material Basis of Evolution, Yale location: 2-0.
University Press, p. 325 (fig.). discoverer: Bridges, 16c27.
Villee, 1942, Univ. Calif. (Berkeley) Publ. Zool. references: Stern and Bridges, 1926, Genetics 11:
49: 125—84. 507 (fig.), 508-10.
Goldschmidt, Hannah, and Piternick, 1951, Univ. phenotype: Vein L2 has one or more gaps or thin
Calif. (Berkeley) Publ. Zool. 55: 67-294. sections. Postscutellar bristles erect or misdi-
phenotype: Wings reduced, with tendency to be rected. Overlaps wild type. RK3.
halterelike. Mesonotum, scutellum, and bristles Tg; Tegula
may be altered or absent; mesothorax becomes location: 2- [o.O to 4.0; associated with Jn(2L)Tg\.
me ta thorax like. According to Lewis, however, origin: X ray induced.
dorsal posterior half of mesothorax (including discoverer E. B. Lewis, 1962.
wing) converted in varying degrees to structure re- references: Mora, 1963, DIS 38: 32.
sembling mirror image of anterior half of meso- phenotype: Wings extended at 90° from body axis,
thorax. Requires a s 8 for expression (Lewis). Ex- often drooping. The tegula (small plate at base of
pression variable; overlaps wild type. Penetrance wing) uniformly duplicated and adjoining anterior
temperature sensitive: 0—1 percent at 29^C» 35 supra-alar bristle usually twinned as well. Horaozy-
percent at 15°. Partially suppressed by Cy and gcus lethal. RK2A.
cytology: Associated with In(2L)Tg = In(2L)21C;22F origin: Induced by D-p-NN-di-(2-chloroethyl)amino-

(Lewis and Mora). phenylalanine (CB. 3026).
th: thread discoverer: Fahmy, 1955.
locotion: 3-43.2. references: 1959, DIS 33: 93.
origin: Spontaneous. phenotype: Legs short and swollen, particularly
discoverer: Bridges, 22J31. posterior pair; swelling most pronounced in tibial
phenotype: Aristae threadlike, without side and tarsal regions. Wings small and broad; diver-
branches. RK1. gent or slightly upheld. Body color slightly dusky
cytology: Placed in region between 72A2 and 72E5, and eye color a bit brownish. Male fertile; viability
on basis of its inclusion in Df(3L)th1<>0.105 - about 20 percent wild type. RK3.
Df(3L)72A2-Bl;73A4-5 but not in Df(3L)sti 00.171 = other information: One allele each induced by. CB.
Dt(3L)72E4-S;74C2-3. In(3L)thiO0.293 = 1506 and CB. 1528.
ln(3L)72A2-Bl;76A4-Bl;79A4-Bl implicates *thH: thick legs-darker
72A2-B1 (Ward and Alexander, 1957, Genetics 42: origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
42-54). phenylalanine (CB. 3025).
*f/,I00.705 discoverer: Fahmy, 1954.
origin: X ray induced. synonym: dkl: darker legs.
discoverer: Alexander, references: 1959, DIS 33: 85.
references: Ward and Alexander, 1957, Genetics 42: phenotype: Extra pigment in body and legs. Legs
42-54. slightly shortened, especially in female. Wings
phenotype: Homozygous lethal. RK2A. small and divergent. Eye shape altered. Viability
cytology: Associated with Dt(3L)thl00.1QS = good in both sexes; female fertility reduced. RK3.
Df(3L)72A2-Bl;73A4-5. *thm: thin macros
*tn100.293 location: 1-48.9.
origin: X ray induced. origin: Induced by 2-chloroethyl methansulfonate
discoverer: Alexander. (CB. 1506).
42-54. phenotype: Bristles slightly shorter and thinner than
phenotype: Homozygous lethal. RK2A. normal. Viability and fertility good. RK3.
cytology: Associated with In(3L)thl<>0.293 = thorny: see tny
In(3L)72A2-Bl;76A4-Bl;79A4-Bl. thread: see th
*tha: thin arched thread bristle: see trb
location: 1-27.8. *tht: thickset
origin: Induced by S-2-chloroethylcysteine (CB. location: 1-42.1.
1592). origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
discoverer: Fahmy, 1957. phenylalanine (CB. 3007).
references: 1959, DIS 33: 93. discoverer: Fahmy, 1953.
phenotype: Fly small, with short thin bristles. references: 1959, DIS 33: 93.
Wings arched over abdomen or drooping at sides. phenotype: Fly reduced in size, more in length than
Viability and fertility low. RK3. breadth, giving a stocky appearance. Eye shape
slightly altered; a few deranged facets. Viability
*thb: thin bristle about 10 percent wild type. Male fertile. RK3.
location: 1-48.0. thv: thick vein
origin: Induced by trie thy lenemela mine (CB. 1246). location: 1-49.7.
discoverer: Fahmy, 1951. origin: Induced by 2-chloroethyl methanesulfonate
references: 1958, DIS 32: 76. (CB. 1506).
phenotype: Bristles thin; short in female. Occasion- discoverer: Fahmy, 1956.
ally, vibrissae abnormal and eyes rough. Vein L5 references: 1958, DIS 32: 76.
sometimes faint or missing beyond posterior cross- phenotype: Veins thick, especially at junction of LI
vein. Viability and fertility good in male but re- and L2. Wings short and broad; marginal hairs ir-
duced in female. RK2. regular. Eyes small and dark. Body color rather
thick: see tk pale. Eclosion delayed. Male viable and fertile.
thick legs: see thl Female fertility subnormal. RK2.
thick vein: see thv *tbv<*: thick vein-delta
thick veins: see tkv origin: Induced by L-p-NN-di(2~chloroethyl)amino-
thickoid: see tkd phenylalanine (CB. 3025).
thickset: see tht discoverer: Fahmy, 1955.
thin arched: see tha synonym: dtv: delta vein.
thin bristle: see thb references: 1958, DIS 32: 69-70.
thin macros: see thm phenotype: Wings slightly short and broad and with
*thh thick legs extra venation, especially around L2, which usu-
location: 1-60.7. ally ends in a delta. Anal plates and genital arch
MUTATIONS 251
deformed; genital region protruding. Male fertile Sometimes a blister near posterior crossvein in fe-
but viability about 50 percent normal. RK3. male; L4 sometimes shortened, especially in fe-
*ti: tarsi irregular male. Female more extreme than male. Easier to
location: 2-55.9. identify at 19°C. RK2.
origin: Spontaneous. *tkv2
discoverer: Ives, 38k5. origin: Spontaneous,
references: 1942, DIS 16: 48. discoverer: Bridges, 34e30.
phenotype: Third and fourth tarsal segments more or phenotype: Veins thickened and with deltas. More-
less fused and swollen. Eyes slightly rough. Via- extreme expression in female and at 19°C. RK2.
bility subnormal. RK2.
tiddler: see tdd */mc: tonomacrochaetae
tilt: see tt location: 1-17.5.
tiny: see ty origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
tiny bristle: see tb phenylalanine (CB. 3026).
tiny bristleoid: see tbd discoverer: Fahmy, 1955.
tiny chaetae: see tc references: 1958, DIS 32: 76.
tiny wing: see tyw phenotype: Bristles thin. Abdomen underpigmented,
tinylike: see tyl especially in female. Eclosion slightly delayed.
Viability and fertility good. RK2.
tk: thick *tmo: tammo
location: 2-55.3. location: Not located.
discoverer: Guthrie, 24k. origin: X ray induced.
references: 1925, Am. Naturalist 59: 479—80. discoverer: Ohnishi, 491.
phenotype: Legs and especially tarsi thick. Wings references: 1950, DIS 24: 62.
somewhat short and broad, with slight px-like ef- phenotype: Bristles one-ha If normal length. RK.2.
fect. According to Waddington [l942, Proc. Zool.
Soc. London Ser. A, 111: 181-88 (fig.)], these ef- *tms: tumorous
fects result from inadequate contraction of the legs location: 1-58.7.
and whole pupa after inflation period. RK2. origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
cytology: Placed in region between 42A2 and 42B1, phenylalanine (CB. 3025).
on basis of its inclusion in inverted segment of discoverer: Fahmy, 1954.
In(2R)Cy = In(2R)42A2-3;58A4-Bl as well as in references: 1959, DIS 33: 93.
Dt(2R)M-S2v611 = Df(2R)40F-41Al;42A19-Bl phenotype: Many small, diffuse tumors. Fly slightly
(Morgan, Schultz, Bridges and Curry, 1939, small. Both sexes viable and fertile. RK3.
Carnegie Inst. Wash. Year Book 38: 273—77). *tnt: tent
location: 1-18.0.
tkd: thickoid origin: X ray induced.
location: 2-40 (30 to 50). discoverer. Fahmy, 1956.
discoverer: Bridges, 33d25. references: 1959, DIS 33: 93.
phenotype: Fly large and thickset, with thick legs. phenotype: Wings droop to variable extent. Bristles
Wings blunt at tip. Eyes large and slightly rough. thin. Fly small. Male sterile. RK2.
Male genitalia sometimes rotated. Fertile but via- *iny: thorny
bility about 50 percent wild type. RK3. location: 1-33.5.
origin: Induced by DL-p-NN-di-(2-chloroethyl)ainino-
phenotype: Fly grossly deformed; extremely inviable.
Eyes small, very rough, and dull red. Thoracic
bristles very short. Wings abnormal, spread, incom-
pletely expanded' Male sterile. RK2.
toni: see ix2

tkv: thick veins tomboy: see ix2
Edith M. Wallace, unpublished. *ton: to no chaetae
location: 1-60.1.
tkv: thick veins origin: Induced by l:4-dimethatiesulfonoxybut-2-yne
location: 2-16. (CB. 2058).
origin: Spontaneous. discoverer: Fahmy, 1951.
discoverer: Nichols-Skoog, 33b25. references: 1958, DIS 32: 76.
phenotype: Veins thickened and branched in region phenotype: Bristles short and thin. Eyes large with
of crossveins, near end of L2, and elsewhere. deranged facets. Wings short, with incised inner
margins and abnormal venation. Variable expres- origin: Induced by D-p-NN-di-(2-chloroethyl)amino-

sion of eye and wing effects. Eclosion slightly de- phenylalanine (CB. 3026).
layed. Male infertile; viability about 50 percent discoverer: Fahmy, 1955.
wild type. Female sterile. RK2. references: 1959, DIS 33: 93.
other information: One allele induced by CB. 1506. phenotype: Bristles short and very thin. Hairs small
tonomacrochaetae: see tmc and sparse. Wings more rounded at tips, margins
tpw: see stotPw often incised; veins slightly thickened. Trident
*tr-26\: triangle-261 pattern slightly darker than wild type. Male viable
location: 3- (not located). and fertile; female sterile. RK2.
origin: Spontaneous. other information: One allele induced by CB. 3026.
discoverer: Spencer. *tre: triangle eye
1935, Am. Naturalist 69: 222-38. origin: Induced by 2-chloroethyl methanesulfonate
phenotype: Small extra crossvein between marginal (CB. 1506).
vein and L2, near their juncture. Variable; over- discoverer: Fahmy, 1956.
laps wild type. RK3. references: 1959, DIS 33: 93.
tra: transformer phenotype: Eyes triangular with apex pointing for-
location: 3-45 (between st and cp). ward. Fly large. Wings broad, blunt tipped, and
origin: Spontaneous. slightly divergent. Male viable and fertile; female
discoverer: Sturtevant, 44d. sterile. RK3.
references: 1945, Genetics 30: 297-99. trefoil: see tf
Seidel, 1963, Z. Vererbungslehre 94: 215-41 (fig.). tri: trident
phenotype: Transforms female into sterile male, location: 2-55.
with fully developed sex combs, male-colored ab- origin: Spontaneous.
domen, normal male abdominal tergites and plates, synonym: Probably trj32k. M33d27. a n d ^33^18
external and internal male genitalia. Testes are the same.
rudimentary, without sperm, and with ovarian-nurse- discoverer: Plough, 32k.
cell-like cells [Brown and King, 1961, Genetics 46: references: Plough and Ives, 1934, DIS 1: 34.
143—56 (fig.)]. Mates readily with female. Testes 1935, Genetics 20: 42-69.
reduced in size, but of normal shape and color. phenotype: Dark trident or streak on thorax. Scu-
Transformed female slightly larger than normal tellura and stemopleural plates also dark. Thorax
male; development rate about that of female. often contains bubbles. Variable; overlaps wild
X/X/Y; tra/tra also sterile. X/Y; tra/tra normal type, but also semidominant. RK3.
male, tra/tra/tra triploid and intersex like diploid triangle eye: see tre
but with larger wing cells. Superfemale intersexual. triangle-261: see tr-261
Normal testis anlage transplanted into tra female trident: see tri
becomes attached to duct apparatus and produces trimmed: see fr*
progeny. RK2. *trl: translucent
tra0: transformer-Dominant location: 2-45 or -65 (10 units from B/).
discoverer: Gowen, 1940. discoverer: Bridges, 20bl7.
synonym: Hr: Hermaphrodite. references: Morgan, Bridges, and Sturtevant, 1925,
references: 1942, Anat. Record 84: 458. Bibliog. Genet. 2: 238.
Gowen and Fung, 1957, Heredity 11: 397-402. phenotype: Eye color translucent ruby, like p. RK2.
Fung and Gowen, 1957, J. Exptl. Zool. 134: 515- tnxi: see fr2
32 (fig.). Truncate-51h: s e e dpolM
phenotype: X/X; traD/Jr intersexual. Body size as *ts: telescope
large as female. Abnormal external genitalia male- location: 2-68.
like. Sex combs with six to eight teeth present. discoverer: Bridges, 15127.
Internal genitalia extremely variable. Spermathacae references: Bridges and Morgan, 1919, Carnegie
and ventral receptacle often present, as are sperm Inst. Wash. Publ. No. 278: 291 (fig.).
pump and paragonia. Gonads most often underde- phenotype: Abdominal segments somewhat drawn
veloped ovaries. Triploid, traD/+/+, resembles 3N out. Wings drooping and divergent. Overlaps wild
female; genitalia female but sex combs have four type. RK3.
to six teeth; sterile. X/X; traD/tra has malelike ft: tilt
appearance; internal and external genitalia male, location: 3-40.0.
and sex combs have eight to nine teeth. RK3. discoverer: Bridges, 15h29.
tracheae broken: see tbr references: Bridges and Morgan, 1923, Carnegie
transformer: see tra
translucent: see tri
Mossige, 1938, Hereditas 24: 115.
trb: thread bristle
phenotype: Wings spread, elevated, and warped in a
location: 1-36.3.
compound curve. Vein L3 shows gap. Eye color
MUTATIONS 253
may be slightly dilute. Developmentally, L3 origi- Barigozzi refers to them as pseudotumors. They
nally complete but central section disappears are under multigenic control, but where adequate
during contraction period (Waddington, 1940, J. analysis exists there usually seems to be one pri-
Genet. 41: 75-139). RK2. mary locus and numerous secondary loci respon-
sible for the phenotype. In many instances, dif-
ferent names have been applied to different deriva-
tives of the same tumor line, which have the same
mary gene but different markers and therefore dif-
ferent constellations of modifiers. The present
treatment represents an attempt to define, insofar
as possible, the primary loci.
tu-1: see tuh-1
*tu-la
discoverer: Payne.
references: Wilson, 1924, Genetics 9: 343—62 (fig.).
phenotype: In combination with tu-lb, produces
melanotic masses in posterior third of third-instar
larva; 20 percent of larvae and 14 percent of adults
affected. Produces some effect when tu-lb hetero-
zygous. Eighty-one percent of larvae with tumor
and 57 percent of those without die before eclo-
sion. tu-la/+; tu-lb/+ has infrequent melanotic
masses. RK3.
*tu-lb
tt: tilt discoverer: Payne.
From Bridges and Morgan, 1923, Carnegie Inst. Wash. references: Wilson, 1924, Genetics 9: 343-62 (fig.).
Publ. No. 327: 135. phenotype: In combination with tu-la, produces
melanotic masses in posterior third of third-instar
*ttr: tefrapter larva; produces some effect when tu-la hetero-
location: 3-51.3. zygous. RK3.
discoverer: Tshetverikov, 25b. *tu-2
references: Astaurov, 1929, Arch. Entwicklungs- location: 2- (not located).
mech. Organ. 115: 424-47. origin: Spontaneous.
1930, Z. Induktive Abstammungs- Vererbungslehre discoverer. Payne.
55: 183-262. references: Wilson, 1924, Genetics 9: 343-62 (fig.).
Timofeeff-Ressovsky, 1934, Z. Induktive Abstam- phenotype: Melanotic masses in larval hemocoele;
mungs- Vererbungslehre 67: 248 (fig.). 20—100 percent of larvae affected. Forty-one per-
Villee, 1942, Univ. Calif. (Berkeley) Publ. Zool. cent mortality of affected larvae. RK3.
49: 180-81. other information: Modifiers on third chromosome.
phenotype: Like ox. Halteres tend to become wing- tu-3: see tuh-3
like. Most flies wild type but may have, in place tu-36a
of a haltere, an organ one-half the size of a normal location: 2- (not located).
wing with veins, bristles, and sense organs. RK3. o ri gi n: Spontaneous.
discoverer: Bridges, 36al6.
Tu: Turned-up wing references; Russell, 1940, J. Exptl. Zool. 84: 363—
location: 1-59. 79 (fig.).
discoverer: Muller, 46119. Oftedal, 1953, Z. Induktive Abstammungs-
references: Muller and Valencia, 1947, DIS 21: 70. Vererbungslehre 85: 408—22.
phenotype: Wings curled; somewhat wrinkled in lon- phenotype: Melanotic masses appear in late larval
gitudinal direction. Heterozygous viability good; ins tars. RK3.
homozygote also viable. RK1. tu-36e
tu: tumor location: Not located,
General term used to denote genes that lead to for- origin: Spontaneous.
mation of melanotic masses of tissue, usually in discoverer: Skoog, 36el5.
late larval stages. Masses apparently result from tv-48
cell aggregation rather than proliferation since location: 2-29.5.
cells in division are not observed within them; origin: Spontaneous.
discoverer: Ghelovitch, 1948. references: Burdette, 1954, DIS 28: 73.

references: 1950, Compt. Rend. 230: 1002-4. phenotype: Small melanotic masses under tergites 1
phenotype: Melanotic masses become visible to un- and 2. Tumor incidence 0.44 percent. RK3.
aided eye in third instar; located in abdomen; vary *tu-59h
in size. RK3. location: 2- (not located).
other information: Modifiers on the X and possibly origin: Spontaneous.
the fourth chromosome. discoverer: Oshima.
origin: Spontaneous. phenotype: Small melanotic masses in third-instar
discoverer: Brncic. larval abdomen; persist into adult stage. RK3.
synonym: iu49h. tu-hw: tumor with brown
references: 1950, DIS 24: 57. location: 2-80.5 (E. H. Grell).
other information: Modifier on third chromosome. Al- origin: Spontaneous.
lelism to tu-48 inferred from interaction in hetero- discoverer: Morgan, 1922.
zygote (Burdette, 1959, Texas Univ. Publ. 5914: synonym: ifit^-: melanotic tumor-A,
57-68). references: 1938, DIS 9: 108.
tu-48** Hartung, 1950, J. Heredity 41: 269-72.
references: Barigozzi and di Pasquale, 1956, Rend. Oftedal, 1953, Z. Induktive Abstammungs-
1st. Lombardo Sci. Lettere, Ser. B 90: 484-509. Vererbungslehre 85: 408-22.
Barigozzi, 1962, Atti Assoc. Genet. Ital. 7: 9-76. phenotype: Numerous melanotic masses in posterior
other information: Allelism to tu-48 inferred from its part of hemocoele; increase in size and number
location about 20 units to the left of 6. through third instar; remain as benign bodies in
tu-48j adult abdomen. Characterized by precocious trans-
location: 3-46. formation of plasmatocytes into lamellocytes
origin: Spontaneous. (which usually occurs at time of pupation); subse-
discoverer: Herskowitz. quent encapsulation of tissue by lamellocytes pro-
synonym: tu48L duces melanotic masses (Rizki, 1957, J. Morphol.
references: 1949, DIS 23: 57. 100: 459-72). RK3.
Herskowitz and Burdette, 1951, J. Exptl. Zool.
117: 499-521. origin: Spontaneous.
phenotype: Melanotic growths appear in larva, more discoverer. Mittler, 1950.
often posteriorly than anteriorly, and in adult ab- synonym: tu^^K
domen. Penetrance around 50 percent; about two references: 1951, DIS 25: 74.
masses per fly. RK3. phenotype: Darkly pigmented spots beneath ventral
tv-49k abdominal surface. RK3.
location: 2- (not located). other information: Allelism inferred from location
origin: Spontaneous. between 75 and 90 on chromosome 2.
discoverer: Oftedal. tv-bwSSg
references: 1951, DIS 25: 122-23. origin: Spontaneous.
1953, Z. Induktive Abstammungs- Vererbungslehre discoverer: Jacobs.
85: 408-22 (fig.). references: Jacobs, Bowman, and Walliser, 1958,
phenotype: Black aggregations of blood cells be- DIS 32: 130.
coming macroscopically visible 96 hr after hatch- phenotype: Melanotic masses appear in larva 55 hr
ing. Result from cell aggregation rather than pro- after batching; persist in adults, commonly in ab-
liferation. RK3. domen, occasionally in thorax, and rarely in head.
other information: Modifiers on X and second chro- Nearly 100 percent penetrant. RK3.
mosome. Not an allele of tu-bw, other information: Allelism by Erk and Sang (1966,
tv-53 DIS 41: 95).
location: 1-41.
origin: Beta ray induced. references: Barrigozzi and di Pasquale, 1956, Rend.
discoverer. Darrow, 53120. 1st. Lombardo Sci. Lettere, Ser. B 90: 484-509.
references: King, 1955, DIS 29: 73. Barigozzi, 1962, Atti Assoc. Genet. Ital. 7: 9-76.
phenotype: Small, melanotic masses in 15—20 per- other information: Tentatively considered to be an
cent of adults. Wings occasionally blistered or allele of tu-bw from crossover data, which place it
nicked; veins knotted or abbreviated. Egg hatching about 18 units to the right of v^.
delayed and reduced to about 65 percent normal. other information: Allelism by Erk and Sang (1966,
Viability 30-50 percent normal. RK3. DIS 41: 95).
other information: Not an allele of ivy. tu-bw*
tu-54e origin: Spontaneous,
location: Not located. synonym: ••'•' tu.
origin: Spontaneous. references: Friedman, Harnly, and Goldsmith, 1951,
discoverer Haddox, 54e23. Cancer Res. 11: 904-11.
MUTATIONS 255
Kaplan, 1955, Trans. N.Y. Acad. Sci. 17: 289-93. wings. Overlaps wild type. tuf/T(2;3)dp has ex-
phenotype: Dark masses in posterior regions of treme form of phenotype. RK2.
larva and in abdomen and thorax of imago. RK3. cytology: Included in In(2R)Cy = In(2R)42A2-
tu-g 3;58A4-B1 (Sturtevant, 1949, DIS 23: 98).
location: 2- (not located). Tuft: see Tft
synonym: tu&. tufted: see tuf
references: Burdette, 1951, DIS 25: 101—2. tufts: see tft
Oftedal, 1953, Z. Induktive Abstatnmungs- tuh-1: tumorous head in chromosome 7
Vererbungslehre 85: 408—22. location: 1-64.5.
Burdette, 1959, Texas Univ. Publ. 5914: 57-68. origin: Spontaneous.
phenotype: Penetrance 47 percent. Second- and discoverer: Griff en.
third-instar larvae have melanized large blood synonym: tu-1.
cells and spindle-shaped cells in hemocoele. RK3. references: Gardner, 1949, DIS 23: 57.
tu-h63: see ey'" Gardner and Woolf, 1949, Genetics 34: 573—85
tu-K: tumor from Oregon-K (fig).
location: 2- (not located). Newby, 1949, J. Morphol. 85: 177-95 (fig.).
origin: Spontaneous. Newby and Thelander, 1950, DIS 24: 89-90.
discoverer: Sang and Burnet. phenotype: In presence of tuh-3, produces asymmet-
references: 1963, Genetics 48: 235—53 (fig.). rical growths of variable size in head region;
1964, Genetics 49: 223-35. mostly external but sometimes internal. Penetrance
phenotype: Small black nodules become evident to- responds to selection. Produces maternal effect in
ward end of third instar, either free in abdominal that reciprocal crosses between tumorous-head
hemocoele or associated with the fat body. Pene- flies and wild type produce different results.
trance low but increased by suboptimal balances of Asymmetry of eye and antenna 1 disks evident in 32
pentose nucleotides, cholesterol deficiency, or an hr larva; contains cells with large chromatic in-
excess of L-tryptophan in the larval diet as well clusions not seen in wild type. Viability about 70
as by X irradiation of embryo. RK3. percent normal. RK3.
other information: Not allelic to tu-hw. other information: Modifiers on chromosome 2.
*tu-R: tumor of Rosenberg
location: 1- (not located). tuh-3: tumorous head in chromosome 3
origin: X ray induced. location: 3-58.5.
discoverer Rosenberg, 57c. origin: Spontaneous.
references: Hinton, 1957, DIS 31: 83. discoverer: Griff en.
phenotype: Bristles shortened. Tumor not described, synonym: tu-3.
but penetrance 40 percent. Developmental time references: Gardner, 1949, DIS 23: 57.
lengthened. Viability of male and homozgous fe- Gardner and Woolf, 1949, Genetics 34: 573—85
male about 50 percent normal. RK3. (fig-)-
*fu-W: tumor from We//e$/ey Newby, 1949, J. Morphol. 85: 177-95 (fig.).
location: 2- (not located). Newby and Thelander, 1950, DIS 24: 89-90.
origin: Spontaneous. phenotype: In presence of tuh-1, produces tumorous-
discoverer. Gowen. head phenotype described under tuh-1. Semidomi-
references: Wilson, King, and Lowry, 1955, Growth nant. Suspected by Woolf (1966, Genetics 53: 295-
19: 215-44 (fig.). 302) of contributing in certain crosses, to inhibi-
phenotype: Melanotic masses become visible 97— tion of attachment of testes to duct system during
102 hr after hatching. Located on surface of or im- development, causing formation of small, un-
bedded in fat body. Located in abdominal segment attached, uniform gonads. RK3.
10 and less frequently in 9 and 11. Characterized Tuh Turneduplike
by precocious transformation of plasmatocytes into location: 1-50 (between g and f).
lamellocytes (which usually occurs at time of pupa- origin: Spontaneous,
tion); subsequent encapsulation of tissue by lamel- discoverer: Muller.
locytes produces melanotic masses (Rixki, 1957, references: 1965, DIS 40: 35.
J. Morphol. 100: 459-72). Viability and fecundity phenotype: Like Tu. Wing tips of heterozygote
low. RK3. turned up slightly but definitely not twisted. Male
other information: Modifiers on the X and third chro- and homozygous female more extreme, with
mosomes. Not allelic to tu-bw or tu-4849h- wrinkled wings sometimes held somewhat apart;
tuf: tufted viable and fertile. RK2.
location: 2-55.5 (between pk and ltd). tumor: see to-
origin: Spontaneous. tutnor he&d-63: see ey*™
discoverer: Sturtevant, 1948. tumorous: see tms
references: 1948, DIS 22: 56. tumorous head: see tuh
phenotype: Small tuft of hairs between eyes and Turned-up wing: see Tu
antennae; basal twinning of anterior halves of Turneduplike: see Tul
fw: twisted dies shortly after eclosion and does not breed.
origin: X ray induced. tx: taxi
discoverer: Demerec, 28cl4. location: 3-91.
phenotype: Abdomen twisted clockwise about 30°, origin: Spontaneous.
as viewed from behind, and not overlapping wild discoverer: Collins, 24j30.
type. Body tends to be dwarfed. Viability about 60 references: 1928, Am. Naturalist 62: 127-36 (fig.).
percent wild type. Male usually fertile. RK2. phenotype: Wings held out at about 75° from body
cytology: Locus between 1CS and 2C10 (Demerec, axis, often arched or wavy, somewhat narrow and
Kaufmann, Fano, Sutton, and Sansome, 1942, dusky. RK2.
Carnegie Inst. Wash. Year Book 41: 191).
*fw2
discoverer: Mohr, 32bl.
phenotype: More extreme than fw. Body regularly
dwarfed. Abdomen twisted 30—60° clockwise, as
viewed from behind; male genitalia often twisted
counterclockwise. Viability about 50 percent wild
type. Male usually fertile, tw^/tw like tw*. RK2.
*twg: twisted genitals
location: 1-48.1.
(CB. 1506).
discoverer Fahmy, 1956. tx: taxi
references: 1959, DIS 33: 93-94. From Collins, 1928, Am. Naturalist 62: 127-36.
phenotype: External genitalia abnormally positioned
on extreme tip of abdomen. Tergites often notched origin: Spontaneous.
at mid-dorsal line. Eyes large, abnormally shaped, discoverer: Tsukamoto, 52j.
and slightly rough. Wings vary from almost normal references: 1956, DIS 30: 79.
to small, deformed structures with very abnormal phenotype: Like tx. Good viability; easily classi-
venation. Bristles frequently waved or bent. Male fied. RK2.
viability and fertility subnormal. RK2. ty: tiny
twirl: see twl location: 1-44.5.
twirled tips: see twt discoverer: Bridges, 25kl.
twisted: see fw phenotype: Bristles small. Body small. Eclosion
twisted genitals; see twg delayed. Viability excellent. Female sterile. Yolk
twl: twirl formation in oocytes inhibited [King and Burnett,
location: 2-63.5. 1957, Growth 21: 263-80 (fig.)]. Follicular cells
origin: Ultraviolet induced. form abnormal derivatives of endoplasmic reticulum
discoverer. Meyer, 54d. and migrate abnormally or form excess of normal
references: 1955, DIS 29: 74-75. endoplasmic reticulum derivative [King and
phenotype: Wings strongly curled. Good viability; Vanoucek, 1960, Growth 24: 333-38; Falk and
easy to classify. RK2. King, 1964, Growth 28: 291-324 (fig.)], ty ovaries
other information: Possibly an allele of upw (2-62). in ry + host develop autonomously (King and Boden-
*Two-b: Two bristles stein, 1965, Z. Naturforsch. 20B: 292-97). RK2.
location: 3-58.3. *tyb-2: tiny bristle-2
discoverer: Bridges, 16b22. origin: Spontaneous.
references: Bridges and Morgan, 1923, Carnegie discoverer: Neel, 4119.
lost. Wash. Publ. No. 327: 155. references: 1942, DIS 16: 52.
phenotype: Two postvertical bristles always and phenotype: Bristles small and thin. Viability and
two anterior dorsocentrals usually absent. Hetero- fertility good. RK1.
zygote viability excellent. Homozygous lethal. tyl: tiny I ike
RK1. location: 1-36.
*twt: twirled tips origin: X ray induced in In(l)dl-49.
location: 1-37.1. discoverer: Oliver, 28k4.
origin: Induced by l:4-dimethanesulfonoxybut-2-yne references: 1935, DIS 3: 28.
(CB. 2058). 1942, DIS 16: 53.
discoverer: Fahmy, 1951. phenotype: Bristles short, fine, and stubblelike.
references: 1959, DIS 33: 94. Eclosion delayed. Both sexes viable and fertile.
pfoeuotype;: Wings completely or partially unex- RK2A.
pended; tips frequently twisted. Male inviable, other information: Not separated from In{l)dl-49.
MUTATIONS 257
tyr-h tyros inase-1 U: Upturned

location: 2-52.4 (4.2 units to the right of b). location: 2-70 (based on UH2°, whose allelism is
origin: Spontaneous. uncertain).
discoverer: H. W. and H. S. Lewis, 1960. origin: X ray induced.
synonym: a1: alpha-1. discoverer: Ball, 32a27.
1961, Proc. Natl. Acad. Sci. U.S. 47: 78-86. phenotype: Wings upturned like those of Cy but dark
1963, Ann. N.Y. Acad. Sci. 100: 827-39. and waxy. Postscutellars crossed as in cu. Body
phenotype: Homozygote has much less tyrosinase color darker than normal. Eyes mottled with light
activity than most strains. Tyrosinase in flecks. Homozygous lethal. RK2A.
tyr-1/ try-1 is heat labile relative to wild type and cytology: Associated with In(2LR)U; breakpoints
has a different substrate profile. Probably tyr-1 unknown.
specifies primary structure of the enzyme. Modi- *UH20
fying genes that alter amount of tyrosinase activity discoverer: Tanaka, 35a6.
in other strains have no effect on homozygous references: 1937, DIS 8: 11.
ryr-i. RK3. phenotype: Wings curled like those of Cy. Homozy-
Tyr-2: Tyrosinase-2 gous viable. RK2.
location: 2-57. Ubx: Ultrabithorax
origin: Naturally occurring allele found in location: 3-58.8.
In(2L)Cy + In(2R)Cy. origin: Spontaneous.
discoverer: H. W. and H. S. Lewis. discoverer: Hollander, 1934.
references: 1963, Ann. N.Y. Acad. Sci. 100: 827— synonym: bx®: bithorax-Dominant; bxdP: bi-
39. thoraxoid-Dominant; Bxl: Bithoraxlike.
phenotype: In combination with some modifying references: 1937, DIS 8: 9, 77.
genes, reduces tyrosinase activity about 50 per- Lewis, 1949, Heredity 3: 130.
cent. Dominant. RK3A. 1951, Cold Spring Harbor Symp. Quant. Biol. 16:
Tyr-3: Tyrasinase-3 159-74 (fig.).
location: 3- (on the right arm). 1954, Am. Naturalist 88: 225-39.
discoverer: H. W. and H. S. Lewis. 1955, Am. Naturalist 89: 73-89.
references: 1963, Ann. N.Y. Acad. Sci. 100: 827- 1963, Am. Zoologist 3: 33-56.
39. phenotype: Halteres of heterozygote about twice
phenotype: In combination with some modifying normal volume, characteristically with one or more
genes, reduces tyrosinase activity about 35 per- hairs on anterior surface of swollen apical seg-
cent. RK3. ment, or capitellum, of the haltere. No overlap
*tyw: tiny wing with wild type and little variability; accurate
location: 3-0. scoring takes practice. Homozygous larva has, in
discoverer: Bridges, 18c9. addition to normally present mesothoracic pair of
phenotype: Wings small. Postscutellars divergent, spiracles, both a metathoracic and a first abdom-
curving upward and forward. Extra bristles on head inal spiracle pair. Flies homozygous for Ubx
and thorax. Viability 60 percent wild type. RK3. but carrying Dp(3;3)bxd^00 = Dp(3;3y66C;89B5-
6;89E2-3, which carries a normal allele of bx but
none of bxd, have extreme bxd phenotype.
Ubx/bx34e has oval, flat halteres; phenotype more
extreme if the third chromosomes are heterozygous
for a chromosome aberration with a breakpoint be-
tween the centromere and the bx locus; E. B.
Lewis (1954) termed this the transvection effect.
bx3** Ubx/+ + indistinguishable from Ubx/+.
Ubx/bxd has large, fleshy halteres like bxd/bxd;
larva lacks first abdominal ventral row of setae.
The cis types, Ubx bxd/+ + and Ubx phx/+ +, also
indistinguishable from Ubx/+. Ubx/pbx has large
halteres and causes transformation of posterior
metathorax toward posterior mesothorax. Homozy-
gous lethal. Enhances expression of Pc and Sex
(Hannah-Alava, 1964, Z. Vererbungslehre 95: 1—9).
RK2.
cytology: Placed close to, if not in, doublet 89E1-2
(E. B. Lewis). Salivary chromosomes normal.
other information: Middle member of bx pseudoallelic
U: Upturned series including, from left to right, bx, Cbx, Ubx,
Edith M. Wallace, unpublished. bxd, and pbx (see map under bx).
synonym: ro-63.
origin: X ray induced in In(3LR)TM6. phenotype: Like un, but wing margins frayed. RK1.
discoverer: Bacher, 67b.
phenotype: Weak Ubx effect. RK3A. origin: X ray induced.
cytology: Not associated with further rearrangement. discoverer: Dubinin, 1928.
bx'oi phenotype: Less extreme and more viable than un or
origin: X ray induced. un 3 . RK2.
discoverer: E. B. Lewis, 1947. *unK: uneven of Krivshenko
synonym: Bxl101. origin: Spontaneous.
phenotype: Like Ubx but much more extreme in inter- discoverer: Krivshenko, 56b9.
actions with other bx pseudoalleles. RK1A. references: 1956, DIS 30: 75.
cytology: Associated with In(3LR)Ubx101 = phenotype: Eyes slightly small, bulging, and rough.
In(3LR)80;89D9-El. Scutellum long and narrow; scutellar bristles thin,
misdirected, and often deformed. Viability and fer-
origin: X ray induced. tility good. RK1.
discoverer: E. B. Lewis, 511. cytology: Salivary chromosomes appear normal.
references: 1952, DIS 26: 66. *unp: uneven from P^2
1952, Proc. Natl. Acad. Sci. U.S. 38: 953-61. origin: Induced by P 3 2 .
phenotype: Like Ubx but much more extreme in inter- discoverer Bateman.
actions with other bx pseudoalleles. RK1A. references: 1951, DIS 25: 78.
cytology: Associated with In(3LR)Ubx^0 = phenotype: Like un. RK2.
In(3LR)61A-C;74;89D-E;93B;96A. other information: Allelism inferred from phenotype
Ubx* and genetic location.
origin: X ray induced. *unc: uncoordinated
discoverer: Schalet, 1959. location: 1-65.9 (reduced from Fahmy's value of
references: 1960, DIS 34: 53, 55. 68.9 to fit on map).
phenotype: Halteres like Ubx. Most flies have vari- origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
able rough eyes and lack one or both postvertical phenylalanine (CB. 3025).
bristles; a few have a slight upward curvature of discoverer. Fahmy, 1954.
wings, RK1A. references: I960, DIS 34: 49.
cytology: Associated with In(3LR)UbxA (cytological phenotype: Fly unable to walk because of lack of
breakpoints unknown), with one break between h coordination in moving legs. Wings held up and
and s t and another left of e. frequently curled at tips. Dies shortly after eclo-
*uex: unextended sion. RK3.
location: 2-55. undersized: see us
origin: Spontaneous. uneven: see un
discoverer: Maeda, 5813. Uneven wing: see Bgz
synonym: unexpended. unexpended: see uex
references: 1962, DIS 36: 39. unexpanded: see unp
phenotype: Wings incompletely expanded as in un expanded irregular: see unr
newly emerged fly; about one-half normal length unextended: see uex
and frequently inflated. Tibiae and tarsi of third unfolded: see uf
legs irregularly shortened and gnarled. Posterior unp: unexpanded
scute liars convergent. Male viability low. RK2. location: 1-63.1.
Uf: Unfolded origins Induced by DL-p-NN-di-(2~chloroethyl)amino-
location: 2- (to the left of 6). phenylalanine (CB. 3007).
origin: X ray induced. discoverer: Fahmy, 1954.
dJ»cov«rer. Beigovsky, 36c29. references; 19S9, DIS 33: 94.
phenotype: Wings spread in homozygote and hetero- phenotype: Wings always unexpanded; frequently
gerte. Viability and ferdlity good. RK3. droop-. Two symmetrical grooves occur on the
Wfra-tef.' see BB pronotam iuMaediately anterior to wing base. Post-
Ultrabithorax: see Ubx sciitellar bristles often crossed. Eclosion delayed.
un: uneven Male fertile; viability about 10 percent normal. Fe-
location: 1-54.4. male extremely in viable. RK3.
origin: Spontaneous. ofrtei- Information: On© allele each induced by CB.
discoverer: llobr, 25a 14. 1356 and X rays.
references: 1927, Ifyt Mag. Natarv. 65: 266, *um: unexpanded irregular
phenotype: Eyes somewhat smaller than normal; sur- location: 1-52.3.
face roaî. RK1. origin: Induced by 2-chloroethyl niethanesulfonate
(CB. 1506).
origin: X ray induced, d-iscov&rer: Fahmy, 1956.
discover**: Deaeree, reference*: 19S9, DES 33: 94.
MUTATIONS 259
phenotype: Wings usually unexpanded to some de- 184: 1084-85; 1960, Heredity 15: 87-96). Nonpro-
gree; if expanded, they are short, broad, and tein tryptophan accumulated (Green, 1959, Genetics
slightly drooping or divergent. Fertility reduced in 34: 564—72). Suppressed by alleles at the su(s)
both sexes. RK3. locus (Schultz and Bridges, 1932, Am. Naturalist
up: upheld 66: 323—32). Tryptophan pyrrolase of su(s) v
location: 1-41.0. differs kinetically from that of wild type (Marzluf).
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- Some brown pigment formed under conditions of par-
phenylalanine (CB. 3007). tial starvation (Tatum and Beadle, 1939, Biol.
discoverer: Fahmy, 1954. Bull. 77: 415—22). Larval Malpighian tubules pale
references: 1958, DIS 32: 77. yellow (Brehme and Demerec, 1942, Growth 6:
phenotype: Wings held upright. Viability and fer- 351-56). RK1.
tility good. RK1. cytology: Locus in or near 10A1-2 (Green, 1954,
other information: Two alleles induced by CB. 1528. Proc. Natl. Acad. Sci. U.S. 40: 92-99).
*ups: upright scutellars other information: Pseudoallelism at the v locus
location: 1-40.8. demonstrated by recombination between v and v36t
origin: Spontaneous. in which v is to the left of v36i (Green, 1954).
discoverer: Fahmy, 1955. V*
phenotype: Posterior scutellar bristles held verti- discoverer: Plunkett, 24g.
cally. Fly small. Eyes dull, small, and abnormally phenotype: Eye color as bright as v at hatching, but
shaped. Wings short and folded. Male sterile; via- darkens rapidly. Suppressed by alleles of su(s)
bility about 20 percent normal. RK2. (Green, 1952, Proc. Natl. Acad. Sci. U.S. 38: 300-
Upturned: see U 5). Larval Malpighian tubes pale yellow (Brehme
*upw: upward and Demerec, 1942, Growth 6: 351-56). RK2.
location: 2-62. V36f
discoverer: Bridges, 33k21. origin: Spontaneous.
phenotype: Wings turned up at tips. More extreme at discoverer: Williams, 36f.
higher temperatures. Veins sometimes have lumps. phenotype: Eye color may be slightly more yellow
RK3. than v. Not suppressed by alleles of su(s) (Green,
vq: see £*•"<? 1952, Proc. Natl. Acad. Sci. U.S. 38: 300-5). Fly
*us: undersized from starved larva does not form brown eye pigment
location: 1-52.5. (Green, 1954, Proc. Natl. Acad. Sci. U.S. 40: 92-
origin: X ray induced. 99). Malpighian tubes of larva pale yellow (Brehme
discoverer: Fahmy, 1956. and Demerec, 1942, Growth 6: 351-56). RK1.
references: 1959, DIS 33: 94. other information: Located to the right of v (Green,
phenotype: Body small. Viable and fertile. RK3. 1954).
other information: One allele each induced by CB. y48a
1506 and CB. 1528; two by X rays. origin: X ray induced.
UW: see Bg2 discoverer: Fox, 48a7.
v: vermilion references: 1948, DIS 22: 53.
location: 1-33.0. 1949, Genetics 34: 647—64.
origin: Spontaneous. Green, 1952, Proc. Natl. Acad. Sci. U.S. 38;
discoverer Morgan, 10k. 300-5.
references: Morgan and Bridges, 1916, Carnegie Barish and Fox, 1956, Genetics 41: 45-57.
Inst. Wash. Publ. No. 237: 27 (fig.). phenotype: Not suppressed by alleles of su(s). Eyes
Sturtevant and Beadle, 1939, An Introduction to of fly from partially starved larva contain no brown
Genetics, Saunders, p. 64 (fig.). pigment. Fly lacks an antigen produced by wild
phenotype: Eye color bright scarlet owing to ab- type; same antigen removed by rb*s*. v4** fly has
sence of brown ommochrome. Ocelli colorless. a new antigen not shared by rfr**a or wild type.
The combination v; bw has white eyes. Eye color RK1.
wild type in genetically v eyes of gynandromorph cytology: Salivary chromosomes normal.
mosaic for wild type and v tissue (Sturtevant, other information: v*Sa pseudoallelic to r 3 *' and
1932, Proc. Intern. Congr. Genet., 6th. Vol. 1: occupies a position to the left of it. v48* not re-
304—7). v eye disks develop wild-type pigmenta- combinationally separable from v (Barish and Fox,
tion when transplanted into wild-type larva (Beadle 19S45).
and Ephrussi, 1936, Proc. Natl. Acad. Sci. U.S. *v5Io
22: 536—40). The v+ hormone of Beadle and origin: X ray induced.
Ephrussi was identified as kynurenine (Butenandt, references: Green, 1952, Proc. Natl. Acad. Sci. U.S.
Weidel, and Becker, 1940, Naturwissenschaften 28: 38: 300-5.
63-64). Activity of the indueible enzyme (Rizki phenotype: Insuppressible v allele. RK1.
and Ri*ki, 1963, J. Ceil Biol. 17: 87-92) trypto- *y51b
phane pyrrolase, absent (Baglioni, 1959, Nature origin: Spontaneous.
references: Green, 1952, Proc. Natl. Acad. Sci. U.S. phenotype: Vibrissae form tufts of bristles beneath
38: 300-5. eyes. Overlaps wild type. RK2.
phenotype: Insuppressible vallele. RK1.
v 5Jc
references: Green, 1952, Proc. Natl. Acad. Sci. U.S.
38: 300-5.
phenotype: Insuppressible v allele. RK1.
*vSlg
discoverer: Edmondson, 51g.
references: Meyer and Edmondson, 1951, DIS 25: 74.
phenotype: Like v. RK1.
*y267-4
origin: X ray induced. vb: vibrissae
discoverer: Hoover, 35i. From Bridges and Brehme, 1944, Carnegie Inst. Wash.
phenotype: Semilethal. RK2A. Publ. No. 552: 212.
cytology: Associated with 'T(l;2)v™7-4 =
T(l;2)llA7-8;36 (Sutton).
v° f: vermilion of OHermann origin: X ray induced.
origin: X ray induced in In(l)dl-49. discoverer: Muller, 261.
discoverer: Offermann. other information: Associated with aberration.
references: 1935, DIS 3: 28. *Vc: Vortice
phenotype: Like v. RK1A. location: Autosomal.
other information: For practical purposes, insep- origin: Spontaneous.
arable from ln(l)dl-49 and a useful marker for that discoverer: Smith, 37c20.
inversion. references: Novitski, 1937, DIS 8: 10.
*Va: Venae abnormeis phenotype: Enhances dp/dp to give phenotype like
location: 2- (not located). hy. Homozygous lethal. RK3.
discoverer: Timofeeff-Ressovsky. ve: veinlet
references: 1927, Arch. Entwicklungsmech. Organ. location: 3-0.2.
109: 70-109 (fig.). origin: Spontaneous.
Roelofs, 1937, Genetica 19: 518-36. discoverer Duncan, 34a.
phenotype: Veins irregularly branched or interrupted. references: 1935, Am. Naturalist 69: 94-96 (fig.).
Heterozygote overlaps wild type in 50 percent of phenotype: Wing veins L3, L4, and L5 do not reach
flies. RK3. wing margins. Does not overlap wild type. Sup-
*vac: vacuolated pressed by px (Waddington) and su(ve). ve/ve/+
location: 1-58.5. inters exes are veinlet whereas ve/ve/+ triploids
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- are normal, according to Pipkin. Developmentally,
phenylalanine (CB. 3026). veins appear complete in prepupa but distal tips
discoverer: Fahmy, 1955. obliterated during contraction period [Waddington,
references: 1958, DIS 32: 77. 1939, Proc. Natl. Acad. Sci. U.S. 25: 305; 1940,
phenotype: Wings blistered; character varies from J. Genet. 41: 75-139 (fig.)]. RK1.
small vacuole to involvement of entire wing. At cytology: Placed between 61E2 and 62A6, on basis
least one wing affected in 95 percent of flies. Via- of its inclusion in Dt(3L)D = Df(3L)61E2-
bility and fertility good. RK2. Fl;62A4-6 from T(Y;2;3)D.
*voo; varied outspread
phenotype: Wings outspread. Eye color mottled
brown (possibly variegation for car). Male sterile
and short lived. RK3A.
cytology: Associated with ln(l)v@o = In(l)18C5-
6;19B7S.
varied outspread: see vac ve: veinlet

varnished: see vr From Duncan, 1935, Am. Naturalist 69: 94-96.
vb: vibrissae
location: 1-49.3. ve*
discoverer: Bridges, 25122. or i g i n: Sponta ne ous.
MUTATIONS 261
discoverer: Bertschmann, 54a. Edinburgh 58: 787-815). Haltere disks also small
references: 1955, DIS 29: 69-70. [Chen, 1929, J. Morphol. 47: 135-99 (fig.)].
phenotype: Wing veins L2, L3, L4, and L5 do not Goldschmidt [l935, Biol. Zentr. 55: 535-54; 1937,
reach wing margins, ve/ve2 male more extreme Univ. Calif. (Berkeley), Publ. Zool. 41: 277-82]
than female and tends to resemble ve2; female re- claimed that wings are more or less fully formed
sembles ve. RK1. and subsequently eroded by degeneration during
Vein: see Vn pupation. Waddington [l939, Proc. Natl. Acad. Sci.
Vein off: see Vno U.S. 25: 299-307; 1940, J. Genet. 41: 75-139
vein let: see ve (fig.)] found no evidence of erosion and concluded
veins longitudinally shortened: see vli that effect of the gene is during larval period and
*Vel: Velvet involves reduction in size of prospective wing area
location: 1- or 3- (rearrangement). and shift in position of line along which wing area
discoverer: Patterson, 1933. is folded out from imaginal disk. Temperatures of
phenotype: Hairs on eyes conspicuous. RK3A. 29°C or greater appreciably increase wing size
cytology: Associated with T(l;3)Vel; breakpoints (Harnly, 1936, Genetics 21: 84-103; Stanley, 1935,
unknown. J. Exptl. Zool. 69: 459-95). vg/+ with certain
*ven: venation Minutes shows scalloping of wings [.Green and
location: 3- [right arm associated with In(3R)P). Oliver, 1940, Genetics 25: 584-92 (fig.)J. vg/vg/+
origin: Spontaneous. has scalloped wings more often than vg/+ (Green,
discoverer: Bridges, 33gl8. 1946, Genetics 31: 1-20). RK1.
references: 1937, DIS 7: 17. cytology: Placed between 49D3 and 49E6, on basis
Bridges and Bridges, 1938, Genetics 23: 111—14. of its inclusion in both Dt(2R)vgB =Df(2R)49D3-
phenotype: Veins irregularly thickened and branched, 4;50A2-3 and Df(2R)vgD = Df(2R)49Cl-2;49E2-6
especially L3 and crossveins. Eyes bulging and (Morgan, Bridges, and Schultz, 1938, Carnegie
bright. Bristles gnarled. Body small. Often sterile. Inst. Wash. Year Book 37: 304—9).
RK3A.
Venae abnormeis: see Va
venation: see ven
venula: see vn/
vermilion: see v
verticals: see vt
*ves: vestigium
location: 1-1.4.
phenotype: Wings abnormal; vary from small and
curved to almost normal with cut-away inner
margins. Eyes slightly rough and abnormally
shaped. Male infertile; viability about 50 percent
normal. RK2.
vesiculated: see vs
vestar: see vst
vestigial: see vg
vestigium: see ves
vg: vestigial
location: 2-67.0. vg; vestigial
origin: Spontaneous. From Bridges and Morgan, 1919, Carnegie Inst. Wash.
discoverer: Morgan, 101. Publ, No. 278: 148.
Last. Wash. Publ. No. 278: 150 (fig.)-
Mohr, 1932, Proc. Intern. Congr. Genet., 6th. Vol. origin: Spontaneous in In(2R)Cy.
1: 190-212. discoverer: L. Ward, 1920.
phenotype: Wings reduced to vestiges; usually held references: 1923, Genetics 8: 276-300.
at right angles to body. Wing veins still visible. phenotype: Wings and halteres absent or reduced to
Halteres also reduced. Postscutellar bristles tiny knobs. Viability low. Female sterile and male
erect. Viability somewhat reduced. Final size of usually so. RK3A.
larva smaller than wild type; pupation is slightly * V gl?
later. Wing disks of late larva markedly smaller origin: X ray induced simultaneously with
than wild type (Auexbach, 1936, Trans. Roy. Soc.
discoverer: Ruch, 1931. phenotype: Wings nearly full length but heavily
references: Morgan, Bridges, and Schultz, 1938, scalloped and narrowed by excisions. Halteres and
Carnegie Inst. Wash. Year Book 37: 306. postscutellars normal. vga/vg intermediate be-
phenotype: Slightly more extreme than vg. Fully fer- tween the two homozygotes. Wings straplike and
tile. vgil/+has occasional nick in wing. smaller in female than in male. RK2.
M(2)S2V611 vg11/+ + haa small eyes, arclike
wings, and in 65 percent of flies, wing nicks.
RK2.
other information: Originally considered associated
with Dt(2Rya-S2vAH, but Bridges and Curry
showed the two phenotypes to be separable by re-
combination.
vg31c: see vgNo1
Vf£*^ ^ t * S66 V&^ ® ^
*yg33k
discoverer: Ives, 33k30.
references: Plough and Ives, 1934, DIS 1: 33.
1935, Genetics 20: 42-69.
phenotype: Like vg^01, but reported to have greater
dominance and greater variability in heterozygote.
RK2.
*Yg37g
discoverer: Poulson, 37g.
1939, DIS 12: 49.
phenotype: Weak allele. Homozygote has slight nick
at ends and occasionally at sides of wings. Pene- vg°: vestigial-antlered
trance better in male than female. RK2. From Bridges and Morgan, 1919, Carnegie Inst. Wash.
*yg40b Publ. No. 278: 212.
discoverer: Ives, 40b. vg s ; vestigial- Beaded
phenotype: Homozygote like vg. Considerable domi- discoverer: Bridges, 28dll.
nance in heterozygote. RK2. references: Morgan, Bridges, and Schultz, 1938,
*Vg40c Carnegie Inst. Wash. Year Book 37: 305-6.
origin: Spontaneous, phenotype: Heterozygote has terminal nicks and
discoverer: Buzzati-Traverso, 40c20. lateral incisions in a few male and still fewer fe-
references: 1940, DIS 13: 49. male wings. Homozygous lethal; ceases develop-
phenotype: Like vg. RK1. ment at about tenth hour. Mitosis abnormal; chro-
*vg48a mosomes seem sticky and form what appear to be
origin: Spontaneous derivative of vg. anaphase bridges [Bull, 1956, J. Exptl. Zool. 132:
discoverer: R. C. King, 48al. 467-508 (fig.)]. RK2A.
references: Poulson and King, 1948, DIS 22: 55. cytology: Associated with Dt(2R)vgB = Df(2R)49D3-
phenotype: Wings of 15 percent of homozygotes have 4;50A2-3.
slight nick between L3 and L4. vg48a/vg wings vgc: vestigial-Carved
scalloped at tips and usually along sides in fe- origin: X ray induced.
male, only at tips in male. Halteres and post- discoverer: Demerec, 28c3.
scutellars normal in all cases. RK3. references: Morgan, Bridges, and Schultz, 1938,
YgSlh2S Carnegie Inst. Wash. Year Book 37: 305-6.
origin: Spontaneous. phenotype: Heterozygote has terminal nicks or
discoverer: Ives, 51h25. lateral incisions in about one-third the flies. Ho-
references: 1952, DIS 26: 65. mozygous lethal. Most embryos show only partial
phenotype: Similar to vg but wings slightly larger. involution and retraction of larval head; mouthparts
RK1. distorted (Bull, 1952, Genetics 37: 569-70).
RK2A.
*vg°: vestigial-antlered cytology: Associated with Df(2R)vgc = Df(2R)49B2-
origin: Spontaneous (probably a derivative of vg). 3;49E7-F1.
discoverer: Morgan, 12j. *vg c "; vestigial-Clipp&d
references: Morgan and Bridges, 1919, Carnegie origin: Spontaneous.
fast. Wash. Publ. No. 278: 211 (fig.). discoverer: Robertson and Reeve, 1947.
MUTATIONS 263
references: Beatty, 1949, Proc. Roy. Soc. *vg": vestigial-nick

Edinburgh, B 63: 249-70. origin: Spontaneous.
phenotype: Female sterile. Oogenesis seems discoverer: Bridges, 15h7.
normal, but no eggs laid. RK2. references: Morgan and Bridges, 1919, Carnegie
vgD: vestigial-Depilate Inst. Wash. Publ. No. 278: 273 (fig.).
origin: Spontaneous. phenotype: Homozygous vgn is wild type; vgn/vg
discoverer: Bridges, 31a22. shows nicked wing tips or laterally incised wings.
references: Morgan, Bridges, and Schultz, 1938, Penetrance 82 percent in female and 45 percent in
Carnegie Inst. Wash. Year Book 37: 305—6. male. RK3.
phenotype: Heterozygote has hairs sparse over vgnG; vestigial-nick of Green
thorax except above wings. Hairs and bristles ab- origin: Spontaneous derivative of vg.
sent from legs except from tarsi. Postscutellars discoverer: Green, 40j26.
small and erect. Humeral bristles and patches re- references: 1941, DIS 14: 39.
duced. Wings have nicks in about one-third the 1946, Genetics 31: 1-20.
flies; wings smaller at higher temperatures LHarnly, phenotype: Homozygote usually wild type; wings
1940, Genetics 25: 521—33 (fig.)]. Homozygous occasionally nicked; vgnO/vg has notched wings;
lethal. In embryo, there is failure of head involu- vgnG/Df(2R)vgD has scalloped wings. vg"G en-
tion and retraction; components of mouth parts dis- hanced by certain Minutes so than an appreciable
sociate and develop on surface of everted larval proportion of homozygotes have notched or scal-
pharynx (Bull, 1952, Genetics 37: 569-70). RK2A. loped wings. RK3.
cytology: Associated with Df(2R)vgD = Df(2R)49Cl- vgn h vestigial-nicked
2;49E2-6. origin: Spontaneous derivative of vg.
*vgdn: vestigial-double notch discoverer: Mohr, 1926.
origin: Spontaneous derivative of vg. references: 1932, Proc. Intern. Congr. Genet., 6th.
discoverer: Nolte, 1942. Vol. 1: 190-212 (fig.).
references: 1944, DIS 18: 44. phenotype: Homozygote wild type. vg"'/vg shows
phenotype: Wings have one notch between veins L3 wing nicks in 27 percent of flies. Enhanced in
and L4 and a second between L4 and L5, but near homozygote, especially female, by Minutes (Green,
L4 and sometimes including the tip of L4. Halteres 1946, Genetics 31: 1-20). RK3.
and scute liars wild type. vgdtl/vg has straplike vgn 12
wings, often with bent ends and held at right origin: Spontaneous.
angles to body. RK1. discoverer: Mohler, 55b9.
*ygh; vestigial-hemithorax references: 1959, DIS 33: 98.
origin: Spontaneous derivative of vg. phenotype: Homozygote wild type, v ^ ^ / v g h a s
discoverer: Ludwig, 1936. terminal wing nicks in 20—30 percent of flies.
synonym: vg*10*". v£ n i 5 /vg n * r and vgni2/vgnw'J has larger nicks and
references: 1936, Verhandl. Deut. Zool. Ges. 38, scalloping. RK3.
Zool. Anz. Suppl. 9: 21—73 (fig.).
1937, DIS 7: 18.
Schultz, 1938, Arch. Entwicklungsmeeh. Organ.
138: 69-102 (fig.).
phenotype: Half of dorsal thorax missing in about 20
percent of flies. Wings resemble v^"" in 10—20
percent, the majority resemble vg. Developmen-
tally, hemithorax is caused by degeneration of a
dorsal mesothoracic imaginal disk. RK2.
*vghR: vestigial-hemithorax of Reck

origin: Spontaneous derivative of vg.
discoverer: Reck, 1937.
phenotype: Like vg*. RK2.
*vg': vestigial-Incised
discoverer. Bridges, 36d20.
references: Morgan, Bridges, and Schultz, 1938,
Carnegie Inst. Wash. Year Book 37: 305—6.
phenotype: Heterozygote shows nicked and incised
wings in one-third the flies. Homozygous lethal.
RK2A. ygno; vestigial-notched
cytology: Associated with D^R^g1 » D% 2R)49C2- From Mohr, 1932, Proc. Intern. Congr. Genet., 6th. Vol.
Dl;50A2-3. ]: 190-212.
vgno: vestigial-notched references: 1959, DIS 33: 98.

origin: Spontaneous derivative of vg. phenotype: Homozygous lethal. Heterozygote has
discoverer: Mohr, 1926. terminal wing nicks in up to 50 percent of flies.
references: 1932, Proc. Intern, Congr. Genet., 6th. vgnwt/vgnw like homozygous vgnw; sterile. RK2.
Vol. 1: 190-212 (fig.). vgP: vestigial-pennant
phenotype: Wings notched in 45 percent of homozy- origin: Spontaneous derivative of vg.
gotes. vg" o /+is wild type. vgn0/vg has ragged discoverer: Kerr, 3Oal5.
wings with terminal notches and lateral excisions references: Harnly, 1935, DIS 4: 14.
in all flies. RK2. 1936, J. Exptl. Zool. 74: 41-59 (fig.).
*ygHo1: vestigial-Notch phenotype: Homozygote usually wild type but occa-
origin: Recovered among progeny of heat-treated sionally has wing nicks. Postscutellars and hal-
parents. teres wild type. vgP/vg has narrow straplike
discoverer: Swigert, 31c. wings, larger in male than female. Size and form
synonym: vg31 c. vary with temperature. RK3 (vgP/vg RK2).
references: Plough and Ives, 1934, OIS 1: 32. *vgPK: vestigial-pennant of Kutschera
1935, Genetics 20: 42-69 (fig.). origin: Spontaneous derivative of vg.
phenotype: Wings of most heterozygotes have discoverer: Kutschera, 1955.
terminal nicks; variations influenced by tempera- references: Mainx, 1956, DIS 30: 77.
ture. Homozygote has only bent vestiges of wings. 1957, Z. Induktive Abstammungs- Vererbungslehre
Posts cute liars erect; trident pattern dark. Body 88: 286-88 (fig.).
often dwarfed. Viability about 50 percent wild phenotype: Homozygote wild type at 17°C, 25°, and
type. Homozygous female sterile. RK2. 28°; vgPK/vg normal at 30°, has straplike wings at
vgNo2 25°, and wings smaller at 17°. RK3.
discoverer: Plough, 31j.
synonym: vg31t.
references: Plough and Ives, 1934, DIS 1: 32.
1935, Genetics 20: 42-69 (fig.).
phenotype: Like vgNol. Heterozygote enhanced by
Minutes (Green, 1946, Genetics 31: 1-20). RK2.
other information: Bridges suggested that all vg
semidominants reported by Plough and Ives were
really vg^01 carried in the South Amherst stock in
a suppressed condition or linked with a lethal.
vgnP: vestigial-nipped
or i g i n: Spontane ous.
discoverer: E. M. Wallace, 38a5.
phenotype: Wings have terminal and sometimes vg*: vestigial-strap
lateral incisions- Overlaps wild type at 25°C but From Bridges and Morgan, 1919, Carnegie Inst. Wash.
not at 19°. Scalloping visible in prepupal wing bud Publ. No. 278: 200.
[Waddington, 1940, J. Genet. 41: 75-139 (fig.)].
RK2. *vg s : vestigial-strap
ygnw; vestigial-no wing origin: Spontaneous derivative of vg.
origin: Spontaneous. discoverer: Morgan, 12d.
discoverer Morgan, 1924. references: Morgan and Bridges, 1919, Carnegie
references: Morgan, Bridges, and Sturtevant, 1925, Inst. Wash. Publ. No. 278: 200 (fig.),
Bibliog. Genet. 2: 59 (fig.), 232. phenotype: Wings of homozygote narrow and strap-
phenotype; Wings of homozygote smaller than in vg; like but nearly full length; often held at right
often only small, bent hooks or knobs. Balancers angles to body. Halteres reduced in size. Post-
also knobs. Scutellar bristles erect. Fly often scutellars normal. RK2.
dwarf and viability low. Female usually sterile. *vg*2
Heterozygote occasionally has wing nicks; second origin: Spontaneous.
vein often shortened (in 73 percent). vgnw/vg discoverer: Williams, 56c.
similar to vg/vgj; vgnw/vgno has strongly incised references: 1956, DIS 30: 80.
narrow wings; vgnw/vgni has scalloped wings with phenotype: Wings incised and about one-fourth to
shortened L2 (Mohr, 1932, Proc. Intern. Congr. one-half normal width; length varies. RK2.
Genet., 6th. Vol. 1: 190-212). Larval wing buds vgs: vestigial-Snipped
reduced in size (Chen, 1929, J. Morphol. 47: 135- origin: X ray induced,
99). RK2. discoverer: Mullet, 1929.
*ygnwl: vestigial-no wing lethal references: 1930, J. Genet. 22: 299-334 (fig.).
origin; Spontaneous. Morgan, Bridges, and Schultz, 1938, Carnegie Inst.
discoverer: Mahler, 55b9. Wash. Year Book 37: 306.
MUTATIONS 265
phenotype: Heterozygote has wing nick in most phenotype: Second longitudinal wing vein always
males but only a few females. Homozygous lethal. has a sizeable gap. L4 often broken, L5 some-
RK3A. times also affected. Homozygous lethal. RK1A.
cytology: Associated with Df(2R)vgs = Df(2R)49B12- cytology: Associated with Tp(3)Vno =
C1;49F1S-SOA1. Tp(3)89E;93F;97A (Nicoletti and Lewis, 1960, DIS
vgU: yestigial-Ultra 34: 53).
origin: Gamma ray induced. vo-3: see e(dpv)
discoverer: Ives, J5131. vortex in chromosome 3: see e(dpv)
references: 1956, DIS 30: 72-73. Vortice: see Vc
phenotype: Heterozygote has greatly reduced wings; *vr; varnished
halteres like vg; some variability. Homozygous location: 3-44.
lethal. vgu/vg wings reduced to a single segment discoverer: Mohr, 20j22.
and halteres virtually absent. About 5 percent of references: Bridges and Morgan, 1923, Carnegie
vgu/vgD eclose; wing varies from bristled knob to Inst. Wash. Publ. No. 327: 237.
two or three small segments; scutellum bare and phenotype: Eyes small with fused facets. Female
half normal size; halteres extremely rudimentary. sterile. RK2.
RK1A.
cytology: Associated with In(2R)vgu = In(2R)49Cl-
2;50Cl-2.
Vi: seeM(l)Bld
Viability: see M(l)Bld
vibrissae: see vb
*vli: veins longitudinally shortened
discoverer: Buchman, 1936.
phenotype: Veins L2, LA, and L5 tend to be
shortened. Overlaps wild type. Semidominant.
RK3.
*Vn; Vein
location: 3-19.6.
discoverer Mohr, 28j21.
Vol. 1: 190-212.
1938, Avhandl. Norske Videnskaps-Akad. Oslo, I. vs: vesiculated
Mat.-Natur. Kl. 4: 1-7. From Evong, 1925, Z. Induktive Abstammungs-
Mohr and Mossige, 1942, Avhandl. Norske Vererbungslehre 39: 165-83.
Videnskaps-Akad. Oslo, I. Mat.-Natur. Kl. 7:
1-51. vs; vesiculated
phenotype: Vein L4 not complete. Wings slightly location: 1-16.3.
spread. Fly smaller than normal. Homozygous origin: Spontaneous.
lethal. RK2A. discoverer: Mohr, 24c23.
cytology: Associated with Di(3L)Vn = Df(3L)64C12- references: 1927, Hereditas 9: 173.
D1;65D2-El. Evang, 1925, Z. Induktive Abstammungs-
*vnl: venula Vererbungslehre 39: 165—83 (fig.).
location: 2- (not located). phenotype: Wings warped, wrinkled, blistered, rough
origin: Spontaneous. textured, discolored, and divergent. May overlap
discoverer: Plaine, 50h. wild type. May be result of breakage of fibers that
references: 1951, DIS 25: 77. normally hold wing surfaces together during un-
phenotype: Extra veins between L3 and L4 largely folding (Waddington, 1939, Proc. Natl. Acad. Sci.
between anterior and posterior cross veins, some U.S. 25: 307). RK2.
also arise from L4 distal to posterior crossvein. cytology: Salivary chromosome location between
Penetrance in male 1.3 percent; in female 43 per- 5D3 and 6A2 (Demerec, Kaufmann, Fano, Sutton,
cent. With So, penetrance is 63 percent in female; and Sansome, 1942, Carnegie Inst. Wash. Year
expressivity also increased. RK3. Book 41: 191).
Vno: Vein off
location: 3- (rearrangement). * v s 29c
discoverer. E. H. Grell, 56c. discoverer: Oliver, 29c9.
references: 1959, DIS 33: 94. references: 1937, DIB 7: 19.
phenotype: Like vs but probably more extreme. At Sturtevant and Beadle, 1939, An Introduction to
25°C, no overlap of wild type; at 30°, 5 percent of Genetics, Saunders, p. 64 (fig.).
males and 12 percent of females overlap wild type phenotype: Eyes pure white. Ocelli, adult testes
(Green, 1939, DIS 11: 45). RK2. sheaths, and larval Malpighian tubules colorless.
Wild-type alleles not completely dominant to wy
origin: Induced by P32. w/+ has less red pigment than +/+ (Muller, 1935,
discoverer: R. C. King, 52a. J. Genet. 30: 407—14; Ziegler-Gunder and Hadorn,
references: 1952, DIS 26: 65. 1958, Z. Vererbungslehre 89: 235-45; Green,
phenotype: Wings wrinkled and blistered. Viability 1959, Proc. Natl. Acad. Sci. U.S. 45: 549-53). w
40 percent wild type. RK2, adult has very little, if any, pteridine (Hadorn and
Mitchell, 1951, Proc. Natl. Acad. Sci. U.S. 37:
origin: Found among progeny of male treated with 650—65). Isoxanthopterin present in considerable
radio frequency, quantity during pupation but is eliminated during
discoverer: Mickey, 61 j. first 3 days of adult life (Hadorn, 1954, Experientia
synonym: bw6lh bubble wing 61j; 10: 483—84). The meconium contains more pteri-
references: 1963, DIS 38: 28. dine than wild type (Hadorn and Kursteiner, 1955,
1964, DIS 39: 58. Arch. Julius Klaus-Stift. Vererbungsforsch.
phenotype: Not described. Sozialanthropol. Rassenhyg. 30: 494—98). Opto-
motor response absent (Kalmus, 1943, J. Genet.
origin: X ray induced. 45: 206—13) but fly phototactic. Eye color devel-
discoverer: Mayo, 1964. opment autonomous in larval optic disk trans-
references: 1966, DIS 41: 58. planted into wild-type host (Beadle and Ephrussi,
phenotype: One or both wings crumpled and partially 1936, Genetics 21: 230). w is a dominant sup-
expanded; occasionally blistered but not otherwise pressor of z (Gans, 1953, Bull. Biol. France Belg.,
affected. Penetrance about 77 percent. No differ- Suppl. 38: 1—90). w decreases ratio of diameter to
ence between sexes. RK2. thickness in spermathecae (Dobzhansky, 1924, Z.
*vst: vestar Induktive Abstammungs- Vererbungslehre 34: 245—
location: 2-4.3. 48; Dobzhansky and Holz, 1943, Genetics 28:
discoverer: Glass, 41115. 295-303). RK1.
references: 1944, DIS 18: 40. cytology: Placed in band 3C2 by Schultz and con-
phenotype: Wings small and straplike; variable. firmed by Lefevre and Wilkins, on basis of the w
Eyes small, very rough, and somewhat glazed. Fe- nonlethal phenotype of recombinant carrying left
male sterile. Viability low. RK3. end of In(l)w™4 = In(l)3Cl-2;20A and right end of
vf: verticals the 4-centric element of T(l;4)w™J = T(1;4)3C2-
location: 1-2.3. 3;20;102C (1966, Genetics 53: 175). Placed in
origin: Synthetic. 3C2-3 by Demerec and Sutton (Demerec, Kaufmann,
discoverer: Gersh. Fano, Sutton, and Sansome, 1942, Carnegie Inst.
references: 1965, Genetics 51: 477—80. Wash. Year Book 41: 191). Panshin also places it
phenotype: Anterior vertical, anterior dorsocentral, at 3C2-3 (1941, Dokl. Acad. Nauk SSSR 30: 5 7 -
and anterior scutellar bristles often missing, verti- 60). Location by Prokofyeva-Belgovskaya is at
cals being most likely to be affected. RK2. 3C3 (1941, DIS 15: 34-35).
cytology: Placed in 3C5-6, on basis of the v* pheno-
type of the following genotypes: Df(l)rata =
Df(l)3C3-4;3C6-7; the heterozygote between ec2
dP2
Df(l)t8t2 and the synthetic deficiency for 3C5 and
dp
6 produced by combining the XD4P element of CO X-16
T(l;4)w*5*-l* = T(lj4)3C4-5;101 and the 4DXP err X-6 chf ch
element of T(l;4Jf1264-l2 = T(l;4)3C6-7;101F; and cf s-4 a4 h
the synthetic deficiency for 3C5 and 6 produced by bl sat aS 02
combining the X*>4P element of T(l;4)w25«-1* with col a2 e
a recombinant between In(lLR)l-vl39 = Bwx bf a 1
ht(lLR)3C6-7 and the right end of a normal X chro-
mosome. 0.01 0.001 0.01 0.005
other information: Not known as a point mutation.
Map of the w locus
w: white Redrawn from Judd, 1964, Genetics 49: 253-65.
location: 1-1.5.
origin: Spontaneous. other information: First mutant found in Drosophila.
discoverer: Morgan, lOe. Member of a pseudoallelic series containing five
references: 1910, Science 32: 120-22. subloci that have been resolved by recombination
Morgan and Bridges, 1916, Carnegie Inst. Wash. as shown in map (Lewis, 1952, Proc. Natl. Acad.
Publ. No. 237: 25 (fig.). Sci. U.S. 38: 953—61; MacKendrick and Pontecorvo,
MUTATIONS 267
1952, Experientia 8: 390-91; Green, 1959, He- references: 1963, DIS 38: 29.
redity 13: 303-15; Judd, 1959, Genetics 44: 3 4 - phenotype: Like w. RK1A.
42). Mutants occupying the right two sites act as W63b
dominant suppressors of z; those occupying the origin: X ray induced.
left three do not (Green, 1959). w (e.g., w1) lo- synonym: wma: white-marbled.
cated to the right of wa; presumably occupies the references: Lefevre and Wilkins, 1966, Genetics 53:
same site as wc^. 175-87.
WJ1E4 phenotype: w63b/w has brownish eye color like
origin: X ray induced. waP/w. w63b/Di(l)w258-45 has a similar but
discoverer Gans. lighter eye color than waP/Df(l)w258-45€ jjale
references: 1953, Bull. Biol. France Belg., Suppl. lethal because w63b is inseparable from Df(l)N63b.
36: 1-90. RK2A.
phenotype: Eyes white. Suppresses z. RK1. cytology: Associated with Df(l)N63b =Df(l)3C2-
cytology: Salivary chromosomes apparently normal. 3;3E2-3.
W11G3: s e e Z11G3 *W64
*WUG2 origin: Spontaneous in FM6.
origin: X ray induced. discoverer Witten, 1964.
discoverer: Gans. phenotype: Like w. RK1A.
phenotype: Eyes variegated. RK2A. W64g3
cytology: Associated with T(l;2)w^3G2 - T(1;2)3C3- origin: Spontaneous.

5;56F, discoverer: Kidd, 1964.
w30: see we2 references: 1966, DIS 41: 60.
w3 2k; s e e wbf2 phenotype: Eye color dark carnation similar to g.
W33e31: s e e wdil RK1.
w33*: see wsat *w258-3
w40aHl origin: X ray induced.
origin: X ray induced in Dp(l;l)scVi. discoverer Demerec, 33h.
discoverer: Muller, Valencia, and Valencia, 1946-53. phenotype: Homozygous lethal. RK2A.
references: Valencia, 1966, DIS 41: 58. cytology: Associated with Dt(l)w2S8-3 =Dt(l)3B2-
phenotype: Male lethal, w affected. RK1A. 3.3C1-2.
cytology: Associated with but presumably inde-
pendent of ln(l)w40aHl = ln(l)lAl-C3;4C4-7;17B7- W258-11
8;18E2-3. origin: X ray induced.

*w48h discoverer. Demerec, 33j.
origin: Induced by mustard gas; derivative of wm4. phenotype: Lethal and cell lethal. Embryologically,
discoverer. Lindsley, 48hl3. the male is abnormal after 12 hr at 23°C; gut in-
references: 1949, DIS 23: 60. completely developed and mesoderm abnormal
phenotype: Like w. RK1A. (Poulson, 1940, Collecting Net 15: 172). RK2A.
cytology: No rearrangement other than ln(l)wm4 = cytology: Associated with Df(l)w2S8-ll =Df(l)3A2-
In(l)3Cl-2;20A. 3-.3C3-5 (Siizynska, 1938, Genetics 23: 291-99).
W258-12
*w48hS: white-48h of Schultz origin: X ray induced,

origin: X ray induced in inbred Oregon-R. discoverer. Demerec, 33j.
discoverer: Schultz, 48h. phenotype: Like w. RK1.
synonym: w*8** (preoccupied). cytology: Salivary chromosomes normal.
phenotype: Like w. RK1. *W258-14
*^5 To origin: X ray induced.
origin: Spontaneous in highly inbred y Oregon-R discoverer Demerec, 33k.
strain. phenotype: Lethal and cell lethal. Hemizygous male
discoverer Redfield, 51a. embryo abnormal after 12 hr at 23°C; gut incom-
references: 1952, DIS 26: 68. pletely developed and mesoderm abnormal
phenotype: Like w. RK1. (Poolson, 1940, Collecting Net 15: 172). RK2A.
w60 cytology: Associated with Df(l)w258-14 ^Dl(l)3A3-
origin: Spontaneous derivative of w* in 4;3Cl-2 CSlizynska, 1938, Genetics 23: 291-99).
Jn(7>cSi£Sc*<R+S, BCS1 @cs w» B. w2S8-42
discoverer: Hollander, 1960. origin: X ray induced.

references: 1960, DIS 34: 50. discoverer: Demerec, 38i.
phenotype: Like w. RK1A. cytology: Associated with Df(l)w2S8-42 *. Dt(l)3A6"
*w62d 8;3C3-S.
origin: Spontaneous derivative of w* in *w258-43
ln(l)^cSiLMC8R+s, ac$* «c« W* B. origin; X ray induced.
discoverer Mickey, 62d. discoverer: Demerec, 38k.
phenotype: Male lethal. RK2A. phenotype: Eyes of w/w/w+c triploids are reddish,
cytology: Associated with T(l;4)w25S~43 = not maroon as in w/w/w+o (see description of
T(J;4)3C3-5;102F4-5. w+ °). RK3.
w258-45 w +0 ; Oregon-R wild-type aliele of white
origin: X ray induced. origin: In Oregon-R wild type.
discoverer. Demerec, 381. discoverer Green.
phenotype: Development resembles w2S8-*l references: 1959, Proc. Natl. Acad. Sci. U.S. 45:
(Poulson). RK2A. 549-53.
cytology: Associated with Df(l)w258-45 =Df(l)3B4- 1959, Nature 184: 294.
Cl;3Cl-2 (Sutton) =£>f(l)3Cl-2;3C3-4 (Schultz). phenotype: Eyes of w/w/w+o triploids maroon; con-
Most recent analysis shows it to be Df(l)3B4- tain less red pigment than w/w/w+c. Amount of
Cl;3C2-3 (Lefevre and Wilkins, 1966, Genetics 53: pigment in diploid w+o/w less than w+c/w but
175-87). difference is not readily detected visually. Homo-
*w258-47 zygotes of w+c/w+c and iv + o / w + o contain same
origin: X ray induced, amount of red eye pigment. RK3.
discoverer Demerec, 39a. other information: The difference in dominance be-
phenotype: Like w. RK1. tween w + 0 and w + c is located to the right of the
cytology: Salivary chromosomes appear normal. wa sublocus. Crossing over is greater in the y-wch
w2S8-48 and wch-spl regions in the presence of w + o than
origin: X ray induced. w"*"Cj this difference is also located in the region
discoverer Demerec, 39c. to the right of the wa sublocus. w*O and w"*"C
cytology: Associated with Dt(l)w258-48 =Df(l)3A9- seem to have the same X-ray mutability (Green,
B1.3C1-2. 1960, Genet. Res. 1: 452-61).
*w2S8-49 w + R ; Russian wild-type aliele of white
origin: X ray induced. discoverer: Timofeeff-Ressovsky.
discoverer Demerec, 39c. references: 1932, Biol. Zentr. 52: 468—76.
phenotype: Like w. RK1. Muller, 1935, J. Genet. 30: 407-14.
cytology. Salivary chromosomes normal (Sutton). phenotype: Like w+A except by special tests.
*w2S8-S0 w/w/w+R triploids have pinkish eyes at hatching,
origin: X ray induced. which soon darken to normal red. Has greater de-
discoverer Demerec, 39c. gree of dominance than w+A (Muller, 1935). RK3.
phenotype: Like w. RK1. other information: Mutates less frequently to w when
cytology: Salivary chromosomes normal. irradiated than w+A (Timofe'eff-Ressovsky, 1932).
*W258-51 T
w : see Df(l)w'Ji
origin: X ray induced. W: see Df(l)w*J2
discoverer Demerec, 39k. m'N: see Df(l)w'J3
phenotype: Like w. RK1. w": white-apricot
* w 258-52 origin: Spontaneous.
origin: X ray induced. discoverer. Hues Us, 1923.
discoverer Demerec, 40a. references: Morgan, Bridges, and Sturtevant, 1925,
phenotype: Eyes white; texture rough. Not lethal. Bibliog. Genet. 2: 218.
RK2A. synonym: apr.
cytology: Associated with In(J)w2S8S2 =in(l)3C7- phenotype: Eyes of male yellowish with orange
9;8B1UF1 (Sutton). tone; female eyes yellower, somewhat lighter than
male. Larval Malpighian tubules colorless (Brehme
: American wild-type o//e/e of white and Demerec, 1942, Growth 6: 351—56). wa; bw
discoverer Timofe'eff-Ressovsky. slightly lighter than wa. wa; st light pinkish
references: 1932, Biol. Zentr. 52: 468-76. yellow (Mainx, 1938, Z. Induktive Abstammungs-
Muller, 1935, J. Genet. 30: 407-14. Vererbungslehre 75: 256—76), as is wa v. z w*
phenotype: Lower degree of dominance in w/w/+ lighter than either mutant alone, only slightly
triploids than in w*R. Eyes pinkish at eclosion, darker than wb{; therefore, w° does not suppress z
darken to maroon, but never become a normal red. (Green, 1959, Heredity 13: 303-15). w* rb and
RK3. w* g have nearly white eyes; w* wch, wb* wa, and
other information: Mutates more readily to w when w* su(f) all have white eyes. su(wa) w° and
irradiated than w+R (Timofe'eff-Ressovsky, 1932). suCw*)0 wa have browner eyes than w*. wBwx w*
is like wa (Judd). w*/+ has lighter eyes than +/+
+
C; Confon-S wild-type aliele of white in v homozygote (Braver, 1953, DIS 27: 86). Dark-
origin: In Canton-S wild type, ened by P. The amount of pigment formed as a
discoverer: Green, function of gene dose can be determined by use of
references: 1959, Proc. Natl. Acad. Sci. U.S. 45: duplications carrying w* and deficiencies: w* fe-
549-53. male < w0 male = w^/w3 female < w**/w* male ^
1959. Nature 184: 294. wa/wafw* female (Muller, 1932, Proc. Intern.
MUTATIONS 269
Congr. Genet., 6th. Vol. 1: 234). Eye color devel- *wa5Sk

opment autonomous in wa optic disks transplanted origin: X ray induced.
into wild-type hosts (Beadle and Ephrussi, 1936, discoverer: Clark.
Genetics 21: 230). RK1. synonym: apr^Sk,
other information: Occupies middle site in the w references: 1956, DIS 30: 71.
pseudoallelic series; i.e., to the right of wbt and phenotype: Apparently like wa. RK1.
to the left of wch. Gives rise to partial revertants, *wq57l
as wT (Muller), waM (Mossige), and waS7i origin: Spontaneous derivative of wa.
(Green). The white region of chromosomes carrying discoverer: Green, 57H.1.
wa pairs and crosses over regularly with nonhomol- synonym: waR57i.
ogous regions of the homologous chromosome; references: Rasmus on, Green, and Ewertson, I960,
specific regions of nonhomologous involvement Hereditas 46: 635-50.
characterize different homologous chromosomes Rasmus on and Rasmuson, 1961, Hereditas 47:
(Green, 1959, Genetics 44: 1243-56; Judd, 1961, 619-30.
Genetics 46: 1687-97). The products of nonhomol- phenotype: Eye color between wa and wild type.
ogous exchange are deficiencies; e.g., Df(l)wrG, Change from w* is greater for brown than for red
Df(l)wrJl, Df(l)wTJ2, and Df(l)wiJ3, and duplica- pigment. su(wa) decreases the amount of both red
tions; e.g., Dp(l;l)wrG and Dp(l;l)wrJ2. and brown pigment. Enhanced by su(f); not a sup-
w a2 pressor of z. RK2.
origin: Spontaneous. other information: Green (1959, Heredity 13: 303—
discoverer: Bridges, 1929. 15) was unable to recover recombinants with wa2
references: 1938, DIS 9: 114. or wa4.
phenotype: Eye color orange, slightly darker than *wo58/
wB. Eyes of male darker than female. Phenotype origin: Spontaneous derivative of wa.
of wa2 rb and wa2 g like rb or g alone; not affected discoverer: Green, 58112.
by su(w*) and does not suppress z (Green, 1959, references: Rasmuson, Green, and Ewertson, 1960,
Heredity 13: 303—15). Larval Malpighian tubes Hereditas 46: 635-50.
colorless (Brehme and Demerec, 1942, Growth 6: Rasmuson and Rasmuson, 1961, Hereditas 47:
351-56). RK1. 619-30.
other information: Occupies middle site of w pseudo- phenotype: Eye color between wa and wild type.
allelic series. Brown pigment at normal level; red pigment inter-
mediate. suCvi^) decreases brown and slightly in-
origin: Spontaneous. creases red pigment formation. Enhanced by su(f);
discoverer Curry, 34g2. not a suppressor of z. RK2.
references: 1938, DIS 9: 114. *wa59W
phenotype: Eyes brownish orange; slightly darker origin: X-ray-induced derivative of w9.
than either w* or wa2; very little sex difference. discoverer. Green, 59kl.
Eye color of w*3 rb and wa3 g like rb or g; not references: Rasmuson, Green, and Ewertson, 1960,
suppressed by su(w*) and does not suppress z Hereditas 46: 635-50.
(Green, 1959, Heredity 13: 303-15). Larval Mal- Rasmuson and Rasmuson, 1961, Hereditas 47:
pighian tubules colorless (Brehme and Demerec, 619-30.
1942, Growth 6: 351-56). RK1. phenotype: Eye color intermediate between wa and
other information: To the left of wch. Green (1959) wild type. Increase in brown pigment greater than
was unable to recover recombinants between wa3 red pigment over level of w°. Suppressed by
and either w*, w*2, or w*4 and concluded that they mtfw*); enhanced by suff); not a suppressor of z.
occupy the same site of the w pseudoallelic series. RK2.
wa3>; s e e w*1 *wo59k9
origin: X-ray-induced derivative of w a .
origin: Spontaneous. discoverer: Green, 59k9.
discoverer Nichols-Skoog, 35cl2. references: Rasmuson, Green, and Ewertson, 1960,
phenotype: Eyes of male yellowish orange, of fe- Hereditas 46: 635-50.
male lighter and more yellow, of both sexes paler Rasmuson and Rasmuson, 1961, Hereditas 47:
than w* and with less sex difference. Eyes of 619-30.
w*4 rb and wm4 g nearly white. Not suppressed by phenotype: Eye color intermediate between W and
&u(w*) and does not suppress x (Green, 1959, normal. More brown pigment than w*; red pigment
Heredity 13: 303-15). Larval Malpighian tubules virtually unchanged. Suppressed by stifw^J; en-
colorless (Brehme and Deuaerec, 1942, Growth 6: hanced by stiff); not a suppressor of z. RK1.
351-56). RK1. waS9k13
other information: To the left of wch. Green (1959) origin: Spontaneous derivative of w*.
was woable to recover recooabitMtnts between wm4 discoverer: Green, 59kl3.
and either w*>, w*»2, or w®$ and conelode-d that they references: Rasmuson, Green, and Ewertson, 1960,
occupy the I A M site of the w p*eiado*Uellc series. Hereditas 46: 635-50.
Rasmuson and Rasmuson, 1961, Hereditas 47: wbf3

phenotype: Eye color intermediate between wa and discoverer Curry, 36k9.
normal. With respect to the level in wa, brown pig- phenotype: Eye color buff, but slightly darker than
ment increased more than red. Red but not brown wb{. Larval Malpighian tubes pale yellow (Brehme
pigment formation increased by su(wa); enhanced and Demerec, 1942, Growth 6: 351-56). RK1.
by su(f); not a suppressor of z. RK2. wbl: white-blood
*Yfa60a5 origin: Spontaneous.
origin: Spontaneous derivative of wB. discoverer Hyde, 14glO.
discoverer: Sherwood, 60a5. references: 1916, Genetics 1: 535-80 (fig.).
references: Rasmuson, Green, and Ewertson, 1960, phenotype: Eyes yellowish ruby at hatching," darken-
Hereditas 46: 635-50. ing to sepialike with age; female lighter than male.
Rasmuson and Rasmuson, 1961, Hereditas 47: Not a suppressor of z (Green, 1959, Heredity 13:
619-30. 303—15). Sensitive to temperature; at 19°C eye
phenotype: Eye color intermediate between wa and color as dark as pn, at 30° as light as wbt or wl;
normal. More brown but not red pigment than wa. sensitivity greatest 40—48 hr after pupation
Enhanced by su(wa); not a suppressor of z. RK1. (Ephrussi and Herold, 1945, Genetics 30: 62-70).
w ': see we
aE Testes sheaths colorless. Larval Malpighian
woM: white-apricot of Mossige tubules pale yellow at 25°C (Brehme and Demerec,
origin: Spontaneous derivative of w a . 1942, Growth 6: 351-56). RK1.
d i scoverer: Mos sige. other information: Located to the left of w
synonym: wa^^. (MacKendrick and Pontecorvo, 1952, Experientia 8:
references: Rasmuson, Green, and Ewertson, 1960, 390-91) and w° (Judd, 1958, Proc. Intern. Congr.
Hereditas 46: 635-50. Genet., 10th Vol. 2: 137; Green, 1959). Judd
Rasmuson and Rasmuson, 1961, Hereditas 47: (1958) was unable to recover recombinants between
619-30. wb* and either wBwx or wa.
phenotype: Eye color intermediate between wa; con- wBwx; white-Brownex
siderably more brown pigment than in wa; red pig- origin: Spontaneous.
ment virtually unchanged. Suppressed by suCw8); discoverer: Mossige, 52a.
enhanced by su(f); not a suppressor of z. RK1. references: 1953, DIS 27: 59.
other information: Green (1959, Heredity 13: 303— Judd, 1959, Genetics 44: 34-42.
15) was unable to recover recombinants with wa2 phenotype: Eye color like bw; no sexual dimorphism.
and w*4. wBwx/+ duller and darker than wild type, from
w aR57j : see which it is readily distinguishable. Heterozygotes
waRM: s e e between wBwx and other w alleles or deficiencies,
wbf: white-buff e.g., Dt(l)Ns, are indistinguishable from
origin: Spontaneous. wBwx/wBwxm jj o t a suppressor of z (Judd, 1959).
discoverer: Safir, 15g28. The double mutant wBwx wco1 lighter than either
references: 1916, Genetics 1: 584-90. single mutant, but wBwx wa and wBwx wbt indis-
phenotype: Eyes light buff, lighter than w° male. tinguishable from wa and wbf, respectively. Testes
w 6 ' male eyes somewhat lighter than female. sheaths colorless. Larval Malpighian tubes pale
Lighter at 19°C than at 25°. wbl; at has white yellow. RK1.
eyes (Mainx). w^f rb and wbl g eyes are lighter other information: Occupies leftmost site in w
than w b/ , rb, or g (Green, 1959, Heredity 13: 3 0 3 - pseudoallelic series, to the left of wbt (Judd,
15). Larval Malpighian tubes colorless (Brehme 1957, Genetics 42: 379-80); 1959). Reduces re-
and Demerec, 1942, Growth 6: 351-56). RK1. combination in the y-spl interval.
other information: Occupies a re combinational site wef: white-coffee
between wBwx and w* in white pseudoallelic origin: X ray induced.
series (Judd, 1959, Genetics 44: 34-42; Green, discoverer: Nicoletti, 1960.
1959). Spontaneous reversions reported by Redfield references: I960, DIS 34: 52-53.
(1952, DIS 26: 68). phenotype: Eyes deep ruby at hatching, resemble
wbf2 waat, but darken greatly with age and resemble se
origin: X ray induced. in older fly. Female heterozygous for wc? and w,
discoverer: Oliver, 32kl6. w*, wc°, wcf>, wbl, wco1, or waat like wct/wcl,
synonym: w32*. wci/+ wild type. Larval Malpighian tubules bright
references: 1935, DIS 3: 28. yellow. RK1.
1935, DIS 4: 12. cytology: Salivary chromosomes appear normal.
phenotype: Eye color light buff, but slightly darker other information: Occupies a site in w pseudo-
than wbl or wbl$. No sexual dimorphism; not a allelic series very close to the left of w*
suppressor of * (Green, 1959, Heredity 13: 303- (Welshons and Nicoletti, 1963, DIS 38: 80).
15). Larval Malpighian tubules yellow (Brehme and wch: white-cherry
Dewerec, 1942, Growth 6: 351-56). RK1. origin: Spontaneous.
MUTATIONS 271
discoverer: Safir, 12j. references: 1964, DIS 39: 59.

references: 1913, Biol. Bull. 25: 4 5 - 5 1 . phenotype: Eyes reddish brown, lighter than wct,
Morgan and Bridges, 1916, Carnegie Inst. Wash. w^l, or wsat, more orange than wa^, and more red
Publ. No. 237: 51 (fig.). than wBwx, No sexual dimorphism. Not a sup-
phenotype: Eyes translucent pink, only slightly pressor of z. RK1.
yellowish; male distinctly lighter than female. other information: Located to the left of wa.
Ocelli pale; larval Malpighian tubes and adult *wd!l: white-dilute
testes sheaths colorless. Enhanced by P and origin: Spontaneous.
e(we). Eyes light in double mutant with rb or g, discoverer Ives, 33e31.
white with wa. Acts as dominant suppressor of z synonym: w33e31,
(Green, 1959, Heredity 13: 303-15). RK1. references: Plough and Ives, 1934, DIS 1: 31.
other information: Occupies a site to the right of wa phenotype: Eye color like weak pr, dilute red. Over-
(Green, 1959) and to the left of WSP (Lewis, 1956, laps wild type; not readily classified. Darkest
Genetics 41: 651). known w allele. Eye color of w/wdil between pn
*wch2 and g. RK2.
references: Green, 1959, Heredity 13: 303—15. wdP: white-deep purple
phenotype: Differs from wch in that eye color of origin: Induced by chloroethyl methanesulfonate
homozygous female is not darker than male; it is (CB. 1506).
insensitive to e(w°), and is not a suppressor of z. discoverer Auerbach, 1957.
RK1. references: 1957, DIS 31: 107-9.
other information: Located to the left of wch. Green, 1958, DIS 32: 88.
*wch41 i 1959, Heredity 13: 303-15.
origin: Spontaneous. phenotype: Eye color deep purple. No sexual dimor-
discoverer: Ives, 41j9. phism. No interaction with e(we). Does not sup-
references: 1942, DIS 16: 48. press z (Green, 1959). RK1.
phenotype: Resembles wch. RK1. other information: Located to the left of wch (Green,
w co : white-coral 1959).
discoverer: Lancefield, 1917. origin: Induced by chloroethyl methanesulfonate
references: 1918, Am. Naturalist 52: 264—69. (CB. 1506).
phenotype: Eyes of male deep ruby at eclosion, discoverer Auerbach, 1957.
darkening to garnet like with age. Eyes of female references: 1957, DIS 31: 107-9.
somewhat lighter. Not a suppressor of z. En- Green, 1958, DIS 32: 88.
hanced by e(we). Lightens rb and g (Green, 1959, 1959, Heredity 13: 303-15.
Heredity 13: 303—15). wco; st is yellow (Mainx, phenotype: Like wdP. RK1.
1938, Z. Induktive Abstammungs- Vererbungslehre other information: Located to left of wctl,
75: 256—76). Larval Malpighian tubules pale w*: white-eosin
yellow (Brehme and Demerec, 1942, Growth 6: origin: Spontaneous derivative of w.
351-56). RK1. discoverer Morgan, l l h .
other information: Originally shown to be to the left synonym: waE: white-apricot of Edinburgh; stock
of w by MacKendrick and Pontecorvo (1952, labeled wa from Edinburgh behaved like w°; prob-
Experientia 8: 390-91). Green (1959) was unable ably result of mislabeling (see MacKendrick, 1953,
to recover recombinants between wco and either DIS 27: 100).
iv* or w*2 and inferred that it occupies middle site references: Morgan and Bridges, 1916, Carnegie
of w pseudoallelic series. Inst. Wash. Publ. No. 237: 28.
yyco6 I Sturtevant and Beadle, 1939, An Introduction to
origin: Spontaneous derivative of w. Genetics, Saunders, p. 64 (fig.).
discoverer Hollander, 1961. phenotype: Eyes of female yellowish pink, male and
references: 1962, DIS 36: 78. w*/w female lighter. Using duplications and de-
phenotype: Like wao. RK2. ficiencies for we, Muller (1932, Proc. Intern.
wco/; white-colored Congr. Genet., 6th. Vol. 1: 233—35) showed that
origin: X ray induced. the eye pigmentation of we female = w* male <
discoverer: Demerec, 33j6. we/w" male = we/we female < w*/vr*fw* female.
synonym: w2SS~g. Enhanced by P, cru, wttg, and e(w0). Dominant
phenotype: Eye color varies in young male from suppressor of x; lightens rb and g (Green, 1959,
brick-red to dull brownish like pn; female slightly Heredity 13: 303—15). Eye color develops auton-
lighter. Lightens rb or g; not a suppressor of z omously in w® eye disks transplanted into wild-
(Green, 1959, Heredity 13: 303-15). RK1. type hosts (Beadle and Ephrussi, 1936, Genetics
other information: To the left of w*>2 (Green, 1959). 21: 230). Larval Malpighian tubes and adult testes
w c " ; w/»ffe-carro* sheaths colorless. RK1.
origin: Spontaneous. other information: Located to the right of w° (Green,
discoverer: Jodd, 1962. 1959).
W*2 discoverer: Sturtevant, 1918.

origin: Spontaneous derivative of w. references: Muller, 1920, J. Exptl. Zool. 31: 443-
discoverer: Hefner, 1925. 73.
synonym: w30. phenotype: Eyes very light buff or yellowish, lighter
phenotype: Similar to we but distinctly darker in in male than in female. Does not suppress z
both sexes. Less sex difference than in we. (Green, 1959, Heredity 13: 303-15).' Larval Mal-
Lightens rb and g; enhanced by e(wej. Dominant pighian tubules colorless (Brehme and Demerec,
suppressor of z (Green, 1959, Heredity 13: 303 — 1942, Growth 6: 351-56). RK1.
15). Larval Malpighian tubules colorless (Brehme other information: Located to left of w (MacKendrick,
and Demerec, 1942, Growth 6: 351-56). RK1. 1953, DIS 27: 100). Reverts to wild type at high
other information: Located to the right of wa (Green, frequency of 5 X 10~ 5 in female; much more rarely
1959). in male; frequency increased by X irradiation.
Also somatic reversions in patches of eye tissue;
origin: Spontaneous (extracted from a we rb stock). increased by X irradiation of young larvae (Lewis,
discoverer: Nolte, 1953. 1959, Genetics 44: 522). Bowman and Green (1964,
references: 1954, DIS 28: 77. Genetics 50: 237) find a spontaneous reversion
phenotype: Eye color of female slightly darker than rate of 0.25 X 10~ 5 in females heterozygous for a
that of we female. Eyes of male pinker than those deletion and in males; reversions not associated
of the we male. Amount of red pigment 3 times and with crossing over.
brown pigment 1.5 times that of we. RK1. wis: white-isoxanthopterinless
c
*w« : white-ecru origin: Spontaneous.
origin: Spontaneous. discoverer: Rasmuson.
discoverer: Muller, 1918. references: 1962, Hereditas 48: 587-611.
references: 1920, J. Exptl. Zool. 31: 443-73. phenotype: Eye color normal. Male has greatly re-
phenotype: Eyes very pale buff; between w1 and w. duced amount of isoxanthopterin in abdomen. Eye
RK1. color of z w' s male intermediate between z and
wild type and appears variegated. wch only par-
references: Green, 1959, Heredity 13: 303-15. tially suppresses z when heterozygous for wls.
phenotype: Like wec. Eye color same in male and RK3.
female. Lightens rb and g. RK1. other information: Located to the right of wa and
other information: Located to left of wch. probably to the left of wch. wa wctl/wia female
Wec3 yields recombinants between wa and wch with dif-
origin: X ray induced. ferent but characteristic intermediate production of
discoverer: Muller. isoxanthopterin. Postulated to be an intralocus
references: 1946, DIS 20: 68. duplication, based on its interaction with z. Prob-
phenotype: Eyes almost white. RK1. ably the same type of change as wrdP, wzl, and
WFM6: nhife in First Multiple wzm.
origin: Spontaneous in FM6 balancer. * w ml ; white-mottled
discoverer: Kidd, 1964. origin: X ray induced.
references: 1966, DIS 41: 60. discoverer: Muller, 1927.
phenotype: Typical white eyes. Useful as another references: 1930, J. Genet. 22: 299-334 (fig.).
marker in FM6. RK1A. phenotype: Eye color variegated. Variegates for N.
*wG: white of Goldschmidt Male lethal. Lines with dark eye color [an effect
origin; X ray induced in In(l)yG = In(l)lA;lC3-4. of an extra Y chromosome (Gowen and Gay, 1933,
discoverer Goldschmidt. Proc. Natl. Acad. Sci. U.S. 19: 122-26)] produce
references: 1945, Univ. Calif. (Berkeley) Publ. sterile males [T(1;3)W^/Y/Y\. RK2A.
Zool. 49: 522. cytology: Associated with T(l;3)wml; breakpoints
phenotype: Like w. RK1A unknown.
wh: white-honey other information: First case of variegated position
origin: Spontaneous derivative of w. effect recorded; originally termed eversporting dis-
discoverer: Dunn, 34j27. placement.
synonym: wa3. *wm2
references: 1935, Hereditas 21: 113-18. origin: X ray induced.
phenotype: Eye color between wbi and w*4; slightly discoverer Patterson, 1929.
darker in male than in female. Enhanced by e(w0). references: Muller, 1930, J. Genet. 22: 299-334.
Dominant suppressor of z (Green, 1959, Heredity phenotype: Eyes mottled. Male sterile. RK2A.
13: 303—15). Larval Malpighian tubules colorless. other information: Associated with T(l;3)wm2;
RK1. breakpoints unknown.
other information: Located to the right of wa (Green,
1959). origin: X ray induced.
w': white-ivory discoverer: Muller, 1929.
origin: Spontaneous. references: 1930, J. Genet. 22: 299-334.
MUTATIONS 273
phenotype: Eyes mottled. RK3A. discoverer: Ratty, 52bl2.

other information: Fragment of an X chromosome. references: Lefevre, 1953, DIS 27: 57.
The region of the chromosome from the right of fa cytology: Associated with T(l;2)wmS^bl2 -
locus to the proximal heterochromatin is deleted. T(l ;2)1E5-F1 ;3C3-4;20B;40-41.
Survives only in combination with normal X's; *wmS2bl3
mottled effect present when normal X's carry w. origin: X ray induced in In(l)rst3.
wm4 discoverer: Ratty, 52bl3.
origin: X ray induced. references: Lefevre, 1953, DIS 27: 57.
discoverer Muller, 1929. cytology: Associated with T(l;4)wm52b13 =
references: 1930, J. Genet. 22: 299-334. T(l;4)2A2-3;3C3-4;101 superimposed on In(l)rst3 =
phenotyne: Eyes mottled. Male and homozygous fe- In(l)3C3-4;20B.
male viable and fertile. RK2A. *y/m53a
cytology: Associated with In(l)wm4 =In(l)3Cl- origin: X ray induced.
2;20A (Sutton). discoverer: P. Farnsworth, 53a4.
W"5 references: Lefevre, 1953, DIS 27: 57.
origin: X ray induced. cytology: Associated with T(l;2)wm53a = T(1;2)3B2-
discoverer. Muller, 1929. C1;3C9-D1;4O-41.
references: 1930, J. Genet. 22: 299-334.
Bolen, 1931, Am. Naturalist 65: 417-22. origin: Neutron induced.
phenotype: Eyes reddish with white facets. Both discoverer: Mickey, 53ell.
sexes viable and fertile. RK2A. references: 1963, DIS 38: 29.
cytology: Associated with T(l;4)xv°*5 = T(1;4)3C3- cytology: Associated with T(l;2)wm53t> = T(1;2)3C3-
4;101Fl-2. 4;20A2-3;58F8-59Al.
*wml1 * y/rn 531
origin: X ray induced. origin: X ray induced in In(l)EN.
discoverer Panshin. discoverer M. A Bender, 53j.
references: Panshin and Khvostova, 1938, Biol. Zh. references: 1955, DIS 29: 69.
(Moscow) 7: 359-80. phenotype: Eyes of hemi- and homozygote mottled
Panshin, 1938, Nature 142: 837. with small dark brown patches. RK2A.
1941, DIS 15: 33-34. cytology. Associated with In(l)W"S3j =in(i)3C3-
cytology: Associated with T(l;4)w°*ll = T(1;4)3C3- 5;1A-2O superimposed on In(l)EN =
4;101A-D. In(l)lA;20;20B-C.
yfm49a y/m 541
origin: X ray induced. origin: Neutron induced.
discoverer Lefevre, 49a7. discoverer Mickey, 5413.
synonym: wmSP: white-mottled Spotted. references: 1963, DIS 38: 29.
references: 1949, DIS 23: 59. phenotype: Like wm<. RK2A.
1951, DIS 25: 71. cytology: Associated with InCiyw*1*4* = In(l)3C3-
Ratty, 1954, Genetics 39: 513-28. 5,20D.
phenotype: Eyes contain one or a few large red * w m55b
sectors on white background. Extra Y chromosome origin: X ray induced in R(l)2.
converts eyes to nearly wild type. RK2A. discoverer: M. A Bender, 55b.
cytology: Associated with T(l;3)wm4â = references: 1955, DIS 29: 69.
T(l;3)3At0-Bl;3E2-3;80. phenotype: Eyes mottled with large patches of pink
yym5 7 b or white. RK2A.
origin: X ray induced, cytology: Associated with ln(l)w^SSb z*In(l)3C3-
discoverer W. K. Baker, 51bl9. 5;1A-2O superimposed on R(l)2, which has opened
phenotype: Mottled for wand rst. RK2A. out as In(l)lA3-4;19F-20Al.
cytology: Associated with ln(l)w^Slb = ln(l)3Cl- Yftn258- J 8
2;20. origin: X ray induced.
discoverer Demerec, 33k.
origin: X ray induced in In(l)wm*. references: Demerec and Slizynska, 1937, Genetics
discoverer Lefevre, 51c20. 22: 641-49.
references: 1951, DIS 25: 71. phenotype: Eyes cream colored and mottled. Both
1951, DIS 26: 66. sexes viable and fertile. RK2A.
Ratty, 1954, Genetics 39: 513-28. cytology: Associated with T(l;4)w**258-IS =
phenotype: Eyes of w^Slc/w variegated. Male T(l;4)3C4-5;101.
lethal. RK2A. wm258~21
cytology: Associated with T(l;4)wmSlc « T(1;4)3C1- origin: X ray induced.
2;3C4-7;20A;10l. discoverer. Demerec, 1934.
*utWm52bl2 references: Judd, 1955, Genetics 40: 739-744.
origin: X ray induced in In(l)rat3. synonym: wVD3.
phenotype: Eyes and larval Malpighian tubes phenotype: Eyes mottled. Male viable. RK2A.
(Schultz) mottled. Heterozygous female shows cytology: Associated with T(l;2;3)wm258-44 -
Notch phenotype. Nearly lethal. Variegation sen- T(1;2;3)3C3-4;4D2-E1;56E1-F1;8OD.
sitive to temperature; more extreme at lower tem- *wm2S8-53
peratures than at 25°C. Variegation caused by origin: X ray induced.
proximity of the w locus to heterochromatin; non- discoverer: Demerec, 391.
variegating w alleles separable from rearrangement references: Sutton, 1940, Genetics 25: 628-35.
by recombination (Judd). RK2A. phenotype: Eyes mottled. Male viable. RK2A.
cytology: Associated with T(l;4)wm2 58-21 = cytology: Associated with T(l;4)w™258-53 -
T(1;4)3E5-6;1O1F. T(1;4)3C1-2;1O1E-F; 101F through 102F lost.
*wm2S8-31 *wm258-S4
discoverer Demerec, 371. discoverer: Sutton, 40e.
phenotype: Eyes cream colored and mottled for w. phenotype: Eyes cream colored and mottled. Male
Male viable. RK2A. lethal. RK2A.
cytology: Associated with T(l;4)wm2S8-31 = cytology: Associated with T(l;3)wm258-54 =
T(l;4)3C3-5;102F4-17. T(l;3)3B2-Cl;19F2-20Al;20E;63C7-8.
*wm258-32
origin: X ray induced. Y^m2 64-58
discoverer: Demerec, 371. origin: X ray induced simultaneously with N264'58.
phenotype: Eyes cream colored and mottled. Male discoverer: Demerec, 38d.
viable. RK2A. references: 1940, Genetics 25: 618-27.
cytology: Associated with T(l;3)wm2S8-32 - phenotype: Eye color variegated. Exists in three
T(1;3)3C3-5;81. types of lines: wm264'S8 from a lines shows ex-
*wm258-34 treme variegation and produces more pigment when
orig'in: X ray induced. paternally inherited than when maternally inherited;
discoverer Demerec, 38b. from / lines produces more fully pigmented eyes
phenotype: Eyes cream colored and mottled. Male and the converse parental effect; from g lines pro-
viable. RK2A. duces more pigment than f and no parental effect.
cytology: Associated with T(l;2)W»258-34 - In a lines, variegation partially suppressed by
T(1;2)3C3-S;41A. extra heterochromatin in genome (Baker and
Spofford, 1959, Univ. Texas Publ. 5914: 135-54)
origin: X ray induced. and in mother's genome (Spofford, 1959, Proc.
discoverer: Demerec, 38b. Natl. Acad. Sci. U.S. 45: 1003-7). Homozygous
references: Sutton, 1940, Genetics 25: 534—40. viable in female but apparently not in male. Varie-
phenotype: Eyes cream colored and mottled. Male gation less (phenotype more wild type) in homozy-
viable. RK2A. gous female than in heterozygous female. Hetero-
cytology: Associated with T(l;2)wm2S8-36 = zygous offspring of homozygous female less varie-
T(l;2)3C6-7;4C2-3;41A-B;41F5-6. gated than heterozygous offspring of heterozygous
*wm258-37 female (Spofford, 1958, Proc. Intern. Congr.
origin: X ray induced. Genet., 10th. Vol. 2: 270; Hessler, 1961, Genetics
discoverer: Demerec, 33j. 46: 463-84). When w^264-58 inherited from
phenotype: Eyes mottled. Male viable. RK2A. mother, more variegation than when inherited from
cytology: Associated with T(l;2)wta2S8-37 = father (Hessler, 1961; Spofford, 1961, Genetics 46:
T(l;2)3C3-4;40-4lA. 1151-67; Baker, 1963, Am. Zoologist 3: 57-69).
RK2A.
origin: X ray induced. cytology: Associated with T(1;3)N264~S8 =
discoverer: Demerec, 38e. T(l;3)3B2-3;3D6-7;80D-F as well as its derivative
phenotype: Eyes cream colored and mottled. Male Dp(l;3yN264-58 = Dp(l;3)3B2-3;3D6-7;80D-F. The
viable. RK2A. duplication has been used in most of the variega-
cytology: Associated with T(l;2)w™258-39 m tion studies.
T(l;2)3C3-5;40E-F.
*wm2S8~40 origin: X ray induced.
origin: X ray induced. discoverer: Patterson.
discoverer: Demerec, 38e. references: Griffen and Stone, 1938, Genetics 23:
phenotype: Eyes cream colored and mottled; texture 149.
rough. RK2A. phenotype: Eyes variegated. RK1A.
cytology. Associated with T(l;2)wt"2 58-40 - cytology: Associated with T(l;3)wm609e ~
7T1;2)3C3'5;41. T(l;3)3C2-3;100C3-4,
*wm2S8-44 wm4000
discoverer: Demerec, 38k. discoverer: Buzzati-Traverso, 4117.
MUTATIONS 275
references: 1943, Rend. 1st. Lombardo Sci. Lettere, wmM c; white-mottled of McLean
Pt. I: Class. Sci., Mat. e Nat. 77: 61-64. origin: X ray induced.
phenotype: Eyes cream colored, darker in male than discoverer: McLean.
female. Viable and fertile in male and homozygous references: Muller, 1946, DIS 20: 68.
female. Variegated for w, rst, fa, and Co. RK2A. phenotype: Eye color light mottled. Variegated for
cytology: A rearrangement with a break in white rst RK2A.
region. cytology: Associated with In(l)wmMc =ln(l)3Cl-
wma: see w63^ 2;20A-C.
*wmA: white-mottled from Austin *WmMed: white-mottled of Medvedev
origin: X ray induced. discoverer. Medvedev, 1934.
discoverer: Stone. phenotype: Mottling on wa background. RK2A.
references: Griffen and Stone, 1939, Genetics 24: cytology: Associated with T(l;4)wmMed; break-
73. points unknown.
1940, Texas Univ. Publ. 4032: 201-7 (fig.)- w"10; white-mottled orange
phenotype: Eyes variegated. Male viable and fertile. origin: Spontaneous.
RK1A. discoverer. Hanly.
cytology: Associated with T(l;4)wmA = T(1;4)3C2- references: 1963, DIS 38: 30.
3;101A2-3. Wright and Hanly, 1966, Science 152: 533-35.
*wmCi: white-mottled of Cicak phenotype: Eyes light mottled orange at eclosion,
origin: X ray induced in YSX'YL. darkening with age. Red pigments (drosopterins)
discoverer Cicak. reduced to about 10 percent normal; other pteri-
references: Oster, 1957, DIS 31: 150. dines about normal. wmo; bw is pale yellow, indi-
phenotype: Eyes variegated with red and white cating most ommochromes removed. Xanthine dehy-
facets. Introduction of additional Y chromosome drogenase about normal. Some males accumulate
does not seem to alter expression. RK2A. large quantities of drosopterins in abdominal fat
other information: Recombination reduced. body. These pigments appear about the second day
*wmDJ; white-mottled of Dubinin of adult life and disappear on the fourth, fifth, or
origin: X ray induced, sixth. Low temperature increases proportion of
discoverer. Dubinin. males with this trait. Female not affected. wmc>/w
references: Sacharov, 1936, Biol. Zh. (Moscow) 5: has dark homogeneous brown eye color; w m o /w a
293-302. and wmo/wBwx are similar but somewhat lighter.
cytology: Associated with T(l;2)wmD1 = Viability good. RK1.
T(l;2)3B;19-20;21 F. other information: Crossover tests give no indica-
tion of chromosome aberration, w010 probably be-
discoverer: Dubinin. longs to the waP subgroup of the w pseud oa lie lie
references: Sacharov, 1936, Biol. Zh. (Moscow) 5: series.
293-302.
cytology: Associated with T(l;4)w'"D3 =
T(l;4)3C;101. *wM: white of MacKendrick
*wmDGl: white-mottled of Dubinin and Goldat origin: Spontaneous derivative of w* (MacKendrick
discoverer: Dubinin and Goldat. believed it to be we, but she obtained it from
references: Sacharov, 1936, Biol. Zh. (Moscow) 5: Edinburgh, where subsequent results make it seem
293-302. likely w* and w* became interchanged).
cytology: Associated with rearrangement in which discoverer: MacKendrick, 1955.
3B is inserted into chromocenter. references: 1958, DIS 32: 82,
Rasmus on, Green, and Ewertson, 1960, Hereditas
*wmD V4: white-mottled of Dubinin and Vofofov 46: 635-40.
discoverer: Dubinin and Volotov. Rasmus on and Rasmuson, 1961, Hereditas 47:
references: Sacharov, 1936, Biol. Zh. (Moscow) 5: 619-30.
293-302. phenotype: Male and homozygous female alike and
phenotype: Eyes mottled; male lethal; heterozygous practically indistinguishable from wild type. Het-
female 2V. RK2A. eroarygote with w or deficiency for w has slightly
cytology: Associated with T(l;4)wtnDV4 = darker eyes than w*"3. Brown pigment at normal
T(1;4)3C3-7;3D;IQIA-D. level; red pigment intermediate between wfl and
wmJ: white-mottled of Jonsson normal. Enhanced by ®ufw*j and suff); not a sup-
origin: X ray induced, pressor of z. Viability and fertility good. RK2.
discoverer: Jonsson, 61128. *WMS9: white of Muller
references: Lefevre, 1963, DIS 37: 49-50. origin: Spontaneous.
Lefevre and Wilkins, 1966, Genetics 53: 175-87. discoverer: Muller, 59d.
phenotype: Eyes mottled red and white. RK2A. references: Mischaikow, 1959, DIS 33: 9«.
cytology: Associated with T(l;4)w«*J » T(t;4)3G2- phenofype: Eyes cream colored, darkening slightly
3;20;imC. with age. RK1.
*WP: white-pearl *w,3

discoverer: Steinberg, 37bl7. references: Green, 1959, Heredity 13: 303-15.
references: 1937, DIS 8: 11. phenotype: Eyes white. Does not suppress z. RK1.
phenotype: Eyes extremely pale, lighter than w*. other information: Located to the left of wch.
Larval Malpighian tubules colorless (Brehme and *ws4
Demerec, 1942, Growth 6: 351-56). RK1. origin: Spontaneous.
w: white-reddish references: Green, 1959, Heredity 13: 303—15.
origin: Spontaneous partial reversion from wa in phenotype: Eyes white. Does not suppress z. Not
sl
t sc sc B. affected Jby su(wB). RK1.
discoverer: Muller, 1944. other information: Located to the left of wa.
references: 1944, DIS 18: 57. *ws5
phenotype: Eye color nearly normal in homozygote; origin: Spontaneous.
dark maroon in wr/wa heterozygote. RK2A. references: Green, 1959, Heredity 13: 303-15.
wrdf: white-recombinant deficiency phenotype: Eyes white. Does not suppress z. RK1.
origin: Spontaneous product of asymmetrical ex- other information: Located to the left of wch.
change within the w locus; marker distribution *ws9
such that wrdt is also w°. origin: Spontaneous.
discoverer Judd, 1961. references: Green, 1959, Heredity 13: 303-15.
references: 1961, Proc. Natl. Acad. Sci. U.S. 47: phenotype: Eyes white. Does not suppress z. RK1.
545-50. other information: Located to the left of w c/l .
1964, Genetics 49: 253-65. ws10
1964, DIS 39: 59. origin: Spontaneous.

phenotype: Eyes white. Suppresses z. wrdt/wsP references: Green, 1959, Heredity 13: 303-15.
like w*P homozygous. RK1. phenotype: Eyes white. Not a suppressor of z. Not
other information: Deficiency for the site occupied affected by sa(w*). RK1.
by wcil in the w pseudoallelic series. Reciprocal other information: Located to left of wch.
asymmetric exchange product is wrdP. Recombina- w*at: white-satsuma
tion in w region reduced. Judd was unable to re- origin: Spontaneous.
cover recombinants between wTdi and w, w°, or discoverer: Bridges, 33126.
wBw*. synonym: w33t.
wrdP: white-recombinant duplication references: 1935, DIS 3: 18.
origin: Spontaneous product of asymmetrical ex- phenotype: Eye color deep ruby; resembles pr and
change within the w locus; marker distribution ma. No sexual dimorphism. Lightens rb and g. Not
such that w^P is also wbf. modified by P or e(w°) or by temperature. Does not
discoverer Judd, 1961. suppress z (Green, 1959, Heredity 13: 303-15).
references: 1961, Proc. Natl. Acad. Sci. U.S. 47: Larval Malpighian tubules nearly wild type
545-50. (Brehme and Demerec, 1942, Growth 6: 351—56).
1964, Genetics 49: 253-65. RK1.
1964, DIS 39: 59. other information: Occupies a site to the left of W>
phenotype: Eye color lighter than wbl. Enhancer of in the w pseudoallelic series (Green, 1959).
z; z + wrdP/z w+ has reddish brown mottled eyes. W*P: white-spotted
+
z wâ w""/z wbt wrdP female has white eyes. discoverer: Showell, 1944.
RK1. synonym: ap-w: spotted wfiite.
other information: Duplication for site occupied by references: Lewis, 1956, Genetics 41: 651.
wch in the w pseudoallelic series. Reciprocal phenotype: Eyes have fine-grained mottling; facets
asymmetric exchange product is wrdf. Increases range from yellowish to brown. Male darker than
exchange in y-spt interval and within the w locus. female (Green, 1959, Heredity 13: 303-15).
Probably same type of change as w**, w*l, and •w"P/w, w s P/w cJ) , and WBP/W** have homogeneous
w*">. brown eye color. waP heterozygous with a defi-
w r,dup : S ee Dpd;l)wrJ2 ciency for all or part of the w locus produces
*w *': white-spontaneous phenotype like W8P/WSP (Green, 1959, Z.
origin: Spontaneous. Vererbungslehre 90: 375-84). Suppressor of z
references: Green, 1959, Heredity 13: 303-15. (Green, 1959). The double mutant w*» WSP is white
phenotype: Eyes white. Does not suppress z. Not and wctt wBP has pale yellow eyes. RK1.
affected by su(w*). RK1. other information: Occupies rightmost site in the w
other information: Located to the left of wc*. pseudoallelic series, to the right of wch (Lewis,
1956).
origin: Spontaneous. W*P2
references: Green, 1959, Heredity 13: 303-15. origin: Spontaneous.
phenotype: Eves white. Does not suppress z. RK1. discoverer: Mohler, 56c22.
other information: Located to the left of wch. references: 1956, DIS 30: 78-79.
MUTATIONS 277
phenotype: Eyes mottled like waP, WSP2/W and phenotype: Eyes white. Not a suppressor of z.
wsp2/we have uniform brownish eyes. wsP2/w RK1.
s
slightly darker and more red than w P/w. RK1. other information: Located to left of wch.
wsp3 *WX3
discoverer: Green, 59a29. references: Green, 1959, Heredity 13: 303—15.
references: 1959, DIS 33: 94. phenotype: Eyes white. Not a suppressor of z. RK1.
phenotype: Like W S P. RK1. other information: Located to left of wch.
w': white-tinged * W X4
discoverer: Hyde, 14k2. references: Green, 1959, Heredity 13: 303—15.
references: 1916, Genetics 1: 535-80 (fig.). phenotype: Eyes white. Not a suppressor of z. RK1.
phenotype: Eye color light pinkish. Larval Mal- other information: Located to left of wch.
pighian tubes colorless (Brehme and Demerec, *WXS
1942, Growth 6: 351-56). RK1. origin: X ray induced.
wyC: white-variegated of Catcheside references: Green, 1959, Heredity 13: 303—15.
origin: X ray induced in R(l)2. phenotype: Eyes white. Not a suppressor of z. RK1.
discoverer: Catcheside. other information: Located to left of wctl.
phenotype: Eyes mottled; heterozygous female tends *WX6
to show N. Viability of male varies among lines origin: X ray induced.
from nearly zero to fair. RK2A. references: Green, 1959, Heredity 13: 303—15.
cytology: Associated with In(l)w^C =ln(l)3Cl- phenotype: Eyes white. Not a suppressor of z. Not
2;19-20 superimposed on R(l)2. affected by su(W). RK1.
w vco : white-variegated cobbled other information: Located to left of wa.
discoverer: Clausen, *WX8
phenotype: Eyes mostly white, with red mottling. origin: X ray induced.
RK2A. references: Green, 1959, Heredity 13: 303—15.
cytology: Associated with T(1;3)W™° = T(1;3)2B17- phenotype: Eyes white. Not a suppressor of z. RK1.
C1;3C4-5;77D3-5;81. other information: Located to left of wch.
*wvD I: white-variegated of Demerec * W X7 6
discoverer: Demerec, 33jl9. references: Green, 1959, Heredity 13: 303—15.
phenotype: Eyes variegated white and red. Male phenotype: Eyes white. A dominant suppressor of z.
fertile. RK2A. Not affected by su(w*). RK1.
cytology: Associated with Dp(l;4)wvD1 = other information: Located to the right of W.
Dp(l;4)3Cl-4;lQlA-D. wr'.' white-zeste light
*wvD2 origin: Spontaneous derivative of wzm.
origin: X ray induced. discoverer. Becker, 1958.
discoverer: Demerec, 33k27. synonym: zl.
phenotype: Fine-grained variegation of cream with references: 1959, DIS 33: 82.
dark spots. Female occasionally shows rst varie- 1960, Genetics 45: 519-34 (fig.).
gation. RK2A. Judd, 1963, Proc. Intern. Congr. Genet., 11th. Vol.
cytology: Associated with T(l;2;4)wvD2 = 1: 3-4.
T(l;2;4)3C4-5;18F;38;101A-C. 1964, DIS 39: 60.
*wvD4 phenotype: Eyes of z wzl female uniform lemon
origin: X ray induced. yellow at 25°-30°C; same at 14° but with large red
discoverer. Demerec, 33k2. spots, z wzl male raised at high temperature like
phenotype: Eyes of heterozygous female mottled. female raised at low temperature. At low tempera-
X/Y male mottled but rarely survives. X/Y/Y more ture, they have large red spots on lemon back-
viable, but sterile. RK2A. ground with salt-and-pepper mottling. X/0 and X/Y
cytology: Associated with T(l;2)wvD4 _ T(1;2)3D6- males identical. + w*1 is wild type, z w*V+ + is
El;40F. vaguely mottled. RK1.
wVD3; s e e w™258'21 other information: Postulated to be a duplication of
*WX1: white from X irradiation part of white locus. The white locus change is lo-
origin: X ray induced. cated to the right of w*. wxt is unstable and mu-
references: Green, 1959, Heredity 13: 303-15. tates to w*m and a white Qudd, 1963; 1964> Prob-
phenotype: Eyes white. Not a suppressor of z. ably same type of change as wia and wr<iP.
RK1. w*m: white-xeste mottled
other information: Located to left of wch. origin: Spontaneous product of asymmetrical ex-
*WX2 change within the w locus.
origin: X ray induced. discoverer: Green, 54k5.
references: Green, 1959, Heredity 13: 303-15. synonym: zm.
278 GENETIC VARIATIONS OF DROSOPMLA MELAHOGASTER
references: Becker, 1959, DIS 33: 82. other information: Allelism inferred from phenotype
I960, Genetics 45: 519-34 (fig.). and genetic location. One allele each induced by
Judd, 1963, Proc. Intern. Congr. Genet., 11th. Vol. CB. 1540, CB. 3025, and X rays.
1: 3-4. waisted: see ws
1964, DIS 39: 60. warped: see wp
phenotype: Eyes of z wzm female raised at 25°— warty: see wa
30°C are lemon yellow; at 14°—17°, slightly orange Washed eye: see We
with large red spots. Eyes of male raised at 25°— water wings: see wtw
30° have lemon-yellow background and fine-grained wavoid: see wd
red spots; at 14°—17° eye color of male almost wavy: see wy
normal. X/0 and X/Y males identical. Eye color waxy: see wx
of z+ wzm homozygote and hemizygote is wild *wd: wavoid
type. Eyes slightly mottled in + wzm/z wzm. RK1. location: 2-40.
cytology: Salivary chromosomes normal. origin: Spontaneous,
other information: wzm postulated to be a duplica- discoverer: Kellen-Piternick, 1941.
tion of part of the white locus that arose from an references: Kellen, 1945, Genetics 30: 12.
asymmetrical exchange. The change in the white phenotype: Wings waved. Variable penetrance and
locus lies to the right of wa. wzm is unstable and expressivity, especially in male. Partially sup-
mutates to wzt and to white. These forms are in pressed by y in both sexes. RK2.
turn unstable and revert to wztn (Judd, 1963; 1964). *wdn: wings down
Probably the same type of change as wis and wrdP, location: 3-100.
discoverer: Morgan.
W: Wrinkled references: 1929, Carnegie Inst. Wash. Publ. No.
location: 3-46.0. 399: 187.
origin: Recovered among progeny of female exposed phenotype: Wings extended and drooping or even
to stratosphere. directed ventrally, broad with close crossveins.
discoverer. Jolios, 1936. Overlaps wild type. Low viability. RK3.
references: 1936, Natl. Geograph. Soc. Tech. *we: wee
Papers, Stratosphere Ser. No. 2: 153-57. location: 1-3.
Jollos and Waletzky, 1937, DIS 8: 9. origin: X ray induced.
phenotype: Homozygote viable. Wings remain small discoverer: Muller, 2615.
and unexpended. Black spots on head beside pro- references: 1935, DIS 3: 30.
boscis or ocelli. Heterozygous female like homo- phenotype: Fly dwarfed. Eyes rough; bristles fine;
zygote but less extreme. Male much less extreme; and wings spread. Fertility very low. RK2.
wings often expanded but wrinkled, blistered, and *We: Washed eye
surface finely pebbled and grayish; no overlap with location: 3-43.0.
wild type. Suppressed by D in male and nearly so origin: Spontaneous.
in female. From prepupal stage through adult, wing discoverer: Andres, 42e7.
bases abnormally narrow, possibly preventing flow references: 1943, DIS 17: 48.
of body fluid in sufficient quantity to expand wings phenotype: Dominant modifier of w that produces
[Waddington, 1940, J. Genet. 41: 75-139 (fig.)]. partial reversion. Produces spot of dilute red pig-
RK1 as dominant. ment varying in size from dot to nearly whole eye.
W13: see T(1;4)A1 Homozygous lethal. Classification, fertility, and
k
wa: warty viability of heterozygote excellent. RK2.
location: 1-64.4 (based on location of wa2; wa said weoJt: see wk
to be near car). wee: see we
origin: Induced by P 3 2 . welt: see wt
discoverer: Bate man, 1950. welt I ike: see wt/
references: 1950, DIS 24: 56. *wgv: wing variance
phenotype: Eyes rough, with scattered enlarged location: 1-33.0 (no recombinants with v among
facets. Occasional notched wing tip. Penetrance 90S).
low. Viability variable. Male infertile in propor- discoverer: Fahmy.
tion to degree of expression. Heterozygous female references: 1959, DIS 33: 94.
often infertile. RK3. phenotype: Wing position variable; wings drooping,
outspread, or upheld. Male sterile. RK2.
origin: Induced by L«p-NN-di-(2-chloroethyl)amino- wh: whiskers
phenylalanine (CB. 3025). location: Autosomal.
discoverer. Fahmy, 1953. origin: Neutron induced,
references; 1958, DIS 32: 77. discoverer: Mickey, 54a7.
phenotype: Eyes irregularly roughened and of references: 1963, DIS 38: 29.
varying size ami shape; ommatidia deranged. Wing phenotype: Many extra vibrissae, which are longer
tips rarely notched. RK3. than normal. RK3.
MUTATIONS 279
whd: withered phenotype: Acrostichal hairs in irregular rows; in-
location: 2-61. complete whorls on thorax. RK3.
origin: Spontaneous. wo: white ocelli
discoverer: Bridges, 38a6. location: 3-76.2.
phenotype: Wings warped and waved or reduced to discoverer: Bridges, 12f21.
shrunken black pupal pads. RK2. references: 1920, Biol. Bull. 38: 231-36.
*whg: whiting Bridges and Morgan, 1923, Carnegie Inst. Wash.
location: Autosomal. Publ. No. 327: 66.
discoverer: Bridges, J3k21. phenotype: Ocelli colorless. Eye color wild type.
references: 1916, Genetics 1: 148. Modifies we to a lighter and less yellow tone.
1919, J. Exptl. Zool. 28: 337-84 (fig.). RK2.
phenotype: Specific modifier of we. we; whg has
pure white eyes. RK3.
*whh: white head
discoverer: Morgan, 13h.
phenotype: Ocelli surrounded by silvery patch. RK3.
whirly: see w/
whiskers: see wh
white: see w
white-marbled: see w63b
whiting: see whg
wi: witty eye
location: 2-54.9 (not allelic to rh).
discoverer: Whitten, 61 g.
phenotype: Eyes rough on lower half owing to irreg-
ular facets. Extra vibrissae in variable number and
distribution. Removal of closely linked modifiers
gives rise to dominant form. Penetrance and ex-
pression variable and highly sensitive to back- wp: warped
ground genotype. RK3. From Bridges and Morgan, 1923, Carnegie Inst. Wash.
wider wing: see ww Publ. No. 327: 216.
wing variance: see wgv
wings down: see wdn * w p : warped
*with: with trident location: 3-47.5.
location: 3- (near p). discoverer. Bridges, 19kl5.
discoverer: Morgan, 10a. references: Bridges and Morgan, 1923, Carnegie
references: Morgan and Bridges, 1919, J. Gen. Inst. Wash. Publ. No. 327: 215 (fig.).
Physiol. 1: 639-43. phenotype: Wings small and narrow, dusky, diver-
Bridges and Morgan, 1923, Carnegie Inst. Wash. gent, and warped. RK2.
Publ. No. 327: 31 (fig.). wr: see fwwr
phenotype: Dark trident pattern on mesonotum. Vari-
able, with some overlap of wild type. RK3. *¥tr: Vfrinkle
withered: see whd location: 2-76.
witty eye: see wi origin: Spontaneous,
wizened: see wz discoverer. Goldschmidt, 1933.
wk: weak synonym: Wrinkled (preoccupied),
location: 3-42. phenotype: Wings wrinkled and blistered. Homoary-
origin: Spontaneous, gote viable and only slightly more extreme than
discoverer Bridges, 33122. heterozygote. Development retarded. RK1.
phenotype: Bristles small, somewhat Minute, but Wrinkled: see W
variable. Abdomen disproportionately small. Wings Wrinkled: see Fr
somewhat warped. Viability variable. RK3. ws; waisted
*wl: whirly location: 1-1.0.
location: 2- (not located). origin: Induced by L^>-NN-di-(2-chloroethyl)®mino-
origin: Spontaneous, pheaylalanine (CB. 3025).
discoverer: Kill, 43k4. discoverer: Fahray, 1955.
phenotype: Anterior part of abdomen constricted, phenotype: Wings heavy textured, more opaque, and
giving appearance of long, narrow waist. Wings smaller than normal. Male completely sterile; fe-
held abnormally and surface wavy. Most flies die male fertile. RK2.
shortly after eclosion, but occasional male is wxwxt: waxy-waxtex
viable and fertile. RK3. origin: Spontaneous.
other information: One allele induced by CB. 1506. discoverer: R. F. Grell, 56k20.
wt: welt synonym: wxt,
location: 2-82. references: 1957, DIS 31: 81.
discoverer: Bridges, 32119. phenotype: Wings slightly spread and curved down
phenotype: Eyes small and narrow, with horizontal distally, texture heavy and waxy, tips pointed.
seam or welt. Many bristles, especially postver- First posterior wing cell narrow, second posterior
ticals, doubled or even quadrupled in number. Ab- cell broad and flared. Fertile in both sexes. RK2.
domen chunky. Occasional nicks in wing. Expres- other information: Allelism inferred from similarity
sion overlaps wild type at 19°C, but is excellent in phenotype and genetic location (2-69).
at 25° or higher. RK1.
*wtl: weltlike
location: 3-59.5.
phenotype: Eyes seamed and small. Aristae re-
duced. Wings rather broad. Female sterile. Ex-
pression better at 19°C. RK3.
*wtw: wofer wings
location: 1-38.9.
origin: Induced by DL-p-NN-di-(2-chlorethyl)amino-
phenotype: Wings short and broad, frequently with
incomplete cross veins, and often thickened owing
to separation of ventral and dorsal surfaces by
fluid. Eyes small and slightly rough. Male geni-
talia twisted; pigmentation of last abdominal seg- wy: wavy
ment in female patchy. Penetrance and viability From Nachtsheim, 1928, Z. Induktive Abstammungs-
low. Female infertile. RK3. Vererbungslehre 48: 245-58.
*wfwc": wafer wings-cleft end
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- wy: wavy
discoverer Fahmy, 1953. origin: Spontaneous.
synonym: cli. discoverer: Nachtsheim, 26g7.
references: 1958, DIS 32: 68. references: 1928, Z. Induktive Abstammungs-
phenotype: Last male abdominal segment grooved in Vererbungslehre 48: 245—58.
dorsal midline and with abnormal genitalia. Eyes phenotype: Wings transversely waved, usually
small; wings short, broad, and slightly divergent. turned up at tip. Abdomen long and narrow. Mar-
Female fertility low; viability good. Classification ginal vein kinked even when other characters over-
difficult. RK3. lap wild type. RK2.
other information: One allele induced by CB. 3007. cytology: Tentatively placed in 11D-E, on basis of
ww: wider wing the breakpoint of T(l;2;3)wy^74-a - T(1;2)8F-
location: 1-32.9. 9A;20A-B;26B-D + T(1;3)11D-E;65C-D.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- wy2
phenylalanine (CB. 3025). discoverer Ruch.
discoverer: Fahmy, 1953. synonym: ex-6.
references: 1958, DIS 32: 78. references: Parker, 1935, DIS 4: 62.
phenotype: Wings slightly shorter and broader than phenotype: More extreme than wy; more upward curl
normal, frequently upheld, and occasionally trun- to wings. RK2.
cated. Male viability and fertility good but female * wy ^ ® °
viability and fertility reduced. RK3. origin; Spontaneous.
other information: One allele induced by CB. 3026. discoverer. Haskell, 40a.
*wx: waxy references: 1941, DIS 14: 39.
location: 2-69.7. phenotype: More extreme than wy; more upward curl
origin: Spontaneous. towing. wy-'Oa/ivyis intermediate. RK2.
discoverer: Ives, 4Ikl5. *Wy274-2
MUTATIONS 281
discoverer: Demerec, 34a. *y3

phenotype: Male lethal. RK2A. origin: Spontaneous.
cytology: Associated with T(l;2;3)wy274-2 = discoverer: Morgan, 26a.
T(l;2)8F-9A;20A-B;26B-D + T(1;3)11D-E;65C-D. phenotype: Cuticle tannish. Bristles vary from dark
*wz: wizened brown to black, hairs from yellow to black. Larval
location: 3-47.8. mouth parts golden at basal prongs; lateral process
discoverer Bridges, 1921. and mouth hooks light. Classification possible in
synonym: shrunken-3. dissected but not in living larva (Brehme, 1941,
references: Bridges and Morgan, 1923, Carnegie Proc. Natl. Acad. Sci. U.S. 27: 254-61). RK1.
Inst. Wash. Publ. No. 327: 241. y3d: yellow-3 dark
phenotype: Small fly; not filled out. Body color dark origin: Spontaneous.
dull; bristles small. Late hatching. Infertile. RK3. discoverer: Sturtevant, 1933.
X-: see 1( )S phenotype: Wings gray like y2 but bristles yellow.
Xei: see apXa Larval mouth parts golden brown; mouth hooks light
y: yellow (Brehme, 1941, Proc. Natl. Acad. Sci. U.S. 27:
location: 1-0.0. 254-61). RK1.
origin: Spontaneous. y3Ai; yellow-3 of Muller
discoverer: E. M. Wallace, l l a . origin: Spontaneous in YL'SCS1.
references: Morgan and Bridges, 1916, Carnegie discoverer: Muller.
Inst. Wash. Publ. No. 237: 27. references: Muller and Valencia, 1947, DIS 21: 70.
phenotype: Body color yellow; hairs and bristles phenotype: Like y3. RK1A.
brown with yellow tips. Wing veins and hairs
yellow. Tyrosinase formed in adults (Horowitz). y3P. yellow-3 of Patterson
For the most part, y is autonomous in mosaics; origin: X ray induced.
i.e., a fly may show both yellow and nonyellow discoverer: Patterson, 31e25.
tissue; however, over limited distances there is synonym: y-'-'*,
some nonautonomy [Hannah, 1953, J. Exptl. Zool. references: 1934, DIS 1: 31.
123: 523—60 (fig.)]. Larval setae and mouth parts Stone, 1935, DIS 4: 62-63.
yellow to brown, hence distinguishable from the Muller and Prokofyeva, 1935, Proc. Natl. Acad.
dark brown of wild type (Brehme, 1937, Proc. Soc. Sci. U.S. 21: 16-26.
Exptl. Biol. Med. 37: 578-80; 1941, Proc. Natl. phenotype: Body tannish with black bristles. Var-
Acad. Sci. U.S. 27: 254-61). RK1. iegated with patches of yellow bristles and hairs,
cytology: Placed in region 1A5-8 on basis of its these patches being ac, slight Hw variegation.
being carried by the XD3P element of Larval mouth parts light at basal prongs. Classifi-
T(l;3)sc260-20 _ T(l;3)lA8-Bl;61Al-2 and by cation possible in dissected larva, more difficult
Dp(l;f)sc260-27 - Df(l;f)lA8-Bl;19F, but not being in living larva (Brehme, 1941, Proc. Natl. Acad.
lost from Df(l)260-5 =Df(l)lA4-5 (Sutton, 1943, Sci. U.S. 27: 254-61). RK1.
Genetics 28: 210-17). cytology: Associated with In(l)y3P - In(l)lA;20.
*ylS: yelloW'l Of Schultz Y4
discoverer: Schultz, 34kl5. discoverer Serebrovsky.
phenotype: Like y. Larval mouth parts like y references: Dubinin and Friesen, 1932, Biol. Zentr,
(Brehme, 1941, Proc. Natl. Acad. Sci. U.S. 27: 52: 147-62.
254-61). RK1. phenotype: Like y. RK1A.
Y2 cytology. Associated with In(l)y4 *>In(l)lA8-
origin; Spontaneous. Bl;l8A3-4.
discoverer; Bridges, 25J26. *yS
phenotype: Cuticle yellow. Hairs and bristles black. discoverer: Patterson.
Wings and veins gray. y2fy is like y2. y2/y35af phenotype: Male lethal. RK2A.
y2/yc4) a n ( j y2fybl a r e w iid type. Viability excel- cytology: Associated with In(l)y5 »In(l)lA-B;14D.
lent. Larval mouth parts slightly lighter than wild *y6
type at basal prongs, but not enough to enable re- origin: X ray induced in sci2.
liable classification (Brehme, 1941, Proc. Natl. phenotype: Body yellow; bristles brown with yellow
Acad. Sci. U.S. 27: 254-61). RK1. tips. Larval mouth parts like y (Brehme, 1941,
y2S: yellow-2 from Swedish Proc. Natl. Acad. Sci. U.S. 27: 254-61). RK1.
origin: Spontaneous. *y31h
discoverer: Bridges. origin: X ray induced in In(l)scs.
3
phenofype: Body color darker tan than y , but discoverer; Patterson, 31b.
bristles not so dark. Viability excellent. Larval phertotyoe: Like y. Shows some ac variegation. Vi-
mouth parts golden brown; mouth hooks and mentum ability good. Larval mouth parts light enough for
dark. Classifiable in living larva (Breiune, 1941, classification in living larva (Brehme, 1941, Proc.
Proc. Natl. Acad. Sci. U.S. 27: 254-61). RK1. Natl. Acad. Sci. U.S. 27; 254-61). RK1A.
*y31c yS3l
origin: X ray induced in In(l)sc8. origin: X ray induced in y+Y.
discoverer: Patterson, 31c. discoverer: Luning, 53i.
phenotype: Bristles dark as in y2 with some yellow synonym: yS3iY,
variegation. Larval mouth parts light at basal references: 1953, DIS 27: 58.
prongs. Classification difficult (Brehme, 1941, phenotype: y/y53iY a fertile y male. RK1A.
Proc. Natl. Acad. Sci. U.S. 27: 254-61). RK1A. yS4j
y3ld origin: Spontaneous.
origin: X ray induced in In(l)sc8. discoverer: Mohler, 55j24.
discoverer: Patterson, 3Id. references: 1956, DIS 30: 79.
references: 1935, DIS 4: 12. phenotype: Body and antennae yellow; bristles
Stone, 1935, DIS 4: 62-63. brownish (slightly darker than y). Wings nearly
phenotype: Similar to y2, but y31d/y35a is Ufa y31d wild type. Wings of y54i/y intermediate, but over-
whereas y2fy35a j S w iid type. Larval mouth parts laps y54j/y54). RK1.
light at basal prongs, but classification difficult yS9b
(Brehme, 1941, Proc. Natl. Acad. Sci. U.S. 27: origin: X-ray-induced derivative of y 2 . Arose as a
254-61). RK1A. mosaic in which half the descendants of the irra-
y3le: s e e y3P diated y2 gene were y^9b and half were y + .
y34c discoverer: Green.
origin: Spontaneous. references: 1961, Genetics 46: 1385-88.
discoverer: Curry, 34cl3. phenotype: Like y. yS9b/y2 1S wUd type, but y^9b
phenotype: Body color tan, very near wild type. Tan does not complement with y2 sc or other black-
antennae allow slow but reliable classification. bristled alleles of y. RK1.
Excellent viability. Larval mouth parts wild type yS9e
(Brehme). RK2. origin: Spontaneous.
*v35t discoverer: Clancy, 59c.
origin: X ray induced in In(l)A99b = In(l)lD3- references: 1960, DIS 34: 48.
E1;19D-E. phenotype: Like y, RK1.
discoverer: Stone, 35a. y62a
references: 1935, DIS 4: 62-63. origin: Spontaneous.
phenotype: Similar to y. y35a/y { . y35a/y2 discoverer: Ehrlich, 62a.
is wild type; y35a/y31d j s \]ê y31d ( or y2), La r v a i references: McCloskey, 1963, DIS 37: 50.
mouth parts golden. Classifiable in living larva phenotype: Bristles and hairs brown. Body yellow.
(Brehme, 1941, Proc. Natl. Acad. Sci. U.S. 27: Tip of male abdomen black. y62a dominant to
254-61). RK1A. y-type alleles and recessive to y^-type alleles.
*v39t RK1.
origin: Spontaneous. *y62b
discoverer Mather, 39el5. origin: Found among progeny of male treated with
references: 1941, DIS 14: 39. radio frequency.
phenotype: Body yellow; bristles brown. Not so discoverer: Mickey, 62b21.
light as y. RK1. references: 1963, DIS 38: 29.
* y40a y62k
origin: Spontaneous, origin: Spontaneous in In(l)scSiLsc8R+S,
discoverer: Buzzati-Traverso, 40a31. acSl &c& wa B.
references: 1940, DIS 13: 49. discoverer: Mickey, 62k8.
phenotype: Like y2. RK1. references: 1963, DIS 38: 29.
*ySO» y62kl9
origin: Spontaneous. origin: Spontaneous in In(l)scsl+dl-49.
discoverer Thoday. discoverer: Pratt, 62kl9.
references: 1954, DB 28: 78. phenotype: Like y. RK1A.
*ySlg *y94-J
origin: Spontaneous in inbred Oregon-R. origin: Spontaneous.
discoverer. Redfield, 51g. discoverer Moree, 46f6.
phenotype: Body yellow; hairs dark; bristles like 1947, DB 21: 69.
y2. RK1. phenotype: Like y. RK1.
*V5J< y260-4
origin: X ray induced in In(l}McstI'Sc8R+dl~49. origin: X ray induced.
discoverer Luning, 53el2. discoverer: Detnerec, 1938.
references: 1953, DIS 27: 58. references: Sutton, 1943, Genetics 28: 210-17.
phenotype: Homoxyecms lethal. RK2A. phenotype: Like y2. RK1.
other information: Not tested of 1(1 }J1 <w «c. cytology: Salivary chromosomes normal.
MUTATIONS 283
*y260-ll phenotype: Bristles yellow; body color wild type.

origin: X ray induced. ybl/y like yfa'; ybl/y2 w iid type. RK1A.
discoverer: Sutton, 39a. cytology: Associated with Dp(l;l)ybl = Dp(l;l)lB2-
references: 1943, Genetics 28: 210-17. 3;4F8-9;5D4-5.
phenotype: Like y. Male viable but sterile. RK2A. other information: ybi changes to y + and y. These
cytology: Associated with T(l;3)y26°-tl = events are more complicated than gene mutations;
T(l;3)lB2-3;85Fl-5. they involve duplication, chromosome rearrange-
*y260-12 ment, and mutation of neighboring genes such as
origin: X ray induced, sc and ac.
discoverer: Sutton, 1939. yc4: yellow-complementing
references: 1943, Genetics 28: 210—17. origin: Spontaneous in In(l)scsl+S.
phenotype: Like y. RK1. discoverer Muller.
cytology: Salivary chromosomes normal. synonym: ya (Muller, Ifi46; preoccupied); ySi (Green,
*y260-13 1961; error).
discoverer: Sutton, 1939. Frye, 1960, DIS 34: 49.
references: 1943, Genetics 28: 210—17. Green, 1961, Genetics 46: 1385-88.
phenotype: Body color wild type; bristles y. Male phenotype: Like y except bristles slightly darker.
fertility reduced. RK2A. yc4/ySl iê yc4. yc4/y2 wild type; however,
cytology: Associated with T(l;2)y260-13 = yc4/y2 sc like y2. Does not complement with other
T(1;2)1A4-5;36D. black-bristled alleles of y. RK1A.
*y260-21 *yG: yellow of Goldschmidt
discoverer: Sutton, 1939. discoverer: Goldschmidt.
references: 1943, Genetics 28: 210-17. synonym: yPx bh yellow-plexus blistered.
phenotype: Male lethal. y260-21/y nk e y. RK2A. references: 1945, Univ. Calif. (Berkeley) Publ.
cytology: Associated with T(l;3)y2^0-21 = Zool. 49: 307, 398-401.
T(l;3)6C;70E-F + In(l)lA6-7;SD8-Bl. phenotype: Like y2. RK1A.
*y260-24 cytology: Associated with In(l)yG = In(l)lA;lC3-4.
origin: X ray induced. *yH51: yellow from Hakozaki
discoverer: Sutton, 1939. origin: Spontaneous.
references: 1943, Genetics 28: 210—17. discoverer: Tanaka, 37e30.
phenotype: Like y. RK1. references: 1937, DIS 8: 11.
cytology: Salivary chromosomes normal. phenotype: Body, wings and legs yellow; bristles
*y260-28 and hairs black. Like y 2 . RK1.
origin: X ray induced simultaneously with ac 2 * 0 " 2 * *y w : yellow of Heuhaus
discoverer Sutton, 39126. origin: X ray induced.
references: 1943, Genetics 28: 210-17. discoverer: Neuhaus.
phenotype: Like y. Male viability reduced. RK2. references: 1936, DIS 5: 26.
cytology: Salivary chromosomes appear normal. phenotype: Bristles yellow; body color wild type.
*y260-30 yN/y is like yN; yN/y2 is wild type. RK1.
origin: X ray induced, *y°: yellow-orange
discoverer: Bishop, 1940. origin: Spontaneous.
references: Sutton, 1943, Genetics 28: 210—17. discoverer: Kill, 43kl8.
phenotype: Like y. RK1. references: 1946, DIS 20: 66.
cytology: Salivary chromosomes normal. phenotype: Body yellow; bristles, hairs dark. RK1.
other information: ac, sc, and svr not affected. yPS9: yellow of Perkovic
*y260-31 origin: Spontaneous in y+Y.
origin: X ray induced simultaneously with discoverer. Perkovic, 59h.
T(l; 2)260-31. references: Meyer, 1959, DIS 33: 97.
discoverer Fano, 1941. phenotype: Body and wings of y/yP59Y yellow;
references: Sutton, 1943, Genetics 28: 210—17. bristles dark. RK1A.
phenotype: Homozygous and hemizygous lethal. ypx bt: S ee y°
260- 3I/y y. RK2. *y*: yellow-spot
cytology: Salivary chromosomes normal at tip of X. origin: Spontaneous.
T(l;2)260-31 = T(1;2)9A;24;29 induced simulta- discoverer. Cattell, 12d.
neously. references: Morgan and Bridges, 1916, Carnegie
ybl: yellow-bristle Inst. Wash. Publ. No. 237: 33 (fig.).
origin: Spontaneous, phenotype: Large yellow spots on dorsal midline
discoverer Sandier. near tip of abdomen, on scutellum, and in narrow
references: Sandier, Hart, and Nicoletti, 1960, DIS stripe along thorax. Spots on scutellum and thorax
34: 103-4. not obvious except in presence of b. RK2.
y9: see yc4 two doses of the right portion of the white locus
yS1: yellow of Singh seem to be required for expression of zeste. Inter-
origin: Spontaneous in In(l)sc8. actions between duplications for z and w more
discoverer. Singh, 1940. complicated. Eye color develops autonomously in
references: 1940, DIS 13: 75. mosaics and from eye disks transplanted into wild-
phenotype: Like y. RK1A. type hosts. Eye color not affected by addition or
*yS6l: yellow ofShuman subtraction of F chromosomes. RK2.
origin: Spontaneous. cytology: Located in salivary chromosome band 3A3,
discoverer Shuman, 61 f. on basis of its inclusion in Df(l)w2S8-U =
references: Meyer, 1963, DIS 37: 51. Df(l)3A2-3;3C3-5 but not in Df(l)w2S8-14 =
phenotype: Like y. RK1. Dt(l)3A3-4;3Cl-2.
ys*: see yc4 Z11G3
yi<*: yellow-tanoid origin: X-ray-induced derivative of z.
origin: Spontaneous. discoverer: Gans.
discoverer. Spencer, 361. synonym: wl*G3.
references: Bridges, 1937, DIS 7: 16. references: 1953, Bull. Biol. France Belg., Suppl.
phenotype: Body color rich tan; antennae light 38: 1-90.
yellow; bristles black. Larval mouth parts golden phenotype: Eye color wild type. RK3.
brown. Classifiable in living larva (Brehme, 1941, cytology: Salivary chromosomes normal.
Proc. Natl. Acad. Sci. U.S. 27: 254-61). RK2. other information: Maps at z rather than w (Judd);
*yv1: yellow-variegated therefore a reversal of z rather than a suppressor
origin: X ray induced. of z at the w locus.
discoverer: Schultz, 33all. z°
phenotype: Variegated for y. RK2A. origin: X ray induced.
cytology: Associated with T(l;2)yvl = T(1;2)1A;39. discoverer: Gans.
yv2
phenotype: Eye color of both sexes wild type, but
origin: Spontaneous. zB/z female has yellow eyes like z/z. May be con-
discoverer: Schultz, 35kl. sidered a subliminal allele. RK3.
phenotype: Body color mostly wild type; head cytology: Salivary chromosomes normal.
bristles mostly black; thoracic bristles often zl: see wzl
yellow. X/0 male not more yellow than X/Y male. zm: see wzm
Larval mouth parts show basal prongs slightly *Z: Zerknitterf
lighter than wild type, the rest dark; not useful for location: 1-5.5.
classification (Brehme, 1941, Proc. Natl. Acad. discoverer: Gnineberg, 3Oh.
Sci. U.S. 27: 254-61). RK2. references: 1931, Biol. Zentr. 51: 219-25.
*yvS6
1934, DIS 2: 8.
origin: X ray induced in y*Y. phenotype: Wings crumpled or incompletely unfolded,
discoverer: C. Hinton and Schmidt. but majority overlap wild type. Viability 10 per-
references: 1956, DIS 30: 121. cent wild type. RK3.
phenotype: Variegates for y. Suppressed by extra Y zeste: see z
chromosomes. RK2A. Zw*: Zwischenferment-A
cytology: Not known to involve a rearrangement. location: 1-63 (T. Wright).
Ybb: s e e bbY origin: Naturally occurring allele.
yellow: see y discoverer: Young.
Ylt: see Pin?* references: Young, Porter, and Childs, 1964,
z: zeste Science 143: 140-41.
location: 1.0 (to the right of pn and kz). Young, 1966, J. Heredity 57: 58-60.
origin: Spontaneous. phenotype: Produces glucose 6-phosphate dehydro-
discoverer. Gans, 46b. genase that migrates faster in starch gel than that
references: 1948, DIS 22: 69-70. produced by ZwB. Zw*/ ZwB female produces a
Gans-David, 1949, Bull. Biol. France Belg. 83: slow- and a fast-migrating enzyme but no hybrid of
136-57. intermediate mobility. Enzyme level same in male
1953, Bull. Biol. France Belg., Suppl. 38: 1—90. and female. RK3.
phenotype: Male wild type. Eyes of female lemon Zw 8 ; Zwisckenfer merit-B
yellow at 25°C, variegated light yellow and origin: Naturally occurring allele.
brownish red at 19°. Ocelli have normal pigmenta- discoverer. Young.
tion (Welshons). Female heterozygous for a white references: Young, Porter, and Childs, 1964,
allele belonging to one of the two right-hand pseu- Science 143: 140-41.
doallelic subloci (e.g., w, w^h, we, w*P) is wild Young, 1966, J. Heredity 57: 58-60.
type. Male containing a w+ duplication is zeste; phenotype: Produces a slow-migrating glucose 6-
male with an intralocus duplication for one of the phosphate dehydrogenase. Enzyme level same in
right subloci (e.g., w*dP) has mottled eyes. Thus male and female. RK3.
Deficiencies
Duplications
Inversions
Rings
Tr a nslo cations
Transpositions
CHROMOSOME ABERRATIONS
ln(3R) Antp '/+
Le Calves, 1948. Bull. Biot. Franc© B#lg. 82: 97-113.

IDENTIFYING SYMBOLS. The standard chromo- Rings. Rejoining of breaks in opposite arms
some sequence or rearrangement is the one on of the same chromosome may give rise to a
which the standard genetic map and the standard ring-shaped chromosome. In ring designations
salivary gland chromosome map are based. Chro- the symbol is followed in parentheses by the
mosome breakage and reunion can give rise to chromosome involved and then by a specific
new chromosome sequences; i.e., chromosome designation; e.g., R(l)l. Ring-shaped Y chro-
aberrations. As the number of interacting breaks mosomes are described as Y derivatives in the
increases the number and complexity of possible section on special chromosomes. Replacement
chromosome aberrations become immense. of the older Xc harmonizes the terminology for
Rather than a descriptive name and symbol for rings with that of aberrations instead of mutants.
every type, chromosome aberrations are clas- Inversions. Intrachromosomal aberrations that
sified in terms of the elementary rearrangement are not rings and that have at least one section
deficiencies, duplications, inversions, rings, whose map order (either cytological or genetic)
translocations, and transpositions; these are ab- is inverted with respect to adjacent regions are
breviated Df, Dp, In, R, T, and Tp, respectively. designated inversions irrespective of whether
The abbreviation is followed parenthetically by segments are interchanged or lost. Inversions
the necessary chromosomal information and then may involve one arm (paracentric) or both arms
by a specific designation, which may be the sym- (pericentric) of a chromosome; they are sym-
bol of a mutant arising simultaneously with the bolized by the abbreviation followed parenthet-
aberration or simply an experiment number. Infor- ically by the chromosome arm or arms involved
mation on salivary chromosome breakpoints is and then by a specific designation; e.g.,
avoided to permit revising the description without In{2L)Cy, ln(2LR)bwvl. In(lLR) designates
altering the symbol. Except insofar as they are pericentric AT-chromosome inversions rather than
used to designate the associated mutant allele, Inp(l), which was formerly used, and 1L is im-
superscripts are not used in aberration desig- plied in In(l). Sometimes one break of a simple
nations. As with mutant symbols, aberration autosomal inversion is in the pericentric hetero-
symbols are always italicized and never contain chromatin and is not positioned with respect to
Greek letters, subscripts, or spaces. the centromere, so that whether the inversion is
Translocations. Translocations are rearrange- paracentric or pericentric is indeterminable. In
ments in which nonhomologous chromosomes such cases, the parenthetical information con-
interact irrespective of the number of breaks in- sists of chromosome number with no arm desig-
volved or of sequential changes or losses or nation. Recombination between similar inver-
gains of chromosome material within the partic- sions may produce viable recombinant inversions
ipating chromosomes. No distinction is made in with the left end of one and the right end of the
the symbol between simple reciprocal and more- other. Superscripts L and R are used to identify
complex translocations, nor are the involved arms the sources of the two ends; for example,
indicated in the parenthetical chromosomal in- In(2R)CyLbwVDelR.
formation. Participating chromosomes are A segment inserted into a new location is in
separated by semicolons and listed in the inverted order if its numerical order in salivary
following order: 1 (X), Y, 2, 3, and 4 [e.g., gland chromosome terminology is inverted with
T{1;Y;3)127, T(l;2;4)wvD2, and 7Ti;O£ 5 ]. The respect to the adjacent segments; it is in dys-
first distinguishing information in the trans- centric order if its polarity with respect to the
location symbol is within the parentheses. This centromere is altered. When a segment from a
chromosomal information is aligned on the left right arm is inserted into a right arm or left into
margin, the different classes being ordered nu- left, the inverted order is dyscentric; but in left-
merically according to this information (with to-right and right-to-left insertions, the two
the provision that Y falls between 1 and 2); terms are discordant, the inverted order being
within classes translocations are arranged al- eucentric instead of dyscentric. This distinction
phabetically according to specific designation. has not heretofore been made, the omission re-
Individual elements of a translocation are de- sulting in ambiguous descriptions of some aber-
noted by the superscripts D, P, and M. P (prox- rations.
imal) refers to the source of the centromere of Transpositions. Intrachromosomal aberrations
the element, D (distal) to the source of a termi- in which two noninserted segments are inter-
nus of different origin from the centromere, and changed are called transpositions; those in
M (medial) to the source of any material inter- which the order of the interchanged pieces is
calated between D and P [e.g., XD2P from undetermined are also considered transpositions
T{l;2)Bld, XD3M2P of T(l;2;3)Dinl until demonstrated otherwise. In transposition
CHROMOSOME ABERRATIONS 287
symbols the chromosome arm, or arms, involved ambiguous descriptions. Recombination within
is not indicated; e.g., Tp(3)bxd100. tandem repeats can lead to formation of tripli-
Deficiencies. Absence of a chromosome seg- cations and in successive steps to tandem re-
ment that produces a hypoploid genotype (either peats of order higher than three. Such high-order
hypodiploid or hypotriploid) is referred to as a repeats are also symbolized Dp.
deficiency. (There is some question about The elementary categories of chromosome aber-
whether a dipioid genotype with one and a rations are not mutually exclusive, and some
fraction X chromosomes should be considered a aberrations combine several of them. In such
hyperploid male or a hypoploid female and simi- cases the symbol used is the one that stands
larly whether a triploid genotype with two and a highest in the following ranking: T> R> In>
fraction X's should be considered a hyperploid Tp> Dp> Df. This is especially so when the
triploid intersex or a hypoploid triploid female. components are inseparable. A complicated
This question is usually resolvable by the sexual rearrangement may be separable genetically into
phenotype of the fly.) The symbol for deficiency its simpler component aberrations, which are
is followed in parentheses by the chromosome usually sufficiently designated with the distin-
number and arm and then by a specific designa- guishing symbol of the original aberration. When,
tion [e.g., DK2L)G; in D/(1)/Vs, 1L is implied however, the original is named after a phenotype
and in Df(4)M, 4R is implied]. Deficiencies may associated with one of the component aberra-
be formed by the deletion of material, either tions, designation of the other component with
interstitial or terminal, from a chromosome arm; the symbol of the mutant is inappropriate. A
they may also be synthesized in a number of rearrangement superimposed upon another re-
ways from preexisting aberrations. Terminal arrangement may be given a name, which more
duplication-deficiency products resulting from often than not refers to the entire complex since
aneuploid segregations from translocation hetero- the newly induced aberration is likely to be in-
zygotes are not ordinarily listed as either de- separable from the original; e.g., ln(2LR)SMl is
ficiencies or duplications but are discussed with a large pericentric inversion superimposed upon
the translocation entry. Other types of derived In(2L)Cy + In(2R)Cy. Component rearrangements
deficiencies and duplications are listed in the of synthetic combinations of aberrations are
appropriate sections, often with simply a refer- occasionally referred to individually, connected
ence to the aberration from which they were with a plus sign; for example, In(l)sc8 +
derived. In(l)dl-49 or In(2L)Cy + ln{2R)Cy. Collecting
Duplications. A genome that carries a chro- terms in much the same way as algebraic fac-
mosome segment in addition to the normal dipioid toring to further abbreviate the symbol is legit-
complement carries a duplication for the seg- imate; e.g., /n(l)sc*+ dl-49 and In(2L+2R)Cy.
ment. The symbol for duplication follows the Formerly, chromosomes with more than one in-
same plan as for other chromosome aberrations version were symbolized Ins( ); we use instead
except that the parenthetical chromosomal in- /n( ) for both singly and multiply inverted chro-
formation contains the chromosome of origin of mosomes since the presence of more than one
the duplicated segment listed first followed, inversion is indicated by the specific desig-
after a semicolon, by the recipient chromosome; nation; e.g., In(l)scslLsc8R+S. In describing a
e.g., Dp(3;l)O5, Dp(l;l)ybl. When the dupli- chromosome, inclusion of several types of in-
cated segment is carried as a free centric ele- formation is often desirable; e.g., sequence and
ment, the letter / (free) follows the semicolon gene content. Such categories are separated by
within the parentheses; e.g., Dp(l;Ol0l' A a comma followed by a space; e.g., ln(l)dl—49}
small chromosome segment duplicated in situ y w B, which designates an X chromosome
may be referred to as a repeat, even though it carrying the delta-49 inversion, the recessive
is still symbolized as a duplication; e.g., markers yellow and white, and the dominant
Dp{l;l)B* When the duplicated regions are in marker Bar. Marker genes are listed in the order
the same order, the term tandem repeat is suf- of the standard genetic map irrespective of their
ficient to specify accurately the new chromosome order on the chromosome in question. Three
if the limits of the duplicated segment are categories of information may be necessary to
known. When these regions are inverted with describe some special chromosomes; e.g.,
respect to each other, two reversed repeats are YSX-YL, ln{l)EN+dl-49, y B, where, besides
possible, making it necessary to specify which gene content and sequence, it is informative to
end of the segment is at the axis of symmetry of designate an abnormal combination of complete
the repeat; i.e., ABCCBD or ACBBCD. Failure chromosome elements.
to make such a distinction has given rise to
DESCRIPTIVE SYMBOLS. In addition to identi- Breaks rejoin cyclically to produce chromosome

fying symbols just discussed, aberrations are aberrations (e.g., A with B and B with A) and
given alternative descriptive symbols indicating multiple breaks may rejoin in more than a single
points of breakage on the salivary chromosome cycle. Thus four breaks may interact to form
map; breakpoints are listed in numerical order one four-break rearrangement or two two-break
according to the limits within which they must rearrangements. A complex rearrangement con-
lie. Each major chromosome arm is divided into sisting of two or more simple cyclic rearrange-
20 numbered divisions on the salivary gland ments is indicated in the descriptive symbol; e.g.,
chromosome map. The entire map is then num-
bered sequentially from 1 to 102 with 1—20*, T(2;3)OR72 = T(2;3)19E;29F + In(2LR)24F;54B
21^40-41-60, 61-80-81-100, and 101-102 or
representing X •, 2L-2R, 3L- 3R, and 4, re-
spectively, the centerpoints representing cen- T(1;2)C314 = T(l;2)5D;40-41
tromere positions. Each numbered division + T(1;2)9D;51D + T(l;2)20;56F.
is divided into six subdivisions designated by The order in which the component rearrangements
the letters A through F, each of which begins are listed in complex descriptive symbols fol-
with a heavily staining band; within the lettered lows the hierarchy according to which the identi-
subdivisions, the bands are numbered individ- fying symbol is determined. For a rearrangement
ually. Thus the complete designation of a par- superimposed upon a preexisting rearrangement,
ticular band consists of its numbered division, a similarly compound designation is used except
its lettered subdivision, and its number; e.g., that the plus symbol is replaced by the word on.
3C2. Positions of breakpoints are designated If one of the new associations of the preexisting
according to the bands between which or the rearrangement is broken by the superimposed
region or regions within which they are known to aberration, then the descriptive symbol is written
lie; for example, if a break lies between bands as though the entire aberration occurred at one
3C2 and 3C7 its position is designated 3C2-7; time rather than stepwise. An example is:
for the sake of brevity, the redundant information T(l;4)wm52b13, which was superimposed upon
3C is omitted from the second half of the no- In(l)tst3, is designated
tation. Less accurately determined breakpoints
may be given less specific designations; e.g., T(l;4)2A2-3;3C3-4;20B;101
3C or 3. An example of the total designation, since 20B, which was originally adjacent to 3C3,
both identifying and descriptive, is as follows: has become associated with 2A2 and 3C3 with
1K2;3)P - T(2;3)58E3-F2;60D14-E2;96B5-Cl, 101. A cyclic rearrangement was produced in-
items of chromosomal information being sep- volving both the preexisting breakpoints and the
arated by semicolons without spaces. Break- subsequently occurring ones; i.e., the symbol
points are listed in order, Y chromosome break- cannot be written as the old and the new cyclic
points designated Ys or YL or simply Y being rearrangements.
inserted between 20 and 21. Apparent terminal NEW ORDERS. In an aberration having only
deficiencies carry a single breakpoint desig- two breakpoints, the new order follows unambigu-
nation. Descriptions of incompletely analyzed ously from the descriptive symbol. In hetero-
rearrangements incorporate the known infor- chromatic rearrangements, however, an ambig-
mation. uity in the position of the breakpoint with respect
Descriptive symbols are used simply as a to the centromere may lead to ambiguities in or-
shorthand method for providing information about der. Thus, for example, T(l;2)8F;40-41 has chro-
the aberration; they supplement rather than sub- mosome 2 broken into two pieces, one extending
stitute for the identifying symbols. We have from 21 to 40 and the other from 41 to 60. Since
attempted to give breakpoints according to the it is not known which piece is centric, it is not
revised salivary gland chromosome maps pub- possible to state to which portion of chromosome
lished by C. B. and P. N. Bridges rather than 2 the acentric portion of the X extending from 1
according to C. B. Bridges's original maps, in to 8F is attached. With three or more break-
which individual bands were not numbered. No points more than one new order is possible;
special notation is used to designate doublet specifying the breakpoints is therefore not suf-
bands; the member of the doublet closer to the ficient to describe the aberration. We have
breakpoint alone is listed. Insofar as practical, adopted the following conventions for specifying
we avoid using breakpoint information in the sequences of aberrations. The sequence of
identifying symbol on the proposition that sub- each chromosome involved in an aberration is
sequent revision of cytological descriptions not specified from one end to the other according to
require alteration in the name of an aberration. salivary gland chromosome terminology. Points
CHROMOSOME ABERRATIONS 289
of breakage and reunion are indicated by vertical numerical value; e.g., |1A4 - 20-20F - 2OA1|
bars, and segments between these points are des- for R(l)2. In multiple-break rearrangements in
ignated by the most extreme band known to be which there is a break in autosomal hetero-
included at each end, separated by an em dash. chromatin whose position with respect to the
Thus the order of centromere is ambiguous, the new order may be
written in two ways depending on the position
T(2;3)P = T(2j3)58E3-F2;60D12-E2;96B5-Cl assumed for the heterochromatic break. In such
is represented as follows: cases, we have usually assumed (for the sake of
supplying the remainder of the new order) that
21 - 58E3|60E2 - 60F ; the heterochromatic break is in region 40 for
61 - 96B5|60D14 - 58F2|96C1 - 100 . breaks in chromosome 2 and 80 for breaks in
Were the order of the inserted segment 60D14 — chromosome 3.
58F2 not known, the segment would have been FORMAT. The chromosome aberrations are now
included within parentheses; i.e., listed in alphamerical order according to symbol,
61 - 95B5|(58F2 - 6OD14)|96C1 - 100 ; which is in bold face. Names, where necessary,
are listed (also in bold face) with cross-refer-
hierarchies of ambiguities are represented by ences to symbols; synonymic names and symbols
parentheses within parentheses. Salivary termi-
appear in body type with cross-references to
nology is not italicized except when part of an
current symbols. Each aberration description
aberration symbol, either identifying or descrip-
is written in the following format:
tive. Use of information on order depends only
on remembering that chromosome 1 extends from symbol: name
1 through 20 with the centromere in 20F, chro- cytology: The descriptive symbol as dis-
mosome 2 from 21 through 60 with the centromere cussed above.
between 40 and 41, chromosome 3 from 61 through new order: As discussed in the preceding
100 with the centromere between 80 and 81, and paragraphs.
chromosome 4 from 101 through 102 with the origin: The inducing agent is listed; aberra-
centromere in 101D. The first breakpoint in tions recovered from untreated parents are
T(2;3)P is listed as 58E3-F2; the first segment listed as spontaneous or naturally occurring,
indicated in the sequential formula goes through depending on whether recovered as a single
band 58E3, and the inserted segment begins with occurrence or repeatedly.
58F2. Nothing is implied about the position of discoverer: Name, date.
the intervening bands 58E4 to 58F1; unless they synonym: Alternative symbols or names, or
are specifically described as missing, they are both.
assumed to exist in association with one or the references: Sources of descriptions of the
other or both fragments produced by the break. aberrations listed in this section, although
Information on new order is written as follows: bibliographic information may appear under
each chromosomal element starts at the free end other categories as well.
with the lower value and the elements are listed genetics: Effects of the aberration on the
in numerical order, Y falling between 20 and 21. expression of genes near the breakpoints and
When desirable, the centromere position is phenotypic effects not yet attributable to
designated with a centerpoint; in special cases known genes are described. Segregational
where centromeres and breakpoints coincide, as and recombinational behavior may also be
is frequently true with ring-X chromosomes, the described. Descriptions of aneuploid deriv-
centerpoint replaces the vertical line. atives are also included in this category.
Rings are differentiated from rod-shaped chro- other information: In rare instances, infor-
mosomes by vertical bars at the beginning and mation not fitting into other categories is
end of the element; the circle is broken for linear included here.
designation at the breakpoint with the lowest
D«l)64j4
DEFICIENCIES cytology: Df(l)3A6-8;3Bl-2 (Judd) + Df(l)3B4-
C2;3Cl-4 (i.e., w258'45).
Del(l): see Dp(l;f) origin: Spontaneous in w^^*"*^-bearing X chromo-
): see Dp(l;f)R some.
discoverer: Judd, 64j4.
Df-3L«: see Df(3L)K genetics: Deficient for I(l)zw2 and I(l)zw3 but not
Df(l)0-sc,LVM: see D((1)260-1 I(l)zw4 or I(l)zw6. Male lethal.
*Df(\)7ak: Def/c/encyfJ) 7a from Austin *D«J)172
cytology: Df(l)3C3-5;3C7-9; inferred from origin: X ray induced.
Mackensen's fig. 15F (1935). discoverer: Patterson.
origin: X ray induced, references: 1932, Am. Naturalist 66: 193—206.
references: Mackensen, 1935, J. Heredity 26: 163- genetics: Deficient for pn, w, fa, and ec. Male
74 (fig.)- lethal.
genetics: Deficient for fa and spl but not w or ec; *Df(1)231c
female N. Male lethal. origin: X ray induced.
*Df(l)UzA discoverer: Patterson.
origin: X ray induced. references: 1932, Am. Naturalist 66: 193—206.
discoverer: Mackensen. genetics: Deficient for v. Male lethal.
references: 1935, J. Heredity 26: 163-74 (fig.). *D((1)235
genetics: Deficient for / but not fw or r. Male lethal. origin: X ray induced.
*D«J)24a discoverer: Patterson.
discoverer: Mackensen. genetics: Deficient for pn, w, fa, and ec. Male
references: 1935, J. Heredity 26: 163-74 (fig.), lethal.
genetics: Deficient for w but not pn or fa. Male *Dtfl)244
lethal. origin: X ray induced.
*D«l)60b discoverer: Patterson.
discoverer: Mackensen. genetics: Deficient for m. Male lethal.
references: 1935, J. Heredity 26: 163-74 (fig.). *D«l)247a
genetics: Deficient for f but not fw or r. Male lethal. origin: X ray induced.
D«l)62dl8 discoverer: Patterson.
cytology: Df(l)3B2-Cl;C3-5 (JMd). references: 1932, Am. Naturalist 66: 193-206.
origin: X ray induced. genetics: Deficient for m. Male lethal.
discoverer: Judd, 62dl8. *Df(l)247g
genetics: Deficient for I(l)zw6, I(l)zw7, and I(l)zw9 origin: X ray induced.
but not I(l)zw3. Lethal in male and in combination discoverer: Patterson.
with all alleles of I(l)zw6 except I(l)zw6lia. references: 1932, Am. Naturalist 66: 193—206.
Forms a viable heterozygote with In(l)wm4L genetics: Deficient for w. Male lethal.
rst3R =In(l)3Cl-2;20AL3C3-4;20BR (deficient for
3C2-3), which is w rst in phenotype. This combi- Df(l)26Q-l
nation should be homozygous deficient for 3C3 and cytology: Df(l)lB4-6; apparently a terminal
lethal [see l(l)3C3\; the discrepancy is unex- deficiency (Demerec and Hoover).
plained. origin: Spontaneous,
Df(l)62gl8 discoverer. L. V. Morgan, 1932.
origin: X ray induced. synonym: Df(l)0-8C,LVM.
discoverer Judd, 62gl8. references: Demerec and Hoover, 1936, J. Heredity
genetics: Deficient for z and l(l)zwl but not 27: 206-12 (fig.).
I(l)zw8. Male lethal. Sutton, 1943, Genetics 28: 213.
DKl)64c4 genetics: Deficient for y, ac, and sc but not svr.
cytology: Df(l)3A4-6;3C3-5 (Judd). Male lethal but not cell lethal (Ephmssi, 1934,
origin: X ray induced. Proc. Natl. Acad. Sci. U.S. 20: 420-22; Walen,
discoverer. Judd, 64c4. 1961, Genetics 46: 93-103).
genetics: Deficient for all known lethal loci between *Df(l)260>2
z and w. Also deficient for w but not for z. Male cytology: Df(l)lB2-3; apparently a terminal defi-
lethal. ciency (Demerec and Hoover).
DKl)64fl origin: X ray induced.
discoverer. Abrahams on, 64fl. references: Demerec and Hoover, 1936, J. Heredity
genetics: Deficient for J(ljzw3 and I(l)zw6 but not 27: 206-12 (fig.).
I(l)xw2 or I{l)zw7. Male lethal (Judd). Sutton, 1943, Genetics 28: 211.
CHROMOSOME A B E R R A T I O N S - DEFICIENCIES 291
genetics: Deficient for y and ac but not sc. Hemi- *Df(l)308

zygous lethal but not cell lethal. Embryo develops origin: X ray induced.
to fully formed larva but fails to hatch (Kaliss, discoverer: Patterson.
1939, Genetics 24: 244-70). references: 1932, Am. Naturalist 66: 193-206.
*D«l)260.5 genetics: Deficient for fa and ec. Male lethal.
cytology: Df(l)lA4-5; apparently a terminal *Df(l)314
deficiency (Demerec and Hoover). origin: X ray induced.
discoverer: Hoover, 1935. discoverer: Patterson.
references: Demerec and Hoover, 1936, J. Heredity references: 1932, Am. Naturalist 66: 193—206.
27: 206-12 (fig.). Mackensen, 1935, J. Heredity 26: 163-74 (fig.).
Sutton, 1943, Genetics 28: 214. genetics: Deficient for w, fa, and ec but not pn or
genetics: No phenotypic effect. Fertility and via- bi. Male lethal.
bility normal. *Df(l)354
*Df(l)260-10 origin: X ray induced.
cytology: Df(l)lA2-3; apparently a terminal discoverer: Patterson.
deficiency (Sutton). references: 1932, Am. Naturalist 66: 193-206.
origin: X ray induced. genetics: Deficient for pi. Male lethal.
discoverer: Sutton, 39a. *Df(l)Al: Deficiency^) from Austin
references: 1940, Genetics 25: 628—35. cytology: Df(l)9B;20.
genetics: Mutant for y and ac but not sc. Viable. origin: Aneuploid segregant from T(1;4)A1/+.
*Df(1)A12
*Df(l)260-19 cytology: Df(l)7A;7B.
cytology: Df(l)lA2-3; apparently a terminal origin: Aneuploid segregant from T(l;2;4)A12/+.
deficiency (Sutton). *Df(l)A124
origin: Spontaneous, cytology: Df(l)10A;13Al-2.
discoverer: Sutton, 1939. origin: Aneuploid segregant from T(1;2)A124/+.
references: 1940, Genetics 25: 628-35. Df(1)ac: Deficiency(T) achaete
1943, Genetics 28: 214. origin: X ray induced simultaneously with a detach-
genetics: No phenotypic effects. Both sexes viable ment of an attached X.
and fertile. discoverer: Muller.
*D«1)262 references: 1954, DIS 28: 146-47.
origin: X ray induced. genetics: Deficient for ac and probably y. Male
discoverer: Patterson. viable.
references: 1932, Am. Naturalist 66: 193—206. Df(1)B26^20: Deficiency(l) Bar
genetics: Deficient for fa; female TV. Male lethal. cytology: Df(l)15F9-16Al;16A6-Bl superimposed on
*DfCl)267 Dp(l;l)15F9-16Al;16A7-Bl.
origin: X ray induced. new order. 1 - 15F9|l6Bl - 20.-
discoverer: Patterson. origin: X ray induced in B chromosome.
references: 1932, Am. Naturalist 66: 193—206. discoverer: Demerec, 34a.
genetics: Deficient for fa; female N. Male lethal. references: Sutton, 1943, Genetics 28: 97—107
(fig-)-
*D«1)268 genetics: Reversion of B. Deficient for f but not as.
origin: X ray induced. Male lethal.
discoverer: Patterson. Df(l)bb: Deficiency^) bobbed
references: 1932, Am. Naturalist 66: 193—206. cytology: Df(l)20C;20D.
genetics: Deficient for car. Male lethal. origin: Associated with lîybtfl1.
*D«1)271
origin: X ray induced, Df(T)bbG: Deficiency(l) bobbed of Gershenson
discoverer: Patterson. cytology. Df(l)19F-20Cl;20B-Dl.
references: 1932, Am. Naturalist 66: 193—206. origin: Associated with ln(l)sc4Lsc8R.
genetics: Deficient for fa; female TV. Male lethal. Df(l)bbl-3°: Dcficiency(l) bobbed-lethal
Reduces crossing over. origin: X ray induced.
*Df(l)274 discoverer: Lindsley, Edington, and Von Halle.
discoverer: Patterson. genetics: Deficient for bb, Segregates irregularly
references: 1932, Am. Naturalist 66: 193—206. from y*Y in male. X/0 male lethal.
genetics: Deficient for /. Male lethal.
*DK1)3O3 origin: X ray induced.
origin: X ray induced. discoverer: Lindsley, Edington, and Von Halle.
discoverer: Patterson. references: 1960, Genetics 45: 1649-70.
references: 1932, Am. Naturalist 66: 193-206. genetics: Deficient for bb. Segregates irregularly
genetics: Deficient foe fa; female N. Male lethal. from y*Y in male. X/0 male lethal.
*Df(l)cmH2: Deficiency(l) carmine of Hannah

origin: X ray induced, cytology: Di(l)6D8-El;6E6-Fl (Hannah).
discoverer: Lindsley, Edington, and Von Halle. origin: X ray induced.
references: 1960, Genetics 45: 1649-70. discoverer: Hannah.
genetics: Deficient for 6b. Segregates irregularly genetics: Deficient for cm. Male lethal.
from y+Y in male. X/0 male lethal. *Df(J)cmH4
cytology: Df(l)6D8-El;6E6-Fl (Hannah).
discoverer: Lindsley, Edington, and Von Halle. discoverer: Hannah.
references: 1960, Genetics 45: 1649-70. genetics: Deficient for cm. Male lethal.
genetics: Deficient for bb. Segregates irregularly *Df(l)ct2o2: Deficiency^) cut
from y+Y in male. X/0 male lethal. cytology: Dt(l)7B3-6;7B6-7.
*D«l)bb'-*56 origin: X ray induced.
origin: X ray induced. discoverer. Hannah, 1947.
discoverer: Lindsley, Edington, and Von Halle, references: 1949, Proc. Intern. Congr. Genet., 8th.
references: 1960, Genetics 45: 1649-70. pp. 588-89.
genetics: Deficient for bb. Segregates irregularly genetics: Deficient for ct but not cm, sn, or oc.
from y+Y in male. X/0 male lethal. Male lethal.
cytology: Also carries In(l)481 = In(l)12E-F;14B. cytology: Df(l)7B2-3;7Cl-2.

discoverer: Lindsley, Edington, and Von Halle, discoverer: Hannah, 1947.
references: 1960, Genetics 45: 1649—70. references: 1949, Proc. Intern. Congr. Genet., 8th.
genetics: Deficient for 6ft. Segregates irregularly pp. 588-89.
from y+Y in male. X/0 male lethal. genetics: Deficient for ct but not cm, sn, or oc.
Df(l)bis: Deficiency(1) bistre Male lethal.
cytology: Df(l)7B5-6;7B7'8.
origin: Induced by DL-p-NN-di(2-chloroethy)amino- cytology: Dt(l)7B2-4;7C2-4.
phenylalanine (CB. 3007). origin: X ray induced.
discoverer. Fahmy, 1954. discoverer Hannah, 1947.
references: 1958, DIS 32: 67. references: 1949, Proc. Intern. Congr. Genet., 8th.
genetics: Deficient for bis. Male viable but sterile. pp. 588-89.
genetics: Deficient for cr but not cm, sn, or oc.
*Df(l)C-PL: Deficiency(l) C of Peterson and Male lethal.
Laughnan
cytology: Dt(l)15F;16E. cytology: Df(l)7A5-Bl;7C4-9.
origin: Spontaneous; allegedly an asymmetrical ex- origin: X ray induced.
change. discoverer: Hannah, 1947.
discoverer Peterson and Laughnan. references: 1949, Proc. Intern. Congr. Genet., 8th.
references: 1963, Proc. Natl. Acad. Sci. U.S. 50: pp. 588-89.
126-33. genetics: Deficient for ct but not cm, scp, an, or oc.
genetics: Deficient for f and B but not as. Male Male lethal.
lethal.
cytology: Df(l)6Fll-7Al;7B8-Cl.
cytology: Df(l)4A5~Bl;4D2-3 + Df(l)17A6-Bl;20C-D. origin: X ray induced.
origin: Associated with Infill1* btplR. discoverer: Hannah, 1947.
D{(l)CILy4R references: 1949, Proc. Intern. Congr. Genet., 8th.
cytology: Dt(l)17A6-Bl;18A3-4. pp. 588-89.
origin: Associated with In(l)ClLy4R. genetics: Deficient for cf but not cm, scp, or &n.
Male lethal.
DKl)cm**<>H4: Deficiency(l) carmine
cytology: D%1)6E. cytology. Dt(l)7B3-4;7B6-7 (questionable).
origin: X ray induced in R(l)2. origin: X ray induced.
discoverer Muller, Valencia, and Valencia, 1946-53. discoverer De Frank, 1947.
references: Valencia, 1966, DIS 41: 58. references: Hannah, 1949, Proc. Intern. Congr.
genetics: Deficient for cm. Genet, 8th. pp. 588—89.
genetics: Deficient for ct but not cm, scp, or sn.
*Dtfl)cmD5: Deficiency(l) carmine of De Frank Male lethal.
cytology: D%l)6E5-6;6F2-3.
origin: X ray induced. cytology: D%1)6D8-E1:7B7-C1.
discoverer: De Frank. origin: X ray induced,
genetics: Deficient for cm. Male lethal. discoverer Hannah, 1947.
CHROMOSOME ABERRATIONS - DEFICIENCIES 293
references: 1949, Proc. Intern. Congr. Genet., 8th. Di(l)Det lz A: see In(l)lzA
pp. 588-89. Df(l)Del271b: see T(l;2)271h
genetics: Deficient for ct but not cm or sn. 6E and DK\){2S7-S: Deficiency(l) forked
F may be transposed rather than lost; otherwise, it cytology: Df(l)15E7-Fl;15F2-4 (Sutton).
should be deficient for cm. Male lethal. origin: X ray induced.
discoverer: Demerec, 33k.
cytology: Df(l)7B2-3;7B6-7 (possibly). genetics: Deficient for /but not vb, r, or os. Male
origin: X ray induced. lethal and cell lethal.
discoverer Hannah, 1947. *Df(l)f2S7.6
references: 1949, Proc. Intern. Congr. Genet., 8th. cytology: Di(l)15E4-Fl;15F9-16Al;16A7-Bl.
pp. 588-89. new order: 1 - 15E4|l6Al - 20.
genetics: Deficient for cr but not cm, sn, or oc. origin: X ray induced in Dp(l;l)B -Dp(l;l)15F9-
Male lethal. 16A1;16A7-B1.
cytology: Df(l)7B2-3;7C3-4. references: Sutton, 1943, Genetics 28: 97-107
origin: X ray induced. (fig.)-
discoverer Hannah, 1947. genetics: Reversion of B. Deficient for f but not vb,
references: 1949, Proc. Intern. Congr. Genet., 8th. M(l)o, or os. Male lethal.
pp. 588—89.
genetics: Deficient for ct but not cm, sn, or oc. cytology: Df(l)15E7-Fl;16D2-4.
Male lethal. origin: X ray induced.
discoverer: Demerec, 34c.
cytology. Df(l)7B3-4,7B6"9. references: Sutton, 1943, Genetics 28: 97—107
origin: X ray induced. (fig)-
discoverer: Hannah, 1947. genetics: Deficient for f and B but not vb, un, lh, or
references: 1949, Proc. Intern. Congr. Genet., 8th. os. Male lethal.
pp. 588-89.
genetics: Deficient for ct but not cm, sn, or oc. *D«l)t257-27
Male lethal. cytology: D((l)14F6-15Al;15F5-6.
cytology: Di(l)7B2-4;7B6-7. discoverer: Demerec, 381.
origin: X ray induced. genetics: Deficient for f and M(l)o, but B not af-
discoverer Hannah, 1947. fected. Male lethal.
references: 1949, Proc. Intern. Congr. Genet., 8th. *Df(l)f2S7.28
pp. 588-89. cytology: Df(l)15E7-Fl;16E5-Fl.
genetics: Deficient for ct but not cm, sn, or oc. origin: X ray induced.
Male lethal. discoverer: Sutton, 40h.
genetics: Deficient for / and B regions. Male lethal.
cytology: Df(l)2E3-Fl;2F2-3 + D£(l)7B2-3;7B4-5 + *DKl)f2 57-31
Dt(l)19A4-5;19A6-Bl. cytology: Df(l)15E7-Fl;15F5-6.
origin: Associated with In(l)ct268-*3. origin: X ray induced,
discoverer Bishop, 41a.
cytology: D£(l)7B2-3;7B4-5. genetics: Deficient for /but not M(l)o or os. Male
origin: Associated with Jn(l)ct368-lSm lethal.
cytology: Di(l)6Fll-7Al;7B5-6. [>Kl)gh Deficiency(l) garnet-lethal

origin: Associated with In(l)ct26s-2°. cytology: Dt(l)12A;12E (Nicoletti).
41k origin: Spontaneous.
cytology. D%l)7B2-3;7C3-4. discoverer: L. V. Morgan, 24124.
origin: X my induced. genetics: Deficient for g and ty but not wy, s, pi, or
discoverer: Hoover, 38d.
sd. Lethal in male and cell lethal.
genetics: Deficient for ct but not acp or m. Male *D41)lz1: Dmiiciency(T) lozenge
lethal.
cytology: Df(l)7Ell-Fl;8El-2 (Hannah).
cytology. DI(1)5D2-3;7B2*3.
discoverer Green.
origin: AneaploAd segregmst from 7Yl;3)et:l68~37/+
references: Green and Green, 1956, Z. Induktive
Abstammungs- Vererbungslehxe 87: 708—21.
®%1)7A5~6;7M-Cl (Button),
genetics: Deficient for lz and sunx. Male lethal.
•rifift; X ray induced.
*DK1)lz2
*fMi*c, 40*.
cytology: Dl(l)8C14-D2;8E3-4 (Hannah).
*; D»flci*ctt for ct bat not cm, mcp, or m. origin: X ray induced,
Wmhe lttbtl.
discoverer Green.

Abstamraungs- Vererbungslehre 87: 708-21. origin: Spontaneous.
genetics: Deficient for lz and amx. Male lethal. discoverer: Eker, 36el2.
*D«1)lz3 genetics: Deficient for w. Male lethal.
cytology: Dt(l)8Cl-3;8D12-E2 (Hannah).
origin: X ray induced. cytology: Df(l)3C6-7;3D2-3 (Sutton).
discoverer: Green. origin: Spontaneous.
references: Green and Green, 1956, Z. Induktive discoverer: Ives, 33h29.
Abstammungs- Vererbungslehre 87: 708-21. references: Plough and Ives, 1934, DIS 1: 31.
genetics: Deficient for lz and amx. Male lethal. 1934, DIS 2: 10, 34.
*D«l)lz5 Demerec, 1941, Proc. Intern. Congr. Genet., 7th.
cytology: Df(l)8D3-5;8F-9A (Hannah). pp. 99-103.
origin: X ray induced. genetics: Deficient for fa but not pn, w, or ec. Male
discoverer: Green. lethal.
Abstammungs- Vererbungslehre 87: 708—21. cytology: Df(l)3C4-5;3C7-8 (Sutton).
genetics: Deficient for lz and amx. Male lethal. origin: Spontaneous,
*Df(1)lzA discoverer Curry, 38g.
cytology: Df(l)3E;3F + Df(l)9E;9F-10A. genetics: Deficient for fa and spl but not w, rst, or
origin: Associated with In(l)lzA. dm. Male lethal.
DHVm259'4: Deficiency(l) miniature
cytology: Df(l)10C2-3;10E2-3. cytology: Df(l)3C2-3;3E2-3.
discoverer: Demerec, 33i. references: Lefevre and Wilkins, 1966, Genetics 53:
references: Dorn and Bur dick, 1962, Genetics 47: 175-87.
503-18. genetics: Deficient (or N. Carries w^3b, a white
genetics: Male lethal. Heterozygote with m muta- allele causing marbled pigmentation of the eyes.
tions has m phenotype. Heterozygote with dy muta- Df(l)N63b/w resembles WSP/W. Df(l)N63b/
tions is wild type. Recombines with ms9, m, mD, Df(l)w258-4S survives and has lighter eye color
dy61a> dy, and dy5**. than waP/Df(l)w258-45.
*DKm-30: Deficiencyd) Minute-30
origin: Spontaneous. cytology: Df(l)3C6-7;3C7-8.
discoverer: Schultz. origin: X ray induced.
references: 1929, Genetics 14: 366—419. discoverer: Demerec, 33j.
genetics: Deficient for cv and M(1)30. Male lethal. references: Slizynska, 1938, Genetics 23: 291—99.
genetics: Deficient for spl and fa but not w, rst, or
Df(l)mal: Deficiency(l) maroonlike ec. Male lethal.
origin: X ray induced in In(l)scs. *D#l)N264-7
discoverer. E. H. Grell. cytology: Df(l)3C6-7;3C8-9.
references: 1962, Z. Vererbungslehre 93: 371-77. origin: Associated with In(iyii264-7.
genetics: Deficient for sw, mal, and su(f) but not
car or M(l)n. Male lethal. cytology: Df(l)3C6-7;3C10-ll (Demerec and Hoover),
DrTJ)N8; Deficiencyd) Notch origin: X ray induced,
cytology: Df(l)3B4-Cl;3D6-El. discoverer: Demerec, 34a.
origin: Spontaneous, genetics: Deficient for fa, spl, and fan but not w,
discoverer: Mohr, 18j7. rst, dm, or ec. Male lethal.
1932, Proc. Intern. Congr. Genet., 6th. Vol. 1: cytology: Dt(l)3C6-7;3C7-8 (Sutton).
202. origin: X ray induced.
Slizynska, 1938, Genetics 23: 291-99 (fig.), discoverer: Demerec, 34c.
genetics: Deficient for w, rst, fa, spl, and dm. Male genetics: Deficient for fa and spl but not w. Male
lethal. lethal.
*Df(l)N264.19
origin: Spontaneous. cytology: Df(l)3C6-7;3C7-8.
discoverer. Mohr, 28k22. origin: X ray induced,
genetics: Not deficient for w. Male lethal. discoverer: Demerec, 34k.
references: Slizynska, 1938, Genetics 23: 291—99.
cytology: Di(l)3C4-5;3C8-9 (Suttoa). genetics: Deficient for fa but not w or ec. Male
origin: Spontaneous, lethal.
discoverer: Mohr, 28Jc29.
gewetics: Deficient for la and spl but not w, nt, or cytology: Df(l)3A4-S;3C7-9.
dm. Male lethal. origin: X ray induced.
discoverer: Demerec, 36d. genetics: Deficient for fa but not rst or dm. Male
references: Slizynska, 1938, Genetics 23: 291—99 lethal.
(fig-).
genetics: Deficient for w, rst, and fa but not pn, kz, cytology: Df(l)3C4-5;4B4-6 (Hoover).
or dm. Male lethal. origin: X ray induced.
*D«1)N264-31 discoverer: Demerec, 37e.
cytology: Df(l)3B4-Cl;3D2-3, genetics: Deficient for fa, dm, and ec but not w, rst,
origin: X ray induced. or bi. Male lethal.
discoverer: Demerec, 36d. *D#1)N264.46
references: Slizynska, 1938, Genetics 23: 291—99. cytology: Df(l)3C6-7;3C7-8.
genetics: Deficient for w, rst, fa, and dm but not pn origin: X ray induced.
or ec. Male lethal. discoverer: Demerec, 37f.
*D«l)N264-32 genetics: Deficient for fa but not w, rst, or dm. Male
cytology: Df(l)3C3-S;3C7-8. lethal.
origin: X ray induced. *D((J)N264-48
discoverer: Demerec, 36h. cytology: Df(l)lB6-7;lB10-ll.
references: Slizynska, 1938, Genetics 23: 291—99. origin: Associated with In(l)N264-48.
genetics: Deficient for rst and fa but not w or dm. *D%1)N264.49
Male lethal. cytology: Df(l)3C4-5;3E8-Fl (Sutton).
*Df(l)N264.33 origin: X ray induced.
cytology: Df(l)3C6-7;3C7-8. discoverer: Demerec, 37j.
origin: X ray induced, genetics: Deficient for fa, dm, and M(1)3E but not
discoverer: Hoover, 36h. w, rst, ec, or bi. Male lethal.
references: Slizynska, 1938, Genetics 23: 291—99. *DK1)N264-51
genetics: Deficient for fa but not rst or dm. Male cytology. Df(l)3C6-7;3C7-8 (Sutton).
lethal. origin: Found among progeny of radium-treated male.
discoverer: Demerec, 37k.
cytology: Df(l)3A3-4;3D2-3. genetics: Deficient for fa but not w, rst, or dm. Male
origin: X ray induced. lethal.
discoverer. Demerec, 37b. *D«1)N264-S4
references: Slizynska, 1938, Genetics 23: 291—99 cytology: Dt(l)3C3-5;3C7-8 (Hoover).
(fig.). origin: X ray induced.
genetics: Deficient for w, rst, fa, and dm. Male discoverer: Demerec, 38b.
lethal. genetics: Deficient for fa, but not w, rst, or dm.
*W(1)N 264-37 Male lethal.
cytology: Df(l)3C6-7;3C7-8. DHDM264-58
origin: X ray induced. cytology: Df(l)3B2-3;3D6-7.
discoverer: Demerec, 37b. origin: Aneuploid segregant from T(l;3)N264-58/+,
genetics: Deficient for fa but not w, rst, or dm. Male cytology: Df(l)3A10-Bl;3E8-Fl (Demerec).
Di(l )N 2 64-3 8 discoverer Demerec, 38k.
cytology: D%l)2D3-4;3E2-3. genetics: Deficient for w, rst, dm, and M( 1)3E but
origin: X ray induced. not pn, kz, or ec. Male lethal.
discoverer Demerec, 37b. *Df(l)N264.72
references: Slizynska, 1938, Genetics 23: 291—99 cytology: Dt(l)3C6-7;3C7-9 (Sutton).
(fig-). origin: X ray induced.
genetics: Deficient for pn, kz, w, rst, fa, anS dm but discoverer Demerec, 38k.
not br, M(1)3E, or ec. Male lethal. genetics: Deficient for fa but not rst or dm. Male
lethal.
cytology: Df(l)3C6-7;3C7-8 (Slizynska, 1938, *DK7)N2 64-73
Genetics 23: 291-99; Welshons, 1959, Proc. Natl. cytology: D%l)3C3-4;4C6-7 (Demerec).
Acad. Sci. U.S. 44: 254-S8). Recent re-examina- origin: X ray induced.
tion of chromosomes in males from lines marked discoverer: Demerec, 381.
jy264-39 reveals presence of 3C7 (Welshons). genetics: Deficient for fa, rst, dm, M(1)3E, ec, and
origin: Spontaneous. M(l)4BC but not w, bi, or rb. Male lethal.
discoverer: Slizynska, 1937. *D#l}N264-76
genetics: Deficient for /a. Male lethal. cytology. Df(l)3B4-Cl;3E4~5 (Sutton).
cytology: Dt(l)3C6-7;3C8-9 (Sutton). discoverer Demerec, 39b.
origin: Spontaneous. genetics: Deficient for w, rst, fa, dm, and M(1)3E
discoverer: Slizynska, 37®. but not pn or ©c. Male lethal.
genetics: Deficient for spl but not w, rst, or dm.

cytology: Dt(l)3B4-Cl;3C7-8 (Sutton). Male lethal.
discoverer: Demerec, 39b. cytology: Df(l)3C6-7;3D2-3 (Sutton).
geneti cs: Deficient for w, fa, and dm but not pn or origin: X ray induced.
ec. Male lethal. discoverer: Demerec, 40a.
other information: Right break disagrees with inclu- genetics: Deficient for spl and dm but not w, rst, or
sion of dm. Either breakpoint is farther to the right ec. Male lethal.
or dm is mutant instead of missing. *D#l)N264-106
*D1(1)N264-79 cytology: Df(l)3C6-7;3C7-8 (Sutton).
cytology: Di(l)2C10-Dl;3C6-7 (Sutton). origin: X ray induced.
origin: X ray induced. discoverer: Demerec, 40a.
discoverer: Demerec, 39c. genetics: Deficient for spl but not pn, w, rst, or dm.
genetics: Weak Notch phenotype; fa is affected. De- Male lethal.
ficient for kz, pn, w, and rst but not br or dm. Male *D«1)N264.} 08
lethal. cytology: Df(l)3C3-5;3E7-S.
origin: Associated with In(iysi264-10S.
cytology. Df(l)3C6-7;3C7-8 (Sutton).
origin: X ray induced, cytology: Df(l)3B4-Cl;3D2-3 (Sutton).
genetics: Deficient for fa but not rst or dm. Male discoverer: Demerec, 40a.
lethal. genetics: Deficient for w, rst, spl, and dm but not
pn or ec. Male lethal.
cytology: Df(l)3C7-8;3E5-6. *Df(l)N264.111
origin: Aneuploid segregant from T(l;4)N264-86/+. cytology: Df(l)3C3-5;3C12-Dl (Sutton).
Kl)N26489 origin: X ray induced.
cytology: Df(l)3B2-3;3F2-3 (Sutton). discoverer: Demerec, 40b.
origin: X ray induced. genetics: Deficient for spl and rst but not pn, w, dm,
discoverer Demerec, 39j. or ec. Male lethal.
genetics: Deficient for w, rst, fa, M(1)3E, and ec *D«J)N264-114
but not pn. Male lethal. cytology: Df(l)3C6-7;3D4-5 (Sutton).
cytology: Dt(l)3C7-8;3E8-Fl (Sutton). discoverer: Kaufmann, 40d.
origin: X ray induced. genetics: Deficient for rst, spl, and dm but not w or
discoverer: Demerec, 39j. ec. Male lethal.
genetics: Deficient for spl, dm, M(1)3E, and ec but * D«l)N264.n 5
not pn or w. Male lethal. cytology: Df(l)3C3-5;3E2-3 (Sutton).
cytology: Dt(l)3B4-Cl;3F3-4 (Sutton). discoverer Sutton, 40e.
origin: X ray induced. genetics: Deficient for rst, spl, and din but not w,
discoverer: Demerec, 39k. M(1)3E, or ec. Male lethal.
genetics: Deficient for w, spl, dm, M(1)3E, and ec
but not pn or bi. Male lethal. cytology: Dt(l)3A6-7;3E2-3 (Sutton).
cytology: Df(iy3C6-7;3C7-8 (Sutton). discoverer: Demerec, 40g.
origin: X ray induced. genet fcs: Deficient for w, rs t, spl, and dm but not
discoverer Demerec, 39k. pn, ec, or bi. Male lethal.
genetics: Deficient for spl but not w, rst, dm, or ec.
Male lethal. cytology: Df(l)3C6-7;3C7-9 (Sutton).
cytology; Df(l)2D2-3;3Cll-12 (Sutton). discoverer Demerec, 40h.
origin: X ray induced. genetics: Deficient for spl but not pn, w, rst, or dm.
discoverer Demerec, 40«. Male lethal.
genetics: Deficient for pn, kz, w, rst, spl, and dm *D«1)N264 -720
but not ec or bi. Male lethal. cytology: Df(l)3C6-7;3D2-3 (Sutton).
cytology: Dt(l)3B4-Cl;4B4-5. discoverer: Demerec, 40j.
origin: Aaeuploid s«gregant from T(l;3yst*64 genetics: Deficient for spl and dm but not kz, w,
rst, or ec. Male lethal.
cytology: D§(l)3C4'S;3C7'g (Sutton),
origin: X ray induced. cytology: Dt(l)3C4-5;3C7-8 (Sutton).
discoverer: Demerec, 40a. origin: X ray induced.
CHROMOSOME ABERRATIONS _ DEFICIENCIES 297
discoverer: Demerec, 41a. discoverer: Muller, Valencia, and Valencia, 1946-53.

genetics: Deficient for spl but not kz, w, rst, dm, or references: Valencia, 1966, DIS 41: 58.
ec. Male lethal. genetics: Deficient for ras and v.
Df(l)rbKlSBH3: DeficiencyO) ruby
cytology: Df(l)3C3-5;3D4-5 (Sutton). cytology: Df(l)4B4-5;4D5-6.
origin: Spontaneous. origin: X ray induced in R(l)2.
discoverer: Bishop, 401. discoverer: Muller, Valencia, and Valencia, 1946-53.
genetics: Deficient for rst, spl and dm but not w. references: Valencia, 1966, DIS 41: 58.
Male lethal. genetics: Deficient for rb.
*Df(l)N264-127 Df(l)rst2: Deficiency(l) roughest
cytology: D£(l)3C6-7;3C7-8 (Sutton). cytology: Df(l)3C3-4;3C6-7 (Schultz; Sutton).
discoverer: Demerec, 41b. discoverer: Bridges, 33d7.
genetics: Deficient for spl but not kz, w, rst, or dm. references: Gersh, i965, Genetics 51: 477—80 (fig.)-
Male lethal. genetics: Deficient for rst and vt. Homozygous
viable.
cytology: Df(l)3C6-7;3C7-8 (Sutton). Df(l)sc4Lsc8R
origin: X ray induced. cytology: Df(l)19F-20Cl;20B-Dl +Dp(l;l)lB2-
discoverer: Demerec, 41b. 3.1B3-4.
genetics: Deficient for spl but not w, rst, or dm. origin: Associated with In(l)sc4Lsc8R.
Male lethal. Df(1)sc4LscL8R
*Df(l)N264.130 cytology: Df(l)19F-20Cl;20B-C.
cytology: Df(l)3C6-7;3C7-8 (Sutton). origin: Associated with In(l)sc4LscL8R.
origin: Spontaneous. DrT7)sc8; DeficiencyfJ) scute
discoverer: Neel, 41c. origin: Spontaneous in In(l)sca.
references: 1942, Genetics 27: 530. discoverer. Noujdin.
genetics: Deficient for fa but not w, rst, or dm. Male references: 1935, Zool. 2h. 14: 317-52.
lethal. genetics: Deficient for y, ac, andHw. Male lethal;
*D«1)NB: Deficiency^) Notch of Bernstein dies as late embryo; larva nearly complete
cytology: Di(l)3C4-5;3C12-Dl (Sutton). (Poulson, 1940, J. Exptl. Zool. 83: 271-325).
origin: Spontaneous. *Df(J)sc*25b
discoverer: Bernstein, 28a7. cytology: Like In(l)scs in mitotic prophase, but
genetics: Deficient for fa but not w, rst, or dm. Male without heterochromatic segments hB and most of
lethal. hA, are ordinarily carried distally in In(l)sc8,
origin: Spontaneous derivative oiln(l)sc8; Y chro-
cytology: Df(l)3C6-7;3C7-8 (Sutton). mosome not demonstrably involved.
origin: Spontaneous, discoverer: Lindsley, 1952.
discoverer: Morgan, 1929. synonym: sc*c.o. X 25b.
genetics: Deficient for fa but not w. Male lethal. references: 1958, Z. Vererbungslehre: 89: 103—22.
D#1)04: Deficiency(J) of Oliver genetics: Deficient for y and ac; mutant for bb. Male
origin: Aneuploid segregant from T(l;3)O4/+. lethal with normal Y; viable with y+Y.
DfCi)pnnoAc4' Deficiencyd) prune
cytology: Df(l)2C8-9;3Al-2 superimposed on cytology: Like In(l)sc8 in mitotic prophase.
In(l)lB3-4;20B-DlLIB2-3;20B-DlR + In(l)4D7- origin: Spontaneous product of recombination be-
Bl;llF2-4. tween the distal heterochromatin of In(l)sca and
new order: 1A - 1B3|2OB - 11F4|4E1 - 11F2|4D7 -
3A2|2C8 - 1B3|2OD1 - 20F. discoverer: Lindsley, 1952.
origin: X ray induced in In(l)scstLsc8R+dl-49. synonym: sc8c.o. X 89a.
discoverer: Muller, Valencia, and Valencia, 1946-53. references: 1955, Genetics 40: 24—44.
references: Valencia, 1966, DB 41: 58. genetics: Deficient for y and ac but not bb. Carries
genetics: Deficient for pn. KS, the fertility complex of Ys distaliy. Male
Df(l)pn-ec: Defic'tency(1) prune to echinus lethal with normal Y, viable with y*Y.
cytology: Df(l)2Dl-2,-3F7-4A4. other Information: Four similar deficiencies,
origin: Aneuploid segregant from T(l;A)pn-ec/+. D%iysc*Elt D%l)mc8P0t Dfi>c*916, and
D*(J>ras-W7Cc8; Deficiency(l) raspberry to DSCl)»cH7ht also described by Ltadtley (1955).
vermilion *D«1)sc*99c
cytology: Df(l)9E3-4;10A4-5 superimposed on cytology: Like In(l)mc8 in mitotic prophase.
ln(l)lB3-4;20B-DlLlB2-3;20B-DlR 4- In(l)4D7- origin: Spontaneous derivative of In(l)mc8; Y chro-
El;llF2-4. mosome not demonstrably involved,
new order 1A - 1B3J20B - UF4J4E1 - 9E3|l0A5 - discoverer Lindsley, 1952.
11F2|4D7 - 1B3J2OD1 - 20F. synonym: ®c9c,o. X 99c.
origin: X ray induced in ln(l)scSiLmc8R+dI'49. references: 1958, Z. Veitrrbttafslefar®: 89t 103-22.
genetics: Deficient for y and ac but not for bb. Male

lethal with normal Y, viable with y*Y. origin: Spontaneous in In(l)sc8.
discoverer: Mather, 1937.
cytology: Like In(l)sc8LBNR in mitotic prophase genetics: Deficient for y, ac, and bb. Male lethal.
but with the part of heterochromatic segment hA
ordinarily carried distally and a portion of hB cytology: Like In(l)scSL,ENR in mitotic prophase.
missing. origin: Spontaneous product of recombination be-
origin: Spontaneous derivative of In(l)scSLENR; Y tween the distal heterochromatin of In(l)sc8LENR
chromosome not demonstrably involved. Postulated and Ys.
to be a product of exchange between the distal discoverer: Lindsley, 1950.
heterochromatin of one chromatid and the proximal synonym: sc8ENc.o. X P7.
heterochromatin of its sister. references: 1955, Genetics, 40: 24—44.
discoverer. Lindsley, 1950. genetics: Deficient distally for y and ac but not bb.
synonym: sc8ENc.o. X Bl. Carries KS, the fertility complex of Ys distally.
references: 1958, Z. Vererbungslehre: 89: 103—22. Male viable.
genetics: Deficient dis tally for y and ac. Male other information: Three similar deficiencies,
viable. Df(l)sc8L7, Df(l)sc8P0, and Dt(l)sc835a, also
other information: Two similar deficiencies, described .by Lindsley (1955).
Df(l)sc*C6 and Df(l)sc8D8, described (Lindsley, *D«l)sc*Ql
1958). cytology: Like In(l)sc8LENR in mitotic prophase,
but portion of heterochromatic segment hA ordi-
cytology: Like In(l)sc8LiENR in mitotic prophase. narily carried distally missing.
origin: Spontaneous derivative of In(l)sc8LENR; Y origin: Spontaneous derivative of In(l)sc8LENR; Y
chromosome not demonstrably involved. Postulated chromosome not demonstrably involved. Postulated
to be a product of exchange between the distal to result from exchange between distal and prox-
heterochromatin of one chromatid and the proximal imal heterochromatin of sister chromatids.
heterochromatin of its sister. discoverer: Lindsley, 1950.
discoverer: Lindsley, 1950. synonym: sc8ENc.o. X Ql.
synonym: sc8ENc.o. X C4. references: 1958, Z. Vererbungslehre: 89: 103—22.
references: 1958, Z. Vererbungslehre: 89: 103-22. genetics: Deficient distally for y and ac. Male
genetics: Deficient distally for y and ac. Male viable.
viable.
other information: Five similar deficiencies, cytology: Like In(l)sc8LENR in mitotic prophase
Dt(l)sc*C13, Df(l)sc8D6, Df(l)sc*F7, Df(l)sc*O7, but carrying only heterochromatic segments hC and
and Df(l)ac853c also described (Lindsley, 1958). hD distally.
origin: Spontaneous derivative of In(l)sc8LENR; Y
*Df(l)sc*J3 chromosome not demonstrably involved. Postulated
cytology*. Like In(l)sc8LENR in mitotic prophase, result of exchange between distal and proximal
but with part of heterochromatic segment hA, ordi- heterochromatin of sister chromatids.
narily carried distally and portion of hB missing. discoverer: Lindsley, 1950.
origin: Spontaneous derivative of In(l)ac8LENR; Y synonym: sc8ENc.o. X S7.
chromosome not demonstrably involved. Postulated references: 1958, Z. Vererbungslehre: 89: 103—22.
to result from exchange between distal and prox- genetics: Deficient distally for y, ac, and bb. Male
imal heterochromatin of sister chromatids. viable,
discoverer. Lindsley, 1950. other information: A similar deficiency, Df(l)sc8W0,
synonym: ac8ENc.o. X J3. also described (Lindsley; 1958).
references: 1958, Z. Vererbungslehre: 89: 103-22.
genetics: Deficient distally for y and ac; mutant for cytology: Df(l)lB2-3;lB3-4.
66. Male viable. origin: Associated with In(l)sc8Lsc4R.
)
cytology: Like In(l)sc8LENR in mitotic prophase cytology: Df(l)lB2-3;lB3'4.
but carrying only heterochromatic segments hC and origin: Associated with In(l)sc8LscL8R.
hD distally. D«J)sc*L.scS1R
origin: Spontaneous derivative of In(l)sc8LENR; Y cytology: Df(l)lB2-3;lB3-4.
chromosome not demonstrably involved. Postulated origin: Associated with In(l}sc8LacslR.
to result from exchange between distal and prox-
imal heterochromatin of sister chromatids. cytology. Dt(l)lBl-2;lB2-3;lB14-Cl.
discoverer: Lindsley, 1950. new order 1A - 1B1J1B14 - lB3|lCl - 20;
synonym: ac*ENc.o. X Kl. 1B2 missing.
references: 1958, Z. Vererbungslehre: 89: 103-22. origin: X-ray-induced derivative of ln(l}ac3
genetics: Deficient distally for y and ac; mutant for In(l)lB2-3;lB14-Cl.
bb. Male viable. discoverer: Sturtevant, 1930.
references: 1935, DIS 3: 15. *Dfil)t282-l: Deficiencyil) tan

1936, Genetics 21: 444-66. cytology: Df(l)8C2-3;8C14-Dl (Sutton). Green and
genetics: Mutant for sc; viability low. Green (1956, Z. Induktive Abstammungs-
Vererbungslehre 87: 708-21) suggested that the
origin: X ray induced. deficiency probably extends farther to the right.
discoverer: Muller. origin: X ray induced.
references: Patterson and Muller, 1930, Genetics 15: discoverer: Demerec, 34c.
495-577. genetics: Deficiency for t, lz, and amx but not dd,
Dubinin, 1933, J. Genet. 27: 443-64. dvr, tip, ny, or ras. Male lethal.
genetics: Mutant for sc; deficient for y and ac. Ap-
parently, y + and ac* loci were inserted into an *Df(1)vB: Deficiencyil) vermilion of Bridges
autosome and subsequently lost. Originally tested origin: Spontaneous.
as an allele of sc only. Male lethal. discoverer: Bridges, 16e9.
references: 1919, J. Gen. PhysioL 1: 645-56.
cytology: Df(l)lBl-2;lB4-7. genetics: Deficient for v. Male lethal. Genetic map
origin: Aneuploid segregant from T(J;2)sci 9/+. shortened 1—3 units.
cytology. Di(l)lB2-3; terminal deficiency. Df(l)w258.3: Deficiencyil) white

origin: Aneuploid recombinant from cytology: Df(l)3B2-3;3Cl-2 (Sutton).
D1(l)scFah: Deficiencyil) scute of Fahmy discoverer: Demerec, 33h.
cytology: D((l)lA8-Bl;lB2-3. genetics: Claimed to have pn affected, but cytology
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- not in agreement with deficiency for pn; therefore
phenylalanine (CB. 3007). probably an independent mutant, w affected, prob-
discoverer Fahmy, 1954. ably mutant, br and fa not affected. Male lethal.
genetics: Probably mutant for sc. Male viable; cytology: Df(l)3A2-3;3C3-5.
homozygous female lethal. origin: X ray induced.
discoverer: Demerec, 33j.
J': Deficiencyil) scute of Jacobs-Duller references: Slizynska, 1938, Genetics 23: 291—99.
cytology: Df(l)lB;3A3-C2. genetics: Deficient for w but not pn, kz, rst, fa, or
origin: Associated with T(l;3)scJ4. ec. Male lethal.
Df(l)scl-8LSC8R *Df(l)w2S8.U
cytology: Df(l)20B-C;20B-Dl. cytology: Df(l)3A3-4;3Cl-2.
origin: Associated with In(l)scL8Lsc8R. discoverer: Demerec, 33k.
cytology: Df(l)20B-C;20B-Dl. genetics: Mutant or deficient for w but not pn, kz,
origin: Associated with In(l)scL8LscslR. rst, or fa. Male lethal.
Df(l)w2S8-42
V1: Deficiencyil) scute of Valencia cytology: Df(l)3A6-8;3C3-S (Sutton).
cytology: Df(l)lA8-C3; terminal deficiency, origin: X ray induced.
origin: Aneuploid recombinant from In(lLR)scvl/+. discoverer. Demerec, 38i.
Df(l)sn: Deficiency(1) singed genetics: Deficient for w but not pn, kz, or rst.
cytology: Dl(l)7B2-3;7D22-El. Male lethal.
origin: Spontaneous in R(l)2. DfU)w2S8-4S
discoverer: C. Hinton. cytology: Dt(l)3B4-C2;3Cl-4; is Df(l)3B4-Cl;3Cl-2
references: Hinton and Welshons, 1955, DIS 29: according to Sutton and Df(l)3Cl-2;3C3-4 according
125-26. to Schultz. May lack 3C2 (Lefevre and Wilkins,
genetics: Deficient for sn but not ct, oc, or ptg. 1964, Genetics 50: 264).
Male lethal. origin: X ray induced.
Df(l)sta: Deficiencyil) stubarista discoverer Demerec, 381.
cytology: Df(l)lD3-El;2A. genetics: Mutant or deficient for w but not pn, kz,
origin: Aneuploid segregant from T(l;3)sta/+. rst, or fa. Male lethal.
Df(l)svr: Deficiencyil) silver Df(1)w2S8.48
cytology: Df(l)lB10-13; apparently a terminal cytology: Df(l)3A9-Bl;3Cl-2 (Sutton).
deficiency. origin: X ray induced,
origin: Found among progeny of cold-treated female. discoverer Demerec, 39c.
discoverer: L. V. Morgan. genetics: Mutant or deficient for w but not pn, kz,
references: 1940, DIS 13: 51. rst, or fa. Male lethal.
Suttoa, 1943, Genetics 28: 213.
genetics: Deficient for y, ac, sc, and swr but not cytology: Dt(l)3Cl-2;3C3-4 + Dt(l)20A;2QB.
mjfs) or etm. Male lethal. origin: Associated with ln(l)wm4Lrst3Ji.
l(l)zw's; mutant or deficient for w. Male lethal.

cytology: Df(l)3Cl~2;3C2-3 +Df(l)20. Survives as w female when heterozygous with the
origin: Associated with In(l)wm4^wmJR. deficiency for 3C2-3, Df(l)wm*Lrst3R =Df(l)3Cl-
DfOV m 4 9 °: Deficiency(i) white-mottled 2;3C3-4.
cytology: Di(l)3A10-Bl;3E2-3.
origin: Aneuploid segregant from T(l;3)wm49a/+. cytology: Df(l)3Cl-3;3C12-D3.
origin: Product of unequal exchange between the
cytology: Df(l)3B2-Cl;3C9-Dl. 3C1-3 region of a chromosome carrying wa and re-
origin: Aneuploid segregant from T(l;2)wmS3a/+, gion 3C12-D3 of a specific homolog.
*Df(1)wn> 258-44 discoverer: Judd.
cytology: Df(l)3C3-4;4D2-El. synonym: w~r N.
origin: Aneuploid segregant from references: 1961, Genetics 46: 1687-97 (fig.)-
T(l;2;3)w°>258-44/+. 1964, DIS 39: 59.
*D1(1hmDV4: Deficiency^) white-mottled of genetics: Deficient for N; mutant or deficient for w
Dubinin and Volotov and dm. Male lethal. Survives as a w female when
cytology: Df(l)3C3-7;3D, heterozygous with Di(l)w258-45 =Dt(l)3B4-
origin: Associated with T(1;4)W«DV4. C2;3Cl-4, Df(l)WrJl = Df(l)3A2-3;3Cl-3, and
Dt(l)wrJ2 =Df(l)3A6-8;3Cl-3.
cytology: Df(l)3C2-3;3C3-4. Df(l)wvco: Deficiency(l) white-variegated
origin: Associated with In(l)wmJLrst3R. cobbled
*Df(l)wrS: Deficiency(l) white-recombinant of cytology: Dt(l)2B17-Cl;3C4-5.
Green origin: Aneuploid segregant from T(l;3)wvco/+.
cytology: Dt(l)3A3-4;3Cl-2 (E. B. Lewis). Df(l)w-ec: Deficiency(l) white-echinus
origin: A regular product of unequal exchange be- cytology: Df(l)3Cl-2;3E7~8.
tween 3C1-2 of a wa or w*2 chromosome and 3A3-4 origin: Aneuploid segregant from T(l;2)w-ec/+.
of certain specific homologous chromosomes. Re- synonym: Df(w-ec)64d.
ciprocal of Dp(l;l)wrG.
discoverer Green. cytology: Df(l)lA;lB2-3.
references: 1959, Genetics 44: 1243-56. origin: Associated with In(l)y3PLscSR.
genetics: Mutant or deficient for w but not z. Male
lethal. Survives as w female when heterozygous cytology: Df(l)lA8-Bl;lB3-4 + Df(l)18A3-4;19F-
with D£(l)w^4Lrst3R =D%l)3Cl-2;3C3-4. 20C1.
Di(J)wrJ1: DeficiencyiV white-recombinant of origin: Associated with In(l)y4Lsc4R.
Judd
cytology: Df(l)3A2-3;3Cl'3. cytology: Di(l)lA8-Bl;lB2-3 + Df(l)18A3-4;20B-Dl.
origin: A regular product of unequal exchange be- origin: Associated with In(l)y4Lsc8R.
tween the 3C1-3 region of a chromosome carrying D«l)y4Lsc9R
w« and the 3A2-3 region of certain specific homol- cytology. Df(l)lA8-Bl;lB2-3 + Dt(l)18A3-4;18B8-9.
ogous chromosomes. origin: Associated with In(l)y4Lsc9R.
discoverer: Judd. *Df(l)zJ: DeficienCy(l) zeste
synonym: w~*. cytology: Di(l)2C2-3;3E2-3.
references: 1961, Genetics 46: 1687-97 (fig.). origin: X ray induced.
1964, DIS 39: 60. discoverer: Gans.
genetics: Deficient for z and the lethals between z references: 1953, Bull. Biol. France Belg. Suppl.
and w; mutant or deficient for w. Male lethal. Sur- 38: 1-90 (fig.),
vives as a if female when heterozygous for the de- genetics: Deficient for pn, z, and w. Male lethal.
ficiency for 3C2-3, Dt(l)w«>4Z<mt3R = Df(l)3Cl-
2;3C3-4. cytology: Df(l)2D4-5;3C3-4.
cytology: Df(l)3A6-8;3Cl-3 [could be the same as discoverer: Gans.
references: 1953, Bull. Biol. France Belg. Suppl.
origin: A regular product of unequal exchange be- 38: 1-90 (fig.),
tween the 3C1-3 region of a chromosome carrying genetics: Deficient for pn, z, and w. Male lethal.
w* or w*dP with the 3A4-8 region of specific homol- *D«l)x3
ogous chromosomes. Probably reciprocal recombi- cytology: Df(l)2CS-6;3B2-3.
nant of Dp(l;l)w*J2. origin: X ray induced.
discoverer: Judd. discoverer: Gans.
•yrtonyutt *r-*. reference*: 1953, Bull. Biol. France Belg. Suppl.
references: 1961, Genetics 46: 1687-97 (fig.)- 38: 1-90 (fig.),
1964, DB 39: 59. genetics: Deficient for pn and z. Male lethal.
genetics: Carries normal alleles of z, l(l)zwl, *Df(1)z4
Jfijcwtf, and t(l)zw4; deficient for the other cytology: D%1)2C5-6;3A9-B1,

discoverer: Gans. Inst. Wash. Publ. No. 278: 277.
references: 1953, Bull. Biol. de France Belg. Suppl. genetics: Homozygous lethal. Gives decreased
38: 1-90 (fig.), crossing over in d-b region.
genetics: Deficient for pn and z. Male lethal. D«2L)G
*D«1)z5 cytology: Dt(2L)36B5-6;40F.
cytology: Df(l)3Al-3;3A4-6. origin: Aneuploid segregant from T(Y;2)G/+.
origin: X ray induced. *Df(2L)H
discoverer: Gans. cytology: Df(2L)37B2-3;40B2-3.
references: 1953, Bull. Biol. France Belg. Suppl. origin: Aneuploid segregant from T(Y;2)H/+.
38: 1-90 (fig.). Df(2L)M-B: see Bi(2L)U-zB
genetics: Deficient for z. Male lethal. Df(2L)M-C: see Df(2L)M-zC
*D«l)z6 D«2L)M-zB: Deficiency(2L) Minute-z-B
cytology: Di(l)3A6-8;3C10-ll. cytology: Df(2L)24E2-Fl;25Al-2.
origin: X ray induced in z. origin: Spontaneous.
discoverer: Gans. discoverer: Bridges, 38dl2.
references: 1953, Bull. Biol. France Belg. Suppl. synonym: Df(2L)M-B.
38: 1-90 (fig.). references: Morgan, Schultz, Bridges, and Curry,
genetics: Deficient for w but not z. Male lethal. 1939, Carnegie Inst. Wash. Year Book 38: 276—
77.
*D«Y)bb: Deficiency(Y) bobbed Curry, 1939, DIS 12: 46.
cytology: Metaphase chromosomes show short arm of 1941, DIS 14: 50.
Y reduced to one-third normal size. genetics: Deficient for M(2)z, dp, and dw-24F but
origin: Spontaneous, not ed, ft, G, M(2)S1, l(2)cg, or tkv. Homozygous
discoverer: Schultz, 33k8. lethal.
genetics: Deficient for bb but not KL or KS.
cytology: Df(2L)24D2-5;25A2-3.
Dt(2)M-33a: see Df(2R)M-c33a origin: Spontaneous.
D«2L)64j discoverer: Curry, 37g27.
cytology: Dt(2L)34E5-Fl;35C3-Dl. synonym: Df(2L)M-C.
origin: X ray induced. references: Morgan, Bridges, and Schultz, 1938,
discoverer: E. H. Grell, 1964. Carnegie Inst. Wash. Year Book 37: 307.
genetics: Deficient for rk, b, j, el, Sco, Su(H), pu, Morgan, Schultz, Bridges, and Curry, 1939,
and Adh but not for nub, M(2)e, rd, or M(2)m. Homo- Carnegie Inst. Wash. Year Book 38: 276—77.
zygous lethal. genetics: Deficient for ed, ft, G, M(2)z, dp, and
Df(2L)al: Deficiency(2L) aristaless dw-24F but not M(2)S1, l(2)cg, or tkv. Homozygous
cytology: Df(2L)21B8-Cl;21C8-Dl. lethal.
origin: X ray induced. DK2L)SS6f: Deficiency(2L) Star
discoverer: E. B. Lewis, 1940. cytology: D%2L)21C6-D1;22A3-B1.
references: 1945, Genetics 30: 147—51, origin: Associated with In(2LR)S56f.
genetics: Deficient for al, ex and ds but not for *DK2L)S-dmr: Deficiency(2L) Star derived
l(2)gl, net, or S. Homozygous lethal. cytology: Di(2L)21D4-El;21E2~3.
*DK2L)bt>: Deficiency(2L) black-Dominant new order: YJ21D4 - 21 A;
cytology: Df(2L)3SC;35D (Kodani). 60 - 2 1 E 3 | l 0 1 .
origin: Spontaneous. origin: Synthetic; a combination of 2LDYP from
discoverer: Goldschmidt, 1945. T(Y;2)21E = T(Y;2)21D4-E1 and 4^2P from
references: 1945, Univ. Calif. (Berkeley) Publ. T(2;4)astv - T(2;4)21E2-3;101.
Zool. 49: 504, 520. discoverer: E. B. Lewis,
genetics: Possibly deficient for 6. Homozygous references: 1945, Genetics 30: 137—66.
lethal. genetics: Deficient for S and ast. Homozygous
Df(2L)C263: Deficlency(2L) Crossover lethal.
suppressor *D«2L)S1
cytology: DB(2L)25F;26F. cy to I ogy: Df(2L)21 C3~4;22A2-3.
origin: Associated with ln(2L)C263. origin: X ray induced,
discoverer E. B. Lewis, 1940,
cytology: Dt(2L)22Dl-2;22D3-El + Dt(2L)33FS- references: 1945, Genetics 30: 147—51.
34Al;34A8-9. genetics: Deficient for ds, S, and ast but not 1(2)0,
origin: Associated with Jn(2L)CyLtR. net, al, ex, shr, or ho. Homozygous lethal.
*EM(2L)d: Defici«ncy(2L) dachs DK2L)S2
origin; Spcntaaeoos in d stock. cyfology: Df(2L)21 C6-D1;22A6-B1.
dis«©'vee®« Bridges, 15j6. origin: X my induced,
synonym: dh discoverer: E. B. Lewis, 1940.
references: 1945, Genetics 30: 147-51. *Df(2R)bwR4<>: Deficiency(2R) brown-Rearranged

genetics: Deficient for ds, S, ast, and shr but not cytology: Dt(2R)59C5-6;59E2-3.
l(2)gl, net, al, ex, or ho. Homozygous lethal. origin: X ray induced.
D«2L)S3 discoverer: Slatis.
cytology: Df(2L)21D2-3;21 F2-22A1. references: 1955, Genetics 40: 5-23.
origin: X ray induced. genetics: Associated with bwR40,
discoverer: E. B. Lewis, 1940.
references: 1945, Genetics 30: 147—51. cytology: Df(2R)41;42A2-3.
genetics: Deficient for S and ast but not l(2)gl, net, origin: Aneuploid recombinant from
al, ex, ds, shr, or ho. Homozygous lethal. In(2R)bwV 34k+cy/+,
*D«2L)S4
cytology: Df(2L)21C3-4;22B2-3. cytology: Df(2R)41B2-Cl;42A2-3.
origin: X ray induced, origin: Associated with In(2R)bwVDe>1LCyR.
discoverer: E. B. Lewis, 1940.
references: 1945, Genetics 30: 147—51. cytology: Di(2R)41A-B;42A2-3.
genetics: Deficient for ds, S, ast, and shr but not origin: Associated with In(2R)bwVDe2LCyR.
l(2)gl, net, al, ex, or ho. Homozygous lethal.
*D«2L)S5 cytology: Di(2R)58A4-Bl;59E2-4.
cytology: Df(2L)21C2-3;22A3-4. origin: Associated with In(2R)CyLbwvêlR.
discoverer: E. B. Lewis, 1940. cytology: Df(2R)58A4-Bl;59D6-El.
references: 1945, Genetics 30: 147—51. origin: Associated with In(2R)CyLbwVDe2R.
genetics: Deficient for ex, ds, S, and ast but not
1(2)0, net, al, shr, or ho. Homozygous lethal. cytology: Df(2R)58A4-Bl;59.
*DK2L)S7 origin: Aneuploid recombinant from
cytol ogy: Df(2L)21 C3-4;21 F2-22A1.
origin: X ray induced in net ho. DfT2R)M-c"a: Deficiency^/?) Minute-c
discoverer: E. B. Lewis, 1940. cytology: Df(2R)60E2-3;60Ell-12.
genetics: Deficient for ds, S, and ast but not l(2)gl, discoverer Schultz, 33a7.
al, ex, or shr. Homozygous lethal. synonym: Df(2)M-33a.
D«2L)Sw-L: Deficiency(2L) Swedish-L references: Bridges, 1937, Cytologia (Tokyo), Fujii
cytology: Tip of 2L contains deficiency, Jub. Vol. 2: 745-55.
origin: Naturally occurring condition in some genetics: Deficient for M(2)c. Homozygous lethal.
Swedish strains,
discoverer: Gustafson, 1937. cytology: Dt(2R)57Fll-S8Al;58F8-59Al.
genetics: No phenotypic effect. origin: Spontaneous.
D«2R)42 references: Morgan, Bridges, and Sturtevant, 1925,
cytology: Dt(2R)42C3-8;42D2-3. Bibliog. Genet. 2: 231.
origin: Probably X ray induced. Found on chromo- Bridges, 1937, Cytologia (Tokyo), Fujii Jub. Vol.
some with vgs. 2: 745-55.
discoverer: Bridges, 36b. genetics: Deficient for px, l(2)Su(H), M(2)l, and
references: Morgan, Bridges, and Schultz, 1938, probably a. Homozygous lethal.
Carnegie Inst. Wash. Year Book 37: 304-9. DK2R)M-S2: Deficiency(2R) Minute of Schultz 2
genetics: Deficient for no tested loci. Homozygous cytology: Salivary chromosomes apparently normal.
lethal. Located in chromocentric region of 41A.
*Df(2R)a*"»2: DeiiciencyUR) arc-broad angular origin: X ray induced.
cytology: Dt(2R)S8D5-6;5SD7-8. discoverer: Schultz, 33al2.
origin: Spontaneous. references: Morgan, Bridges, and Schultz, 1938,
discoverer: Goldschmidt. Carnegie Inst. Wash. Year Book 37: 306.
references: 1945, Univ. Calif. (Berkeley) Publ. Morgan, Schultz, Bridges, and Curry, 1939,
Zool. 49: 363-73, 388-89. Carnegie Inst. Wash. Year Book 38: 273—77.
genetics: Associated with a 6 * 2 . Morgan, Schultz, and Curry, 1940, Carnegie Inst.
D«2R)bwS: Deficiency(2R) brown Wash. Year Book 39: 251-55.
cytology: Dt(2R)59D10-El;59E4-Fl. genetics: Lethal homozygous; mutant phenotype
origin: Spontaneous. with stw but not Jag, It, ti, ap, tk, std, or mat;
discoverer. Motor, 31k28. judged a deficiency on this basis.
genetics: Deficient for bw. Homozygous lethal.
cytology. Salivary chromosomes apparently normal,
cytology: D%ZR)41;42A2-3 origin: X ray induced,
ofigin: Associated with discoverer Schultz, 33a5.
synonym: Df(2R)M-S4. discoverer: Nichols-Skoog, 33b20.

references: Morgan, Bridges, and Schultz, 1938, references: Bridges, Skoog, and Li, 1936, Genetics
Carnegie Inst. Wash. Year Book 37: 306. 21: 788-95 (fig.).
Morgan, Schultz, Bridges, and Curry, 1939, Li, 1936, Peking Nat. Hist. Bull. 11: 39-48.
Carnegie Inst. Wash. Year Book 38: 273—77. genetics: Not deficient for en, Wo, or en.
Morgan, Schultz, and Curry, 1940, Carnegie Inst. Df(2R)Np/T(2;3)dp lethal and homozygous lethal.
Wash. Year Book 39: 251-55. Df(2R)0re-R: Deficiency(2R) Oregon-R
genetics: Gives mutant phenotype with stw and ap. cytology: Df(2R)60F2-3; terminal deficiency.
Lethal homozygous and with M(2)S2, l(2)Sp9c, origin: Naturally occurring in Oregon-R stock,
l(2)Spll, and l(2)SpJ5. No interaction with Jag, It, discoverer: Bridges, 3615.
rl, tk, std, or msf. Judged a deficiency on basis of genetics: No detectable phenotypic effect in homo-
genetic evidence. zygote.
DH2R)P
cytology: Salivary chromosomes appear normal. cytology: Dt(2R)58E3-Fl;60D14-E2.
origin: X ray induced. origin: Aneuploid segregant from T(2;3)P/+.
discoverer: Schultz, 33a3. D«2R)Px: Deficiency^/?) Plexate
synonym: Df(2R)M-S8. cytology: Df(2R)60B8-10;60Dl-2.
references: Morgan, Bridges, and Schultz, 1938, origin: Spontaneous.
Carnegie Inst. Wash. Year Book 37: 306. discoverer: Bridges, 22f6.
Morgan, Schultz, Bridges, and Curry, 1939, references: 1937, Cytologia (Tokyo), Fujii Jub. Vol.
Carnegie Inst. Wash. Year Book 38: 273—77. 2: 745-55.
genetics: Lethal homozygous and with M(2)S2, genetics: Deficient for 1(2)NS, sp, bs, 6a, and Pin
l(2)Sp9c, l(2)Spll, and l(2)Spl5. Gives mutant but not or, Fo, pd, 11, mr, or l(2)ax. Homozygous
phenotype with stw but not r/ or ap. lethal.
Df(2R)Px2
cytology: Df(2R)41A. In mitotic metaphase 2R about cytology: Df(2R)60C5-6;60D9-10.
three-fourths normal size, origin: X ray induced.
origin: X ray induced, discoverer. Schultz, 3211.
discoverer. Schultz, 32k22. references: Bridges, 1937, Cytologia (Tokyo), Fujii
synonym: Dt(2R)M-S10. Jub. Vol. 2: 745-55.
references: Morgan, Schultz, Bridges, and Curry, genetics: Deficient for 6s, ba, and Pin but not
1939, Carnegie Inst. Wash. Year Book 38: 2 7 3 - 1(2)NS, sp, or M(2)c. Homozygous lethal.
77. Df(2R)Px4
Morgan, Schultz, and Curry, 1940, Carnegie Inst. cytology: Dt(2R)60B;60Dl-2.
Wash. Year Book, 39: 251-55. origin: Associated with In(2LR)Px4.
k
1941, Carnegie Inst. Wash. Year Book 40: 282—87. Df(2R)PxS
genetics: Lethal homozygous and with M(2)S2, origin: Spontaneous in In(2LR)bwvl.
l(2)Sp9c, l(2)Spll, and l(2)Spl5. Gives mutant discoverer Thompson, 1957.
phenotype with rl but not stw or ap. references: 1963, DIS 38: 28.
genetics: Deficient for bs and ba but not sp. Homo-
cytology: Df(2R)41A;41C. zygous lethal.
origin: Associated with T(Y;2;3)D. Df(2R)Sw: Deficiency(2R) Swedish
*D«2RM-$2v9ih Deficiency^/?) Minute of Schultz cytology: Dl(2R)60F3-4; terminal deficiency.
2 from vestigial 77 origin: Natural condition of Swedish-b.
cytology: Df(2R)40F-41Al;42A19-B1. discoverer Catcheside, 36120.
origin: X ray induced; arose simultaneously with genetics: No phenotypic effect.
vgU. Df(2R)vgB: Defictency(2R) vestigial-Beaded
discoverer Ruch, 1931. cytology: Df(2R)49D3-4;S0A2-3.
synonym: Df(2R)M-vg11. ori g i n: Spontaneou s.
references: Morgan, Bridges, and Schultz, 1938, discoverer Bridges, 28dll.
Carnegie Inst. Wash. Year Book 37: 306. references: Morgan, Bridges, and Schultz, 1938,
Morgan, Schultz, Bridges, and Curry, 1939, Carnegie Inst. Wash. Year Book 37: 304—6.
Carnegie Inst. Wash. Year Book 38: 275. genetics: Deficient for vg and 1(2)C but not sea.
genetics: Deficient for rl, M(2)S2, stw, ap, tk, and Homozygous lethal.
msf but not ltd. Homozygous lethal. D1(2R)vgC: Deficiency(2R) vestigial-Carved
Dt(2R)MS4: see Di(2R)M-S2* cytology: Df(2R)49B2-3;49E7-Fl.
Df(2R)MS8: see Df(2R)M-S2s origin: X ray induced.
Df(2R)M-S10: see Df(2R)MS2*<> discoverer. Demerec, 28c3.
Df(2R)M-vgi i: see Df(2R)M-$2v6i' references: Morgan, Bridges, and Schultz, 1938,
*DK2R)Np: Deficiency^/?) Ho top I euro! Carnegie Inst. Wash. Year Book 37: 304—6.
cytology: Df(2R)44Fl-2;45El-2 (Bridges), genetics: Deficient for mca, vg, and I(2)C, Homozy-
origin: Spontaneous. lethal.
DK2R)vgt>: Deficiency(2R) vestigial-Depilate origin: X ray induced,

cytology: Df(2R)49Cl-2;49E2-6. discoverer: Demerec, 33J25.
discoverer: Bridges, 31a22. Coyne, 1935, DIS 4: 59.
references: Morgan, Bridges, and Schultz, 1938, Mossige, 1938, Hereditas 24: 110-16.
Carnegie Inst. Wash. Year Book 37: 304—6. genetics: Deficient for M(3)h; lethal homozygous
genetics: Deficient for sea, vg, and 1(2)C. Homozy- and in combination with Df(3L)Ly.
gous lethaf. *Df(3L)Mz: Deficiency(3L) from Minute(2) z stock
*DK2R)vgh Deficiency(2R) vestigial-Incised cytology: Loss of several bands from tip of 3L.
cytology: Dt(2R)49C2-Dl;S0A2-3. origin: Spontaneous.
discoverer Bridges, 36d20. discoverer: Bridges, 36h28.
references: Morgan, Bridges, and Schultz, 1938, DH3DN264-6: Deficiency(3L) Notch
Carnegie Inst. Wash. Year Book 37: 304—6. cytology: Df(3L)73E;8OC.
genetics: Deficient for sea, vg, and 1(2)C. Homozy- origin: Aneuploid segregant from T(l;3)N264-6/+.
gous lethal. *Df(3L)rul00.392: Deficiencyi3L) roughoid
Df(2R)vgS: DeiiciencyHR) vestigial-Snipped cytology: Df(3L)61Ej62A10-Bl.
cytology: Di(2R)49B12-Cl;49F15-50Al. origin: X ray induced.
origin: X ray induced. discoverer: Alexander.
discoverer: Muller, 1929. references: Ward and Alexander, 1957, Genetics 42:
references: 1930, J. Genet. 22: 299—334. 42-54.
Morgan, Bridges, and Schultz, 1938, Carnegie Inst. genetics: Deficient for ru. Homozygous lethal.
Wash. Year Book 37: 304-6. *Df(3L)rul00.393
genetics: Deficient for sea, vg, and l(2)C. Homozy- cytology: Df(3L)61F2-3;62A4-6.
gous lethal. origin: X ray induced.
*D«3L)D: Deficiency(3L) from T(Y;2;3)D references: Ward and Alexander, 1957, Genetics 42:
cytology. Df(3L)61E2-Fl;62A4-6. 42-54.
origin: Associated with T(Y;2;3)D. genetics: Deficient for ru. Homozygous lethal.
*D«3L)hl00.390: Deficiency(3L) hairy *D«3L)ru300.234
cytology: Di(3L)66D2-5;66D14-El. cytology: Dt(3L)61E;62A2-4.
discoverer. Alexander, discoverer. Alexander.
references: Ward and Alexander, 1957, Genetics 42: references: Ward and Alexander, 1957, Genetics 42:
42-54. 42-54.
genetics: Deficient for ft. Hornozygous lethal. genetics: Deficient for ru. Homozygous lethal.
Df(3L)Hn: Deficiency(3L) Henna
cytology: Df(3L)66A;66B. *Df(3L)ru-Kl: Deficiency(3L) roughoid of
origin: X ray induced simultaneously with Krivshenko
T(2;3ytin = T(2;3)53E-54A;77A;94F;96A. cytology: Di(3L)62A12-Bl;62B2-3.
discoverer. Van Atta, 30k. origin: X ray induced,
references: 1932, Am. Naturalist 66: 93—95. discoverer: Krivshenko, 1957.
1932, Genetics 17: 637-59. references: 1958, DIS 32: 81.
Lewis, 1956, DIS 30: 130. genetics: Has rough and slightly reduced eyes in
genetics: Mutant or deficient for Hn. Homozygous combination with ru but, judging from cytology,
lethal. probably not deficient for ru.
*Df(3L)K: Deficiency(3L) of Krivshenko *Df(3L)ru-K2
cytology: Df(3L)61 Cl-2; apparently a terminal de- cytology: Df(3L)61F4-S;62A10-Bl.
ficiency. origin: X ray induced.
origin: Probably X ray induced, discoverer: Krivshenko, 1957.
discoverer: Krivshenko, 5614. references: 1958, DIS 32: 81.
synonym: Df-SL^-. genetics: Deficient for ru.
references: 1959, DIS 33: 95. *Df(3L)sf100.62: Deficiency(3L) scarlet
D^3L)Ly: Defidency(3L) Lyra cytology: D£(3L)73A2-3;73A10-Bl.
cytology: Dt(3L)70A2-3;70A5-6. origin: X ray induced,
origin: X ray induced. discoverer: Alexander,
discoverer: Dubinin, 1929. references: Ward and Alexander, 1957, Genetics 42:
references: Morgan, Bridges, and Schultz, 1937, 42-54.
Carnegie Inst. Wash. Year Book 36: 301. genetics: Deficient for at. Homozygous lethal.
genetics: Associated with the mutant, Ly. Lethal *Df(3L)sfi oo.i7i
homosrfgous and in combination with cytology: D%3L)72E4-5;74C2-3.
*DH3L)M-h*3l: Deficimcy(3L) Minuto-h origin: X ray induced,
cytology: Probably includes bands in 70A. discoverer: Alexander.
references: Ward and Alexander, 1957, Genetics 42: genetics: Found in Dl but, judging from other infor-
42-54. mation placing the locus of Dl in region 91,
genetics: Deficient for st. Homozygous lethal. Df(3R)Dl is separable from Dl. Homozygous lethal.
*D«3L)sti 00.200
cytology: Df(3L)72E4-5;73A10-Bl. *DK3R)DIH
origin: X ray induced. cytology: Df(3R)91C6-Dl;92A2-3 (Slizynski).
discoverer: Alexander. origin: Induced by unspecified chemical,
references: Ward and Alexander, 1957, Genetics 42: discoverer: Auerbach.
42-54. references: 1943, DIS 17: 49.
genetics: Deficient for st. Homozygous lethal. genetics: Deficient for Dl. Homozygous lethal.
*Df(3R)e4.39: Deficiency(3R) ebony
*D«3L)thl00.i0S: Deficiency<3L) thread cytology: Df(3R)93B;93F.
cytology: Df(3L)72A2-Bl;73A4-5. origin: X ray induced.
origin: X ray induced. discoverer: Alexander.
discoverer: Alexander. references: 1960, Genetics 45: 1019-22.
references: Ward and Alexander, 1957, Genetics 42: genetics: Deficient for e. Homozygous lethal.
42-54. l 00.172
genetics: Deficient for th and st. Homozygous cytology: Df(3R)93B7-10;93F10-94Al.
*Df(3L)Vn: Deficiency(3L) Vein discoverer: Alexander.
cytology: Df(3L)64C12-Dl;65D2-El. references: Ward and Alexander, 1957, Genetics 42:
origin: Spontaneous. 42-54.
discoverer Mohr, 28J21. genetics: Deficient for e. Homozygous lethal.
references: 1932, Proc. Intern. Congr. Genet., 6th. *D«3R)ei 00.256
Vol. 1: 190-212. cytology: Df(3R)93A5-Bl;93F5-9.
1938, Norske Videnskaps-Akad. 4: 1-7. origin: X ray induced.
Mohr and Mossige, 1943, Norske Videnskaps-Akad. discoverer: Alexander.
7: 1-51 (fig.). references: Ward and Alexander, 1957, Genetics 42:
genetics: Deficient for jv, dv, and Me but not for 42-54.
a-3, Hn, or se. Mutant or deficient for Vn. Homo- genetics: Deficient for e. Homozygous lethal.
zygous lethal. D«3R)M-S31: De(iciency(3R) Minute of Schultz
*D«3R)89EF discoverer: Schultz, 33alO.
cytology: Dt(3R)89D7-El;90A2-3. references: 1940, DIS 13: 51.
origin: Synthetic. Made by combining the 3RD4P genetics: Termed deficiency on basis of Minute
element of one T(3;4) with the 4D3RP element of phenotype and mutant interaction with cu but not
another. ma. Homozygous lethal.
discoverer: Dubovsky and Kelstein. D%3R)Na: Deficiency(3R) from Naples
references: 1936, Eksperim. Med. No. 11: 65—84. cytology: Dt(3R)96Fll-97Al;97A2-S.
Kelstein, 1938, Biol. Zh. (Moscow) 7: 1145-69. origin: Associated with In(3R)Na.
Pipkin, 1959, Texas Univ. Publ. 5914: 69-88. Df(3R)PJ4: Deficiency(3R) from Pasadena
phenotype: Heterozygote (presumably with two X's) cytology: Df(3R)9OC2-Dl;91A2-3,
resembles female intersexes and is sterile. Male origin: X ray induced.
has rotated genitalia. discoverer. £. B. Lewis.
genetics: One of a series of deficiencies for the genetics: Deficient for sr and gl but not k or Dl.
middle of 3R synthesized and carefully studied by Homozygous lethal.
Dubovsky and Kelstein. Heterozygous male has Df(3R)ry: Deficiency(3R) rosy
rotated genitalia which may be feminized both in cytology: Df(3R)87D-E;87E-F,
structure and color; has sex combs. Heterozygous origin: X ray induced.
female sterile. discoverer: E. H. Grell, 1960.
references: 1962, Z. Vererbungslehre 93: 371—77.
Df(3R)bxdl00; D*ficiency(3R) bithoraxoid genetics: Deficient for kmr and ry. Homozygous
cytology: D%3R)89B5-6;89E2-3. lethal.
origin: Aneuploid recombinant from Tp(3)bxdi00/+. Df(3R)rY27
cytology: Dt(3R)9lC7-Dl;92A2-3. discoverer: Schalet.
origin: Aneuploid recombinant from Tp(3)bxdllo/+. references: 1964, DIS 39: 62-64.
Schalet, Kemaghan, and Chovnick, 1964, Genetics
D#3R)Dh Deficiency{3R) Delta 50: 1261-68.
cytology: D§(3R)90D2-4;9QD5-El. genetics: Deficient for 1(3)S3, m i , ry, pic, 1(3)54,
origin: Spontaneous. 1(3)S5, l(3)S6, 1(3)26, and 1(3)S7 but not 1(3)S1,
d i s c overer: Schultx. l(3)S2, or kar. Homozygous lethal.
*D«3R)ry28 references: 1964, DIS 39: 62-64.

origin: X ray induced in cu kar chromosome. Schalet, Kernaghan, and Chovnick, 1964, Genetics
discoverer: Schalet. 50: 1261-68.
references: 1964, DIS 39: 62-64. genetics: Deficient for kar, mes, arid ry but not for
Schalet, Kernaghan, and Chovnick, 1964, Genetics 1(3)S2, pic, 1(3)84, 1(3)S5, 1(3)S6, 1(3)26, or 1(3)S7.
50: 1261-68. Homozygous lethal.
genetics: Deficient for ry and 1(3)26. Homozygous
lethal. origin: X ray induced in cu kar chromosome.
*D«3R)ry29 discoverer: Schalet.
origin: X ray induced in cu kar chromosome. references: 1964, DIS 39: 62-64.
discoverer Schalet. Schalet, Kernaghan, and Chovnick, 1964, Genetics
references: 1964, DIS 39: 62-64. 50: 1261-68.
Schalet, Kernaghan, and Chovnick, 1964, Genetics genetics: Deficient for l(3)S2, kar, mes, ry, pic,
50: 1261-68. 1(3)S4, 1(3)S5, 1(3)S6, and 1(3)26 but not 1(3)S7.
genetics: Deficient for kar, mes, ry, pic, 1(3)S4, Homozygous lethal.
1(3)SS, and 1(3)S6 but not 1(3)S2, 1(3)26, or 1(3)S7. Df(3R)ryS2
Homozygous lethal. origin: X ray induced in cu kar chromosome.
*D«3R)ry3° discoverer: Schalet.
origin: X ray induced in cu kar chromosome. references: 1964, DIS 39: 62-64.
discoverer: Schalet. Schalet, Kernaghan, and Chovnick, 1964, Genetics
references: 1964, DIS 39: 62-64. 50: 1261-68.
Schalet, Kernaghan, and Chovnick, 1964, Genetics genetics: Deficient for 1(3)S3, mes, ry, pic, l(3)S4,
50: 1261-68. 1(3)S5, and 1(3)S6 but not 1(3)S2, kar, 1(3)26, or
genetics: Deficient for ry and 1(3)26 and probably 1(3)S7. Homozygous lethal.
for mes and pic. Homozygous lethal. *Df(3R)ry66
origin: X ray induced in cu kar chromosome. discoverer: Kernaghan.
discoverer: Schalet. references: Schalet, 1964, DIS 39: 62-64.
50: 1261-68. genetics: Deficient for mes, ry, pic, 1(3)S4, 1(3)S5,
genetics: Deficient for ry and probably mes and pic 1(3)S6, and 1(3)26 but not 1(3)S2, kar, or 1(3)S7.
but not for 1(3)26. Homozygous lethal. Homozygous lethal.
*DK3R)ry70
origin: X ray induced in cu kar chromosome. discoverer: Kernaghan.
discoverer: Schalet. references: Schalet, 1964, DIS 39: 62-64.
50: 1261-68. genetics: Deficient for mes, ry, pic, 1(3)S4, 1(3)S5,
genetics: Deficient for ry and 1(3)26 and probably and 1(3)S6 but not for 1(3)S2, kar, 1(3)26, or 1(3)S7.
formes and p i c Homozygous lethal. Homozygous lethal.
origin: X ray induced in cu kar chromosome. origin: X ray induced.

references: 1964, DIS 39: 62-64. genetics: Deficient for the right portion of the mes
Schalet, Kernaghan, and Chovnick, 1964, Genetics region and for ry but not for 1(3)S3, the left portion
50: 1261-68. of the mes region, or pic. Homozygous lethal.
genetics: Deficient for kar, mes, ry, pic, 1(3)S4, *Di(3R)ry75
1(3)S5, and 1(3)S6 but not 1(3)S2, 1(3)26, or 1(3)S7. origin: X ray induced in kar2 chromosome.
Homozygous lethal. discoverer: Schalet.
genetics: Deficient for 1(3)S3, mes, ry, pic, 1(3)S4,
origin: X ray induced in cu kar chromosome. and 1(3)S5 but not kar or l(3)S6. Homozygous
discoverer. Schalet. lethal.
Schalet, Kernaghan, and Chovnick, 1964, Genetics origin: X ray induced in kar2 chromosome.
50: 1261-68. discoverer Schalet.
genetics: Deficient for ry and probably mes and pic genetics: Deficient for 1(3)S2, kar, 1(3)S3, mes, ry,
but not for 1(3)26. Homozygous lethal. pic, 1(3)S4, and 1(3)S5 but not for 1(3)S1, 1(3)S6, or
1(3)26. Homozygous lethal.
discoverer: Schalet. origin: X ray induced in kar2 chromosome.
discoverer: Schalet. discoverer: Gloor and Green, 1957.

genetics: Deficient for 1(3)S3, mes, ry, and pic but synonym: 1(4)11.
not 1(3)S1, 1(3)S2, kar, or 1(3)26. Homozygous references: Hochman, Gloor, and Green, 1964,
lethal. Genetica 35: 109-26.
*Df(3R)ry78 genetics: Permits pseudodominant expression of sv
origin: X ray induced in kar2 chromosome. and spa alleles. Heterozygote with spa shows
discoverer: Schalet. slight sparkling effect but the effect is extreme in
genetics: Deficient for mes, ry, pic, and 1(3)26 but heterozygote with spaPoJ. Lethal homozygous and
not for kar, or 1(3)S3. Homozygous lethal. with spaCat, 1(4)9, 1(4)29, Df(4)3, Di(4)12, Df(4)24,
Df(3R)ryK: Deficiency(3R) rosy of Kernaghan Df(4)34, and Df(4)G.
origin: X ray induced in cu kar chromosome. D«4)12
discoverer: Kernaghan. origin: X ray induced.
references: 1964, DIS 39: 62—64. discoverer: Gloor and Green, 1957.
Schalet, Kernaghan, and Chovnick, 1964, Genetics synonym: 1(4)12.
50: 1261-68. references: Hochman, Gloor, and Green, 1964,
genetics: Deficient for 1(3)S1, 1(3)S2, kar, l(3)S3, Genetica 35: 109—26.
mes, ry, pic, 1(3)S4, 1(3)S5, 1(3)S6, and 1(3)26 but genetics: Heterozygote with spa is wild type and
not l(3)S7. Homozygous lethal. with spaPo1 is slight sparkling. Lethal homozy-
Df(3R)sbdlOS: Deficiency(3R) stubbloid gous and with spa^**, 1(4)9, 1(4)29, Df(4)3,
cytology: Df(3R)88F9-89Al;89B4-5. Df(4)ll, Dt(4)24, Di(4)34, and Df(4)G.
origin: X ray induced. Df(4)17
discoverer: E. B. Lewis. origin: X ray induced.
references: 1948, DIS 22: 72-73. discoverer: Gloor and Green.
genetics: Deficient for c(3)G, sbd, and Sb but not synonym: 1(4)17.
kar, cv-c, ss, or bx. Df(3R)sbd* 05/+ shows de- references: Hochman, Gloor, and Green, 1964,
creased crossing over in X (Hinton, 1966, Genetics Genetica 35: 109-26.
55: 157—64). Homozygous lethal. genetics: Lethal homozygous and when heterozy-
*Df(3R)sri00.394; Deficiency(3R) stripe gous with ciD or Ce 2 . Does not interact with any
cytology: Df(3R)90C2-7;90F3-7. other factor in Df(4)M.
origin: X ray induced. Df(4)24
discoverer: Alexander. origin: X ray induced.
references: Ward and Alexander, 1957, Genetics 42: discoverer: Gloor and Green, 1957.
42-54. synonym: 1(4)24.
genetics: Deficient for sr. Homozygous lethal. references: Hochman, Gloor, and Green, 1964,
*D«3R)sr300.24 Genetica 35: 109-26.
cytology: Df(3R)90C2-4;91A2-5. genetics: Heterozygote with spa is sparkling and
origin: X ray induced. with apaPo1 is poliert. Lethal homozygous and
discoverer: Alexander. with spaCat, 1(4)9, 1(4)29, D((4)3, Di(4)ll, Df(4)12,
references: Ward and Alexander, 1957, Genetics 42: Df(4)34, and D£(4)G.
42-54. D«4)31
genetics: Deficient for sr. Homozygous lethal. origin: X ray induced.
*D«3R)sr300.J0i discoverer: Gloor and Green, 1957.
cytology: Df(3R)90D2-4;91A6-8. synonym: 1(4)31.
origin: X ray induced. references: Hochman, Gloor, and Green, 1964,
discoverer: Alexander. Genetica 35: 109-26.
references: Ward and Alexander, 1957, Genetics 42: genetics: Lethal homozygous and when heterozy-
42-54. gous with ciD or Ce 2 . Does not interact with any
genetics: Deficient for sr. Homozygous lethal. other factor in Df(4)M.
D«4)3 D«4)34
cytology: Di(4)102E;102F; inferred from genetic origin: X ray induced.
data. discoverer: Gloor and Green, 1957.
origin: X ray induced, synonym: 1(4)34.
discoverer. Gloor and Green, 1957. references: Hochman, Gloor, and Green, 1964,
genetics: Lethal in homozygote and in combination Genetica 35: 109-26.
with Df(4)G, Dt(4)ll, Dt(4)34, and spac«* but not genetics: Lethal homozygous and when heterozy-
1(4)9 or 1(4)29 (Hochman). gous with Df(4)G, Df(4)ll, Di(4)3, and ®paCat but
not with 1(4)9 or 1(4)29.
cytology: Df(4)102E2-10;102F2-10 (Hochman); sali- D44)G: Deficiency(4) of Gloor and Green
vary chromosome bands missing in sections 102E cytology: D£(4)102E2-10; tip of 4R lost and re-
and perhaps some from 102F, but distal tip of 4R mainder of chromosome 4 capped with X-chromo-
is present. somal material, including 1A (Hochman).

references: Hochman, Gloor, and Green, 1964, cytology: Df(4)101F2-102Al;102A2-5 (combined from
Genetica 35: 109-26. observations of Fahmy and Hochman).
genetics: Lethal homozygous and in heterozygous origin: Recovered from progeny of male injected
combination with Df(4)3, Df(4)ll, Df(4)12, Df(4)24, with thymus extract from leukemic mice (Gross
Df(4)34, spac*t, 1(4)9, and 1(4)29. Recessives at Factor).
sv and spa loci expressed fully in heterozygote. discoverer Fahmy, 63a.
Loci of y + and ac + linked to the chromosome. genetics: Deficient for ci and M(4) but not ar, gvl,
Df(4)M: Deficiency) Minute bt, ey, sv, spa, or the lethal effect of ciD. Homo-
cytology: Df(4)101E-F;102B6-17; right break to the zygous lethal.
left of 102B9-10 according to Bridges, to the right Df(w-ec)6*d: see Df(l)w-ec
according to Slyzinski (1944, J. Heredity 35: 3 2 2 - Doublet: see Dp(l;l)BSRMG
24). Hochman has not succeeded in distinguishing Dp(l)3Cl: see Dp(l;l)wTG
between these alternatives.
origin: Spontaneous. DUPLICATIONS
synonym: Df(4)M-4. *Dp(l;l)IOO: Duplication(l;l) 100
references: 1935, Biol. Zh. (Moscow) 4: 401—20. origin: Spontaneous product of exchange between
1935, Tr. Dinam. Razvit. 10: 469. Dp(l;f)100 and proximal heterochromatin of C(1)RM.
genetics: Deficient for at, ci, gvl, 1(4)1, 1(4)13, *Dp(l;1)105
1(4)18, 1(4)25, M(4), and Sen but not bt, 1(4)2, sv, cytology: Metaphase X chromosome has one arm of
or spa. Homozygous lethal. normal length and one about 40 percent normal
length.
origin: X ray induced. new order: 1 — 20|6 — 1.
discoverer Schultz, 32k29. origin: X-ray-induced deletion of most of A" euchro-
references: Bridges, 1935, Biol. Zh. (Moscow) 4: matin was recovered as a C(l)JRM/Dp(l;f)105 fe-
401-20. male, which by detachment produced Dp(l;l)105 in
genetics: Deficient for at, ci, and ciD. Homozygous the succeeding generation.
lethal. discoverer: Dobzhansky, 1930.
references: 1932, Biol. Zentr. 52: 493-509.
cytology: Dt(4)101E;102B; similar to Df(4)M genetics: Contains wild-type alleles of y through dx
(Bridges, 1935, Tr. Dinam. Razvit 10: 470). and also probably bb.
origin: X ray induced. Dpd;l)U2
discoverer: Schultz, 33a8. origin: Spontaneous product of exchange between
synonym: Df(4)M-43. Dp(l;i)112 and proximal heterochromatin of an
genetics: Deficient for at, ci, gvl, and M(4) but not attached X.
bt, ey, or sv. Homozygous lethal.
*Dp(l;l)m
origin: X-ray-induced deletion of most of X euchro-
matin that was recovered as a C(1)RM/Dp(l;f)138
discoverer: Glass, 42hl2.
female, which by detachment produced Dp(l;l)138
in the subsequent generations.
genetics: Deficient for Ce, ci, and M(4) but not for
discoverer: Dobzhansky, 1930.
ey or sv. Homozygous lethal.
cytology. Df(4)101E; 102D13-E1. Vererbungslehre 68: 134—62.
genetics: Extends from locus of r to base of X;
origin: Recovered among progeny of male injected
carries B. Female nearly wild type, but male has
with Drosophila DNA.
low viability and is sterile.
discoverer: Fahmy, 62e.
genetics: Deficient for at, ci, gvl, and ey; Fahmy
*Dp(1;1)258-46
cytology: Dp(l;l)2B4-7;3A4-6; reversed repeat
claimed it was also deficient for spa but not sv. If
(Sutton).
we assume that Df(4)M62e is a simple interstitial
new order 1 - 2B4|3A4 - 2B7|2B7 - 20 or
deficiency, Fahmy's claim contradicts all other
1 - 3A4J3A4 - 2B7|3A6 - 20.
evidence on the order of sv and spa. Hochman
finds that Dt(4^462e/BpaP°l is poliert. Homozy-
gous lethal.
genetics: Originally appeared as w but reverted to
w+. M(l}Bld, tw, bt, pn, kz, and gt not affected.
cytology: Dt(4)101E;W2B10-17 (Fahmy); Dt(4)101E-
Dp(l;l)B: Duplication(l;l) Bar
F;102B2-5 (Hochman).
cytology: Dp(l;l)15F9-16Al;16A7-Bl; a tandem du-
origin: Gamma ray induced,
plication (Bridges, 1936, Science 83: 210-11;
discoverer. Fahmy, 62f.
Muller, Prokofyeva-Belgovskaya, and Kossikov,
genetics: Deficient for BT, ci, gvl, and M(4) but not
1936, Dokl. Acad. Nauk SSSR 1: 87-88).
bt, ey, av, or spa. Homozygous lethal.
new order: 1 - 16A7J16A1 - 20.
CHROMOSOME ABERRATIONS - DUPLICATIONS 309
origin: Spontaneous. and the proximal heterochromatin of the 4DXP ele-

discoverer: Tice, 13b. ment of T(1;4)BS.
references: 1914, Biol. Bull. 26: 221-30. discoverer: Lindsley and Sandier.
genetics: Position effect for B, apparently resulting references: 1963. In Methodology in Basic Genetics,
from juxtaposition of 16A1 with 16A7, which may W. J. Burdette, ed. Holden-Day, Inc., pp. 3 9 0 -
undergo mutation to less extreme forms (e.g., B*). 403.
Produces normal and triplicated \_Dp(l;l)BB\ genetics: Ineffective in generating tandem acro-
products by unequal crossing over. centric compound X chromosomes.
*Dp(1;l)B263.28
cytology: Dp(l;l)15F9-16Al;16A3-4;16A6- Dp(l;l)BsTMG: Duplication(l;l) Bar of Stone
7;16A7-B1. Tandem Metacentrigenie
new order: 1 - 16A3|l6A7|l6Al - 20. cytology: Dp(l;l)15F9-16Al;20 added as a second
origin: X-ray-induced deletional derivative of arm to In(l)sc*LENR.
Dp(l;l)BlBi = Dp(l;l)15F9-16Al;16A7-Bl. new order: 1A - B2|20B - lA|20-20 - 16Al|lO2F.
discoverer: Demerec, 34b. origin: Spontaneous recombinant between the X in
references: Sutton, 1943, Genetics 28: 97—107. normal sequence of a C(1)TM and the 4DXP ele-
*Dp(l;l)B263.48 ment of T(1;4)BS.
cytology: Dp(l;l)3E2-3;15F9-16Al;20A2-3. discoverer: Lindsley and Sandier.
origin: Recombinant product from Tp(l)B263"•**/+. references: 1963. In Methodology in Basic Genetics,
W. J. Burdette, ed. Holden-Day, Inc., pp. 390—
Dp(l;l)BSRAG: Duplication(l;l) Bar of Stone 403.
Reversed Acrocentrigenic genetics: Generates tandem metacentric compound X
cytology: Dp(l;l)15F9-16Al;20. chromosomes in Dp(l;l)BsTMG/+ female by recom-
new order: -20|lA - 20|20 - 16Al|lO2F. bination between the duplication and the base of a
origin: Spontaneous recombinant between the distal homolog in normal sequence, at a rate of about
AT of a C(1)RA and 4DXP from T(1;4)BS. 20 X 1 0 - 4 .
discoverer: Lindsley and Sandier. Dp(l;l)BSTRG: Duplication(l;1) Bar of Stone
references: 1963. In Methology in Basic Genetics, Tanalem Ring-genic
W. J. Burdette, ed. Holden-Day, Inc., pp. 390— See C(l)TMBs, subsection on compound chromo-
403.
genetics: Generates reversed acrocentric compound
X chromosomes in Dp(l;l)BsRAG/+ female, usually cytology: Dp(l;l)4A5-Bl;4D2-3 + Dp(l;l)17A6-
by a double exchange in which one exchange oc- Bl;20B-C.
curs between the duplicated segment of one strand origin: Associated with In(l)bbDtl^ClR.
and the homologous region of its sister and the Dp(J;1)bbDfLy4R
other between the duplication-bearing X and its cytology: Dp(l;l)lA8-Bl;4D2-3 + Dp(l;l)18A3-
normal homolog. Rate of C(1)RA generation about 4;20B-C.
6 X10-4. origin: Associated with In(l)bbD{Ly4R.
Dp(l;l)BsRMG: Duplication(T;l) Bar of Stone Dp(l;l)BB: Duplication(l;1) Bar Bar
Reversed Metacentrigenie cytology: Dp(l;l)15F9-16Al;16A7-Bl; a tandem trip-
cytology: Dp(l;l)15F9-16Al;20. lication [Bridges, 1936, Science 83: 210-11 (fig.)].
new order: 1 - 20-20 - 16Al|lO2F. new order: 1 - 16A7|l6Al - 16A7J16A1 - 20.
origin: Spontaneous recombinant between C(1)RM origin: Spontaneous through unequal crossing over
and the 4®XP element of T(1;4)BS. i n f i / B female.
discoverer Muller. discoverer Zeleny.
synonym: Doubter, references: 1920, J. Exptl. Zool. 30: 292—324,
references: 1936, DIS 6: 8. Sturtevant, 1925, Genetics 10: 117-47.
Lindsley and Sandier, 1963. In Methodology in genetics: Either or both B regions may carry a less
Basic Genetics, W. J. Burdette, ed. Holden-Day, extreme derivative of B; e.g., BlB, BB1, or BlBl.
Inc., pp. 390-403. Number of duplicated segments may be either in-
genetics: Generates reversed metacentric compound creased or decreased by unequal crossing over.
X chromosomes in Dp(l;l)BsRMG/+ female by *Dp(1;l)Bt: Duplication(l;1) Branch!et
crossing over between the duplicated segment and cytology: Dp(l;l)3B2-Cl;6F6-7; tandem repeat
either the X to which it is attached or the homolo- (Darby).
gous X at a rate of about 2.5 X 10—4. new order. 1 — 6F4|3C1 — 20.
Dp(1;l)BSTAG: Duplication(ljl) Bar of Stone origin: Induced by P 3 2 .
Tandem Acrocentrigenic discoverer: Bateman, 1950.
cytology: Dp(l;!)15F9-16Al;20. references: 1950, DIS 24: 54.
new order «20 - 1AJ20 - 16Al|lO2F. 1951, DIS 25: 77.
origin: X-ray-induced recombinant between Hi® distal Dp(1;1)Bx': Dup!ication(J;1) Bzadex-recessive
heterochrot&atin of an X chromosome with a ter- cytology: Dp(l;l)17A;17E-F (E. B. Lewis).
minal beterochromatic segment derived from y*Y origin: Spontaneous.
discoverer: Ives, 35k.

references: 1937, DIS 7: 6. cytology: Dp(l;l)20B-C;20B-Dl.
Green, 1952, Proc. Natl. Acad. Sci. U.S. 38: 949- origin: Associated with In(l)scSLscL8R.
53.
1953, Genetics 38: 91-105. cytology: Dp(l;l)lA8-Bl;lB2-3 + Dp(l;l)18A3-
1953, Z. Induktive Abstammungs- Vererbungslehre 4;2OB-D1.
85: 435-49. origin: Associated with In(l)sc8Ly4R.
genetics: Duplicated for os+, Bx+, and iu+. Does Dp(l;l)sc260.2S
not yield unequal crossovers as does Bxr49k. origin: Aneuploid recombinant from In(lLR)sc26°-25m
Dp(l;l)Bxr49k Dp(l;l)sci-aL5c4R
cytology: Dp(l;l)17A;17C (E. B. Lewis). cytology: Dp(l;l)19F-20Cl;20B-C.
origin: Spontaneous. origin: Associated with In(l)scL8Lsc4R.
discoverer: Mossige, 49k22. Dp(T;l)sct-8LSc8R
references: 1950, DIS 24: 61. cytology: Dp(l;l)lB2-3;lB3-4.
Green, 1953, Z. Induktive Abstammungs- origin: Associated with In(l)scL8Lsc8R.
Vererbungslehre 85: 435—49. Dp(1;l)scS"-sc4R
genetics: Duplicated for Bx + but not for os+ or fu+. cytology: Dp(l;l)19F-20Cl;20B-Dl.
Unequal crossing over yields wild types and tripli- origin: Associated with In(l)scslLsc4R.
cations. Quadruplications have also been pro- Dp(T;l)scSii-sc8R
duced. cytology: Dp(l;l)lB2-3;lB3-4,
Dp(l;1)Cll-y4R origin: Associated with In(l)scSl^sc8R.
cytology: Dp(l;l)lA8-Bl;4A5-Bl. Dp(1;l)scS1LscL8R
origin: Associated with In(l)C]Ly4R. cytology: Dp(l;l)20B-C;20B-Dl.
Dp(l;l)Co: Duplication(l;l) Confluens origin: Associated with In(l)scslI-'scL8R.
cytology: Dp(l;l)3C4-5;3D6-El; tandem duplication
(Schultz, 1941, DIS 14: 54-55). : Duplication(l;l) scute of Valencia
new order: 1 — 3D6|3C5 — 20. cytology. Dp(l;l)lA8-C3.
origin: Recovered among progeny of cold-treated fly. origin: Aneuploid recombinant from In(lLR)scvl/+.
discoverer: Gottschewski, 34c. *Dp(l;1)Th: Duplication(l;l) Theta
references: 1937, Z. Induktive Abstammungs- origin: X-ray-induced detachment of C(1)RM with
Vererbungslehre 73: 131—42. X-ray-induced deletion of most of the X euchro-
1935, DIS 4: 7, 14, 16. matin.
1937, DIS 8: 12. d i s cov erer: Muller.
genetics: The Co phenotype arises from a duplica- references: Muller and Painter, 1929, Am. Naturalist
tion of the Notch locus (salivary band 3C7). 63: 197.
Dp(l;1)Hw: Duplication(l;l) Hairy wing Patterson, 1930, Genetics 15: 141-49.
cytology: Dp(l;l)lA8-Bl;lB2-3; tandem repeat Muller, 1932, Proc.Intern. Congr. Genet., 6th. Vol.
(Demerec and Hoover, 1939, Genetics 24: 68). 1: 213-55.
new order: 1A1 - lB2|lBl - 20. genetics: Fragment of X chromosome, including y + ,
origin: Spontaneous. sc+, and bb+ attached to right of X centromere.
discoverer: Bridges, 23cl2. Causes development of interalar bristle not ordi-
genetics: Duplication produces Hw phenotype. narily present in D. melanogaster (Stern, 1956,
*Dp(h1)NBB-8 Arch. Entwicklungsmech. Organ. 149: 1—25).
cytology: Dp(l;l)16A;17E; tandem duplication. Dp(J;J)w: Duplication(l;l) white
new order: 1 - 17E|l6A - 20. cytology: Dp(l;l)3A;3C.
origin: Spontaneous as nonrecombinant strand from new order 1 - 3C|3A - 20.
/ B os/+ B + mother. origin: Spontaneous as a recombinant from wctt/wsP.
discoverer: Peterson and Laughnan. discoverer. E. B. Lewis, 55j.
references: 1963, Proc. Natl. Acad. Sci. U.S. 50: references: 1957, DIS 31: 84.
126-33. genetics: Loci of w and r&t within duplicated sec-
genetics: Male viability reduced. Has a BB pheno- tion. Unequal crossing over gives normal and trip-
type. licated products. Quintuplication also produced.
Dp(l;l)w60h2i: see Dp(l;l)wG
cytology. Dp(l;l)lB2-3;lB3-4. Dp(l;l)w6 0h3o: seeDp(l}-l)wrG2
origin: Associated with In(l)sc4Lsc8R.
Dp(1;1)$c4i-y4R Dp(J;1)w": Duplication(l;l) white-apricot
cytology: Dp(l;l)lA8-Bl;lB3-4 + Dp(l;l)18A3- cytology: Dp(l;l)3A10-Bl;3C3-5 (Gersh, 1962,
4;19F-2OC1. Genetics 47: 1393-98).
origin; Associated with In(l)&c4I*y4R. new order: 1 - 3C3|3B1 - 20.
Dp(1;1)sc8LSc4R origin: Spontaneous from w^/w* female; recovered
cytology: Dp(l;l)l9F-20Cl;20B-Dl. once as a recombinant and once as a presumed
origin: Associated with Irv(l)scSLac4R. recombinant.
discoverer Green. *Dp(l'l)z4

references: 1959, Genetics 44: 1243-56. cytology: Dp(l;l)2B16-Cl;3B-Cl.
Dp(7;7VrG-* Duplication(l;l) white-recombinant new order: 1 - 3BJ3C1 - 2Cl|3Cl - 20.
of Green origin: X ray induced.
cytology: Dp(l;l)3A3-4;3Cl-2. discoverer: Gans.
new order: 1 - 3Cl|3A4 - 20. references: 1953, Bull. Biol. France Belg., Suppl.
origin: A regular product of asymmetric exchange 38: 1-90.
between 3C1-2 of a wa or wa2 chromosome and genetics: Duplication male viable and fertile. Homo-
3A3-4 of specific homologs. Reciprocal of zygous female viable but poorly fertile.
Dt(l)w*G. *Dp(hl)z8
discoverer: Green, 60h21. cytology: Dp(l;l)2B18-Cl;4B4-S.
synonym: Dp(l)3Cl; Dp(l;l)w60h21, new order 1 - 4B4|2C1 - 20.
Gersh, 1962, Genetics 47: 1393-98 (fig.). discoverer: Gans.
Dp(J;l)wrG2 references: 1953, Bull. Biol. France Belg., Suppl.
cytology: Dp(l;l)3B2-Cl;3C3-5 [Gersh, 1962, 38: 1-90.
Genetics 47: 1393-98 (fig.)]. genetics: Lethal in male.
new order: 1 - 3C3J3C1 - 20. Dp(1;l)z59d
origin: Spontaneous by recombination. cytology: Dp(l;l)2F5-3Al;3A4-5 (Gersh).
discoverer: Green, 60h30. new order: 1 - 3A4|3A1 - 20.
synonym: Dp(l;l)w^°h3 0^ origin: X ray induced in y2 su(wa) z.
Dp(l;l)wrJ2: Duplication(l;l) white-recombinant discoverer Green, 59dl5.
ofJudd references: 1961, Genetics 46: 1555-60.
cytology: Dp(l;l)3A6-8;3Cl-3 [could be same as Gersh, 1962, Genetics 47: 1393-98 (fig.).
Dp(l;l)wrG\.
new order: 1 - 3Cl|3A8 - 20. Dp(1;2)51b
origin: A regular product of unequal exchange be- cytology: Dp(l;2)3Cl-2;3D6-7;52F.
tween the 3C1-3 region of a chromosome carrying origin: Aneuploid segregant from T(l;2)51b/+.
wrdp with the 3A4-8 region of specific homologs. *Dp(l;2)A12: Duplication(l;2) from Austin
Probably reciprocal recombinant of Dt(l)wrJ3. cytology: Dp(l;2)7A;7B.
discoverer: Judd, 1961. origin: Aneuploid segregant from T(1;2;4)A12/+.
synonym: wT>duP. *Dp(l;2)A124
references: 1961, Proc. Natl. Acad. Sci. U.S. 47: cytology: Dp(l;2)10A;13Al-2;59.
545-50. origin: Segregant from T(1;2)A124/+.
*Dp(I;2)cf7c?; Duplication(l;2) cut
cytology: Dp(l;l)lA;lB2-3. cytology: Dp(l;2)7B2-3;8E2-3;25C.
origin: Associated with In(l)y3PLsc8R. origin: Aneuploid segregant from T(l;2)ct7cl/+.
Dp(l;l)yt>l: Duplication(l;l) yellow-bristle Dp(l;2)Kl: Duplication(T;2) of Krivshenko
cytology: Dp(l;l)lB2-3;4F8-9;5D4-5 (Nicoletti, cytology: Dp(l;2)lA5-B3;2Q;29A.
Lindsley). origin: Associated with T(1;2)K1.
new order 1A - 1B2J5D4 - 4F9|lB3 - 20. Dp(l;2)scl9: Duplication(l;2) scute
origin: Spontaneous. cytology: Dp(l;2)l Bl-2;lB4-7;25-26.
discoverer: Sandier. origin: Aneuploid segregant from T(l;2)sciS>/+.
references: Sandier, Hart, and Nicoletti, I960, DIS *Dp(l;2)w"'52b12: Duplication(l;2) white-mottled
34: 103-4. cytology: Dp(l;2)lE5-Fl;3C3-4;40-41.
genetics: Mutant for y; duplicated for cv. Regularly origin: Aneuploid segregant from TXl;2)wtaS2bl2/+.
generates further rearrangements; has produced
losses of the duplicated segment, which are cytol ogy: Dp(l;2)3B2-Cl;3C9-D1;4O-41,
accompanied by changes In phenotype from ybl to origin: Aneuploid segregant from Tf7;2>wn'53«/-K
y-like and a translocation between the tips of X Dp(l;2)wm2S8.44
and 2L accompanied by a change from ybl to y + , cytology: Dp(l;2)3C3-4;4D2-El;56El-Fl,
*Dp(1;J)zh Duplication(l;1) zeste origin: Aneuploid segregant from
cytology: Dp(l;l)lE2-3;4B4-5; tandem repeat.
new order 1A - 4B4|lE3 - 20.
origin: X ray induced. Dp(l;3)126
discoverer Gans. origin: X ray induced.
references: 1953, Bull. Biol. France Belg., Suppl. discoverer Dobzhansky, 1930.
38: 1-90. references: 1935, Z. Induktive Abstammungs-
genetic*: Male lethal. Vererbungslehre 68: 143.
*Dp(l;l)z2 genetics: Duplicated for r, M(l)o, f, and B but not si
cytology: Dp(l;l)2ClQ-Dl;4D2'4. or o«; variegates for / and Mffjo (Schultz). Dupli-
origin: Associated with Jn(l)m2. cated section inserted into chromosome 3 between
st and cu. Also an inversion in 3L. Viability.,fer- *Dp(1;4)N264.86

tility, and phenotype of Dp(l;3)126/+ male and cytology: Dp(l;4)3C6-7;3E5-6;10lF-102.
female normal. origin: Aneuploid segregant from T(l;4)N264-86/+.
*Dp(J;3)ct11<>: Duplication(l;3) cut
cytology: Dp(l;3)lB;7B2-3;84B. Dp(l;4)r+: Duplication(l;4) rudimentary-wild type
origin: Aneuploid segregant from T(l;3)ctlla/+. origin: X-ray-induced derivative of T(1;4)BS =
*Dp(J;3)cti2ci T(l;4)16A6-Al;102F2-3; probably a deletion of
cytology: Dp(l;3)7B2-3;7D2-6;85. most of the X euchromatin from the XD4P element.
origin: Aneuploid segregant from T(l;3)ct12cl/+. discoverer Green.
references: 1963, Genetica 34: 242-53.
cytology: Dp(l;3)5D2-3;7B2-3;80C-F. genetics: Carries normal alleles of r and /appended
origin: Aneuploid segregant from T(l;3)ct26s'37. to the right end of chromosome 4.
Dp(J;3)K2: Duplication(l;3) of Krivshenko Dp(J;4V m57e .' Duplication(l;4) white-mottled
cytology: Dp(l;3)20A-B;20D-F;80-8l. cytology: Dp(l;4)3C2-3;3C4-7;101.
origin: Associated with T(1;3)K2. origin: Aneuploid segregant from T(l;4)wm51c/+-
Dpd;3)NS0k11: Duplication(l;3) Nofch
cytology: Dp(l;3)lE3-4;3C6-7;89A. Dp(l;4)wvD1: Duplication(l;4) white-variegated
origin: Aneuploid segregant from T(l;3)Ns0kl1 /+. of Demerec
cytology: Dp(l;4)3Cl-4;101A-D.
cytology: Dp(l;3)3B2-3;3D6-7;80D-F. origin: X ray induced in y.
origin: Aneuploid segregant from T(l;3)N264-58/+, discoverer: Demerec 33J19.
synonym: Dp(l;3)wm264-S8. genetics: Variegated for w but not ci. X broken be-
*Dp(l;3)N264.ioo tween w and rst; 4 probably broken in left arm.
cytology: Dp(l;3)3B4-Cl;4B4-5;80.
origin: Aneupioid segregant from T(l;3)N264-100/+. *Dp(l;A)pn-ec: Duplicationfl;Autosome) prune to
Dp(l;3)04: Duplication(l;3) of Oliver echinus
origin: Aneuploid segregant from T(1;3)O4/+. cytology: Dp(l;A)2Dl-2;3F7-4A4;40-41 or 80-81.
Dp(l;3)rasv: Duplication(l;3) raspberry-variegated origin: Aneuploid segregant from T(l;A)pn-ec.
cytology: Dp(l;3)9E;13C;81F.
origin: Aneuploid segregant from T(l;3)rasv/+. *Dp(l;f)l: Duplicationd;free)
*Dp(1;3)sc260-20: Duplica1ion(l;3) scute origin: X-ray-induced deletion of most of X euchro-
cytology: Dp(l;3)lA8-Bl;61Al-2. matin.
origin: Aneuploid segregant from T(l;3)sc260-20/+. discoverer: Muller.
synonym: Del(l)l.
Dp(1;3)scJ4: Duplication(l;3) scute of Jacobs- references: Painter and Muller, 1929, J. Heredity
duller 20: 287-98.
cytology: Dp(l;3)lB;61A. Muller and Painter, 1932, Z. Induktive Abstam-
origin: Aneuploid segregant from T(l;3)scJ*/+. mungs- Vererbungslehre 62: 316—65.
DtfJtfhnl**1: Duplication(l;3) singed genetics: Contains wild-type alleles of y, sc, br,
cytology: Dp(l;3)6C;7C9'Dl;72D2-El. pn, and 66.
origin: Aneuploid segregant from T(l;3)sni3al/+. *0p(l:f)2
Dp(l;3)sta: Duplication(l;3) stubarista origin: X-ray-induced deletion of most of X euchro-
cytology: Dp(l;3)lD3-El;2A;89B21-C4. matin.
origin: Aneuploid segregant from T(l;3)8ta/+. discoverer. Muller.
Dp(J;3)w»49a: Duplication(l;3) white-mottled synonym: Det(l)2.
cytol ogy: Dp(l;3)3A 10-Bl;3E2-3;80. references: Painter and Muller, 1929, J. Heredity
origin: Aneuploid segregant from T(l;3)wm49a/+^ 20: 287-98.
Muller and Painter, 1932, Z. Induktive Abstam-
Dp(1;3)wvce: Dvplication(l;3) white-variegated mungs- Vererbungslehre 62: 316—65.
cobbled genetics: Contains wild-type alleles of y, sc, br,
cytology: Dp(l;3)2B17-Cl;3C4-5;77D3-5;81. and 66.
origin: Aneuploid segregant from T(l;3)wyco/+. Dp(l;f)3
cytology: Dp(l;f)lD;19-20 (Gersh) 3.7-4 times the
*Dp(l;4)Ah buplicatioMb*) from Austin size of chromosome 4 at metaphase; lacks only the
cytology: Dp(l;4)9B;20;101-102. distalmost part of heterochromatic segment hD
origin: Aneuploid segregant from T(1;4)A1/+. (Cooper).
*Dp(l;4)A12 origin: X-ray-induced deletion of most of X euchro-
cyrology: Dp(l;4)lB-C;7A;7B;13B1 -5; 101-102. matin.
origin: Anettploid segregant from T(1;2;4)A12/+. discoverer: Weltroan, 1954.
*Dp(l;4)N2"'*S; QwpUcation(h4) Notch references; Lindsley and Sandier, 1958, Genetics
cytol of y: Dp(l;4)3B4-Cl?6A 2-Bl;101 F-102A. 43: 547-63.
origin: Anettpioid segregaot from TXl;2;4yNi*4'95. Grell, 1964, Genetics 50: 151-66.
genetics: Carries wild-type alleles of y, ac, sc, Dp(l;f)52

su(f), and 66 but not s^w*), dor, pn, or car. Dis- cytology: Dp(l;f)lB10-C4;19-20 (Gersh). 3.7-4
joins regularly from XY, 3 percent nondisjunction times the size of chromosome 4 at metaphase;
from C(l)RMt and causes 18 percent nondis junction lacks only the distalmost part of heterochromatic
otIn(l)dl-49 from + in In(l)dt-49/+/Dp(l;f)3 female. segment hD (Cooper).
*Dp(l;f)10 origin: X-ray-induced deletion of X euchromatin.
origin: X-ray-induced deletion of most of X euchro- discoverer: Weltman, 1954.
matin. references: Lindsley and Sandier, 1958, Genetics
discoverer: Weltman, 1954. 43: 547-63.
references: Lindsley and Sandier, 1958, Genetics genetics: Contains wild-type alleles of y, ac, sc,
43: 547-63. su(f), and 66 but not su(wa), pn, or car. Segregates
genetics: Carries wild-type alleles of y, ac, and sc normally from XY, 3 percent nondisjunction from
but not su(f) or 66. 61 percent nondisjunction from C(1)RM, and causes 13 percent nondisjunction of +
XY, 45 percent from C(1)RM, and regular disjunc- from In(l)dl-49 in In(l)dl-49/+/Dp(l;f)52.
tion of + from In(l)dl-49 in In(l)dl-49/+/Dp(l;f)10 Dp(l;f)60g
female. origin: A spontaneous exchange between the distally
located heterochromatin of In(l)sc8, y3ld anc j fae
*Dp(l;f)12 proximal heterochromatin of a normal X. Occurred
cytology: 3.4—4 times the size of chromosome 4 at in a triploid female.
metaphase; lacks only the distalmost part of heter- discoverer: Mohler, 60g.
ochromatic segment hD (Cooper). references: 1960, DIS 34: 52.
origin: X-ray-induced deletion of most of X euchro- genetics: Carries y3id, ac + , and su(f)+ but not car*.
matin. other information: The reciprocal product, a reversed
discoverer. Weltman, 1954. acrocentric compound X [C(l)RA60g\ was recovered
references: Lindsley and Sandier, 1958, Genetics from the same fly.
43: 547-63.
genetics: Contains wild-type alleles of y, ac, sc, cytology: Two-thirds the length of normal X at meta-
su(wa), su(f), and 66 but not pn or car. Disjoins phase.
regularly from XY, 4 percent nondisjunction from origin: Spontaneous deletion of most of X euchro-
C(1)RM, and causes 19 percent nondisjunction of + matin.
from In(l)dl-49 in In(l)dl-49/+/Dp(l;f)12 female. discoverer: L. V. Morgan, 221.
*Dp(l;f)U synonym: sc-Dp.
origin: X-ray-induced deletion of most of X euchro- references: 1938, Genetics 23: 423—62.
matin. genetics: Contains wild-type alleles of y, ac, sc,
discoverer: Muller. svr, sta, tw, br, pn, hi, car, and 66 but not gt, w,
synonym: Del(l)14. os, or Bx. Phenotype of duplication-bearing female
references: Painter and Muller, 1929, J. Heredity nearly wild type, but occipital bristles and hairs
20: 287-98. are present, eyes are a trifle smaller and rougher,
Muller and Painter, 1932, Z. Lnduktive Abstam- and wings have straighter outer margins and some-
mungs- Vererbungslehre 62: 316—65. times scalloped inner margins. In male, duplica-
genetics: Contains wild-type alleles of y, sc, and tion more than 99 percent lethal.
66 but not 6r.
cytology: Dp(l;f)2A2-Bl;19F5-2QA; one-fourth the
Dp(1;f)18 length of normal X at metaphase.
cytology: Dp(l;t)lF-2A; 19-20 (Gersh). origin: X-ray-induced deletion of most of X euchro-
origin: X-ray-induced deletion of most of X euchromatin.
matin. discoverer Dobzhansky, 1930.
discoverer Weltman, 1954. references: 1932, Tr. Lab. Genet. (Leningrad) 9:
references: Lindsley and Sandier, 1958, Genetics 193-216.
43: 547-63. 1935, Z. lnduktive Abstammungs- Vererbungslefare
genetics: Contains wild-type alleles of y, ac, sc, 68: 134-62.
mi(w*), and 66 but not pn or car. genetics: Contains wild-type alleles of y, sc, svr,
Dp(J;f)24 ®u(a), dor, and 66 but not kz or car (Schulte and
cytology: Dp(l;f)lA-B;l9~20 (Gersh). Bridges, 1932, Am. Naturalist 66: 323—34; Lewis,
origin: X-ray-induced deletion of most of X euchro- 1954, J. Exptl. Zool. 126: 235-75). With duplica-
matin. tion, both sexes viable and wild type except for
discoverer: Muller. presence of occipital bristles.
synonym: Del(l)24, *Dp(1;f)W2
references: 1932, Proc. Intern. Congr. Genet., 6th. cytology: One-fifth the length of normal X at meta-
Vol. 1: 213-55. phase.
genetics: Contains wild-type alleles of 1(1}JI, y, origin: X-ray-induced deletion of most of X euchro-
and ac but not matin.
discoverer: Dobzhansky, 1930. references: 1935, Z. Induktive Abstammungs-

references: 1932, Biol. Zentr. 52: 493-509. Vererbungslehre 68: 134—62.
1935, Z. Induktive Abstammungs- Vererbungslehre genetics: Contains wild-type alleles of y, sc, svr,
68: 134-62. and br; variegates for y, sc, and svr. Both sexes
genetics: Contains y+ to rb+ inclusive and not bb+. viable and wild type except for presence of occip-
Usually male lethal, but female survives and has ital bristles.
occipital bristles, narrow parallel-sided wings, Dp(l;{)135
branched posterior crossveins, and heavier bristles origin: X-ray-induced deletion of most of X euchro-
on thorax. matin from y2.
*DP(l;f)l06 discoverer: Dobzhansky, 1930.
cytology: Metaphase length about four times that of references: Sivertzev-Dobzhansky and Dobzhansky,
chromosome 4. 1933, Genetics 18: 173-92.
origin: X-ray-induced deletion of most of X euchro- genetics: Contains y2 and wild-type alleles of sc,
matin. SVT, and bb. Both sexes wild type except for
discoverer. Dobzhnasky, 1930. presence of occipital bristles.
genetics: Contains wild-type alleles of y, sc, and cytology: Metaphase length about one-fourth that of
svr but not bb. normal X.
Dp(l;f)107 origin: X-ray-induced deletion of most of X euchro-
cytology: Metaphase length about one-fifth that of a matin from y2.
normal X. discoverer: Dobzhansky, 1930.
origin: X-ray-induced deletion of most of X euchro- references: 1932, Biol. Zentr. 52: 493-509.
matin. genetics: Contains y2 and wild-type alleles of sc,
discoverer: Dobzhansky, 1930. svr, kz, pn, and bb. Variegates for y (Schultz).
references: 1932, Biol. Zentr. 52: 493-509. Viability low. Shows spread wings and occipital
genetics: Contains wild-type alleles of y, sc, svr, bristles.
and bb. *Dp(l;f)137
Dp(l;f)112 cytology: About one-fifth the length of normal X at
cytology: Dp(l;t)lE4-Fl;19-20 (Gersh); slightly metaphase.
longer than chromosome 4 at metaphase. origin: X-ray-induced deletion of most of X euchro-
origin: X-ray-induced deletion of most of X euchromatin from y2.
matin. discoverer: Dobzhansky, 1931.
discoverer. Dobzhansky, 1930. references: 1932, Biol. Zentr. 52: 493-509.
references: 1932, Biol. Zentr. 52: 493-509. genetics: Contains y2 through w+ but not bb+.
genetics: Contains wild-type alleles of y, sc, svr,
and au(w*) but not bb. Both sexes viable and have *Dp(l;f)U3-3
occipital bristles. origin: Associated with T(l;3)143-3.
Dp(l;0164
cytology: About one-fourth the length of normal X at cytology: Dp(l;f)lB; 19-20 (Gersh).
metaphase. origin: X-ray-induced deletion of most of X euchro-
matin. discoverer. Weltman, 1954.
discoverer Dobzhansky, 1930. references: Lindsley and Sandier, 1958, Genetics
references: 1932, Biol. Zentr. 52: 493-509. 43: 547-63-
genetics: Contains wild-type alleles of y, sc, svr, genetics: Carries wild-type alleles of y and ac but
and bb but not kz. not sc, su(wa), pn, car, su(f), or bb. Disjoins
Dp(I;f)122 essentially randomly from XY, 36 percent nondis-
cytology: Dp(l;t)lE4-Fl;19-20 (Gersh). junction from C(1)RM, and does not interfere with
origin: X-ray-induced deletion of most of X euchro- disjunction of + from In(l)dl-49 in In(l)dl-
matin. 49/+/Dp(l;t)164 female.
discoverer Weltman, 1954. *Dpd;f)U7
references: Lindsley and Sandier, 1958, Genetics cytology: 3.7—4 times the size of chromosome 4 at
43: 547-63. metaphase; lacks only the distalmost heterochro-
genetics: Carries wild-type alleles of y, ac, sc, and matic segment hD (Cooper).
bb but not &u(w*), pn, car, or su(f). Disjoins reg- origin: X-ray-induced deletion of most of X euchro-
ularly from XY, 6 percent nondisjunction from matin.
C(t)RM, and causes 9 percent nondisjunction of + discoverer Weltman, 1954.
from In(l)dl-49 in In(l)dl-49/+/Dp(l,f)122. references: Lindsley and Sandier, 1958, Genetics
*Dp(l;0134 43: 547-63.
origin: X-ray-induced deletion of most of X euchro- genetics: Carries wild-type alleles of y, ac, sc,
matin. suCw*), &u(f), and bb but not pn or car. Disjoins
discoverer: Dobzhansky, 1931. regularly from XY, 3 percent nondisjunction from
CHROMOSOME ABERRATIONS- DUPLICATIONS 315
C(1)RM, and causes 16 percent nondisjunction of + origin: X-ray-induced deletion of most of X euchro-
from In(l)dl-49 in In(l)dl-49/+/Dp(l;f)167 female. matin.
Dp(l;f)J79 discoverer Krivshenko and Cooper, 1954.
origin: X-ray-induced deletion of most of X euchro- references: Grell, 1964, Genetics 50: 151-66.
matin. 1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32.
discoverer: Weltman, 1954. genetics: Carries wild-type alleles of y, ac, sc, svr,
references: Lindsley and Sandier, 1958, Genetics and 66 but not su(wa), dor, pn, or stiff). Causes
43: 547-63. 2.4 percent nondisjunction of chromosome 4 when
genetics: Carries wild-type alleles of y, ac, sc, and added to normal diploid female genotype.
8u(w°) but not pn, car, su(f), or 66. Disjoins reg- Dp(l;M144
ularly from XY, 20 percent nondisjunction from cytology: Invisible in salivary preparations
C(1)RM, and causes 2 percent nondisjunction of + (Krivshenko); 1.1 times the length of chromosome 4
from In(l)dl-49 in In(l)dl-49/+/Dp(l;f)179 female. at metaphase; lacks nucleolus organizer (Cooper).
*Dp(l;{)749 origin: X-ray-induced deletion of most of X euchro-
cytology: Dp(l;t)lB12-13;2Q; frequently associated matin.
with nucleolus in salivary preparations discoverer Krivshenko and Cooper, 1954.
(Krivshenko); 3—4 times the length of chromosome references: Grell, 1964, Genetics 50: 151—66.
4 at metaphase; has nucleolus organizer (Cooper). 1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32.
origin: X-ray-induced deletion of most of X euchro- genetics: Carries wild-type alleles of y and ac but
matin. not sc, svr, suCw0), dor, pn, su(f), or 66. Causes
discoverer Krivshenko and Cooper, 1953. 36.6 percent nondisjunction of chromosome 4 when
genetics: Carries wild-type alleles of y, ac, sc, added to normal diploid female genotype.
svr, and 66 but not pn.
Dp(l;f)797 cytology: Twice the length of chromosome 4 at
cytology: Dp(l;f)2B4-5;20; frequently associated metaphase; has nucleolus organizer (Cooper).
with nucleolus in salivary preparations origin: X-ray-induced deletion of most of X euchro-
(Krivshenko); 2—3 times the length of chromosome matin from In(l)sc8.
4 at metaphase; has nucleolus organizer (Cooper). discoverer Krivshenko and Cooper, 1954.
origin: X-ray-induced deletion of most of X euchro- genetics: Carries wild-type alleles of y, ac, and 66
matin. but not sc, svr, or pn.
discoverer Krivshenko and Cooper, 1953. Dp(l;{)1156
genetics: Carries wild-type alleles of y, ac, sc, cytology: 2.6 tiroes the length of chromosome 4 at
svr, and bb but not pn. metaphase; has nucleolus organizer (Cooper).
origin: X-ray-induced deletion of most of X euchro-
matin from In(l)sc8.
cytology: Invisible in salivary preparations discoverer Krivshenko and Cooper, 1955.
(Krivshenko); 0.7 the length of chromosome 4 at genetics: Carries wild-type alleles of y, ac, and 66
metaphase; lacks nucleolus organizer (Cooper). but not sc or pn.
origin: X-ray-induced deletion of majority of X Dp(T;f)1158
euchromatin. cytology: 2.3 times the length of chromosome 4 at
discoverer: Krivsbenko and Cooper, 1953. metaphase; has nucleolus organizer (Cooper).
references: Grell, 1964, Genetics 50: 151—66. origin: X-ray-induced deletion of most of X euchro-
1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32, matin from In(l)sc8.
genetics: Carries wild-type alleles of y, ac, and sc discoverer Krivshenko and Cooper, 1954.
but not mj(wa), dor, pn, su(f), or 66. Causes 10 genetics: Carries wild-type alleles of y, ac, and 66
percent nondisjunction of chromosome 4 when but not sc or pn.
added to normal diploid female genotype.
Dp(1;f)819 cytology: 2.7 times the length of chromosome 4 at
cytology: Dp(l;f)lD3-4;20; usually associated with metaphase; has nucleolus organizer (Cooper).
nucleolus in salivary preparations (Krivshenko); origin: X-ray-induced deletion of most of X euchro-
2.9 times the length of chromosome 4 at metaphase; matin from In(l)sc8.
has nucleolus organizer (Cooper). discoverer: Krivshenko and Cooper, 1954.
origin: X-ray-induced deletion of most of X euchro- genetics: Carries wild-type alleles of y and ac but
matin. not sc or pn.
discoverer: Krivshenko and Cooper, 1953.
genetics: Carrie* wild-type alleles of y, ac, sc, @vr, cytology: 3.1 times the length of chromosome 4 at
and bb but not pn. metaphase; has nucleolus organizer (Cooper).
Dp(hM56 origin: X-ray-induced deletion of most of X euchro-
cytology; Dp(l;f)lD3-4;2O: frequently associated matin from In(l)scs.
wife raadeoiaa in salivary preparations discoverer Krivshenko and Cooper, 1955.
(Krivsbenko); three times the length of chromosome genetics: Carries wild-type alleles of y, ac, and 66
4 at neta-phase; has nucleolus organiser (Cooper). but not sc or pn.
Dp(l;f)1162 Dpf 7/01797

cytology: One-half the length of chromosome 4 at cytology: Seven-tenths the length of chromosome 4
metaphase; lacks nucleolus organizer (Cooper). at metaphase; lacks nucleolus organizer (Cooper).
origin: X-ray-induced deletion of most of X euchro- origin: X-ray-induced deletion of most of X euchro-
matin from In(l)sc8. matin from In(l)sc8.
discoverer. Krivshenko and Cooper, 1954. discoverer: Krivshenko and Cooper, 1954,
references: Grell, 1964, Genetics 50: 151—66. genetics: Carries wild-type alleles of y and ac but
1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32. not sc, pn, or 66.
genetics: Carries wild-type alleles of y and ac but Dp(l;f)im
not sc, su(wa), dor, pn, su(f), or 66. Causes 3.8 cytology: Same length as chromosome 4 at meta-
percent nondisjunction of chromosome 4 when phase; lacks nucleolus organizer (Cooper).
added to normal diploid female genotype. origin: X-ray-induced deletion of most of X euchro-
*Dp(hf)1170 matin from In(l)sc8.
cytology: 1.9 times the length of chromosome 4 at discoverer: Krivshenko and Cooper, 1954.
metaphase; lacks nucleolus organizer (Cooper). references: Grell, 1964, Genetics 50: 151-66.
origin: X-ray-induced deletion of most of X euchro- 1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32.
matin from In(l)sc8. genetics: Carries wild-type alleles of y and ac but
discoverer Krivshenko and Cooper, 1954. not sc, su(wa), dor, pn, su(f), or 66. Causes 22.8
genetics: Carries wild-type alleles of y and ac but percent nondisjunction of chromosome 4 when
not sc, pn, or bb. added to normal diploid female genotype.
Dp(J;f)n94
Dp(l;f)1173 cytology: 3.1 times the length of chromosome 4 at
cytology: 3.2—3.6 times the length of chromosome 4 metaphase; has nucleolus organizer (Cooper).
at metaphase; has nucleolus organizer (Cooper). origin: X-ray-induced deletion of most of X euchro-
origin: X-ray-induced deletion of most of X euchromatin from In(l)sc8.
matin from In(l)sc8. discoverer: Krivshenko and Cooper, 1954.
discoverer: Krivshenko and Cooper, 1954. genetics: Carries wild-type alleles of y, ac, and 66
genetics: Carries wild-type alleles of y, ac, and bb but not sc or pn.
but not sc or pn.
Dp(l;f)12Dl
cytology: 1.8 times the length of chromosome 4 at cytology: 2.2 times the length of chromosome 4 at
metaphase; has nucleolus organizer (Cooper). metaphase; has nucleolus organizer (Cooper).
matin from In(l)acs. matin from In(l)sc8.
discoverer. Krivshenko and Cooper, 1954. discoverer Krivshenko and Cooper, 1954.
genetics: Carries wild-type alleles of y, ac, and 66 genetics: Carries wild-type alleles of y, ac, and 66
but not sc or pn. but not sc, or pn.
Dp(hf)1204
Dp(l;f)im cytology: Nine-tenths the length of chromosome 4 at
cytology: 1.6 times the length of chromosome 4 at metaphase; lacks nucleolus organizer (Cooper).
metaphase; lacks nucleolus organizer (Cooper). origin: X-ray-induced deletion of most of X euchro-
matin of In(l)ac8. discoverer: Krivshenko and Cooper, 1954.
discoverer: Krivshenko and Cooper, 1954. references: Grell, 1964, Genetics 50: 151—66.
references: Grell, 1964, Genetics 50: 151-66. 1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32.
1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32.• genetics: Carries wild-type alleles of y and ac but
genetics: Carries wild-type alleles of y and ac but not sc, su(wa.), dor, pn, su(f), or 66. Causes 18.9
not sc, mtfw*), dor, pn, svt(f), or 6b. Causes 14.2 percent nondisjunction of chromosome 4 when
percent nondisjunction of chromosome 4 when added to normal diploid female genotype.
added to normal diploid female genotype. Dp(1;f)1205
Dp(hf}lW cytology: Seven-tenths the length of chromosome 4
cytology: Less than 0.3 the length of chromosome 4 at metaphase; lacks nucleolus organizer (Cooper).
at metaphase; lacks nucleolus organizer (Cooper). origin: X-ray-induced deletion of most of X euchro-
matin from ln(I}acs. discoverer: Krivshenko and Cooper, 1954.
discoverer: Krivahenko and Cooper, 1954. genetics: Carries wild-type alleles of y but not ac,
references: Grell, 1964, Genetics 50: 151—66. sc, pn, or 66.
1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32. Dp(l;f)12O6
genetics: Carries wild-type atleles of y and ac but cytology: One-half the length of chromosome 4 at
not «c, mtCw"), dot, pn, suff), or 6.6. Has no effect metaphase; lacks nucleolus organizer (Cooper).
on disjunction of cforofaosone 4 when added to origin: X-ray-induced deletion of most of X euchro-
normal diploid female genotype. matin from In(l)mc8.
discoverer: Krivshenko and Cooper, 1954. references: Grell, 1964, Genetics 50: 151-66.
genetics: Carries wild-type alleles of y but not ac, 1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32.
sc, pn, or 66. genetics: Carries wild-type alleles of y, ac, sc,
Dp(l;f)1208 su(wa), and dor but not pn, su(t), or bb. Variegates
cytology: Twice the length of chromosome 4 at for dor in male with no Y chromosome. Causes 11.9
metaphase; has nucleolus organizer (Cooper). percent nondisjunction of chromosome 4 when
origin: X-ray-induced deletion of most of X euchro- added to normal diploid female genotype.
matin from In(l)scs. Dp(l;f)1339
discoverer: Krivshenko and Cooper, 1954. cytology: Dp(l;f)lD-E;20; usually associated with
genetics: Carries wild-type alleles of y, ac, and 66 chromocenter in salivary preparations (Krivshenko);
but not sc or pn. 1.1 times the length of chromosome 4 at metaphase;
Dp(l;f)1209 lacks nucleolus organizer (Cooper).
cytology: 1.9 times the length of chromosome 4 at origin: X-ray-induced deletion of most of X euchro-
metaphase; lacks nucleolus organizer (Cooper). matin.
origin: X-ray-induced deletion of most of X euchro- discoverer Krivshenko and Cooper, 1954.
matin from In(l)scs. references: Grell, 1964, Genetics 50: 151-66.
discoverer: Krivshenko and Cooper, 1954. 1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32.
genetics: Carries wild-type alleles of y, ac, and 66 genetics: Carries wild-type alleles of y, ac, sc, svr,
but not sc or pn. and su(wa) but not dor, pn, su(f), or 66. Causes
Dp(l;f)1328 17.5 percent nondisjunction of chromosome 4 when
cytology: Dp(l;f)2A2~3;20; usually associated with added to normal diploid female genotype.
nucleolus in salivary preparations (Krivshenko);
2.1 times the length of chromosome 4 at metaphase; *Dp(l;f)1341
has nucleolus organizer (Cooper). cytology: Dp(l;f)2C-D;20 (Krivshenko); more than
origin: X-ray-induced deletion of most of X euchro- three times the length of chromosome 4 at meta-
matin. phase; has nucleolus organizer (Cooper).
discoverer: Krivshenko and Cooper, 1954. origin: X-ray-induced deletion of most of X euchro-
references: Grell, 1964, Genetics 50: 151—66. matin.
1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32. discoverer: Krivshenko and Cooper, 1954.
genetics: Carries wild-type alleles of y, ac, sc, svr, genetics: Carries wild-type alleles of y, ac, sc, svr,
su(wa), and 66 but not dor, pn, or su(f). and 66 but not pn.
In(l)sc4LscSR/Dp(l;f)1328 dies. Causes 3.8 per- Dp(l;f)J342
cent nondisjunction of chromosome 4 when added cytology: More than three times the length of chro-
to normal diploid female genotype. mosome 4 at metaphase; has nucleolus organizer
*Dp(l;{)1330 (Cooper).
cytology: Dp(l;f)2B10-ll;20 (Krivshenko); 2.6 times origin: X-ray-induced deletion of most of X euchro-
the length of chromosome 4 at metaphase; has matin.
nucleolus organizer (Cooper), discoverer: Krivshenko and Cooper, 1954.
origin: X-ray-induced deletion of most of X euchro- genetics: Carries wild-type alleles of y, ac, sc, svr,
matin. and 66 but not pn.
discoverer: Krivshenko and Cooper, 1954.
genetics: Carries wild-type alleles of y, ac, sc, svr, Dp(l;f)J343
and 66 but not pn. cytology: Dp(l;i)lF;20; usually associated with
*Dp(l;f)1331 chromocenter in salivary preparations (Krivshenko);
cytology: Dp(l;f)lE-F;20 (Krivshenko); 1.9 times 2.6 times the length of chromosome 4 at metaphase;
the length of chromosome 4 at metaphase; has has nucleolus organizer (Cooper).
nucleolus organizer (Cooper), origin: X-ray-induced deletion of most of X euchro-
matin. discoverer: Krivshenko and Cooper, 1954.
discoverer Krivshenko and Cooper, 1954. references: Grell, 1964, Genetics 50: 151—66.
genetics: Carries wild-type alleles of y, ac, sc, svr, genetics: Carries wild-type alleles of y, ac, sc, svr,
and 66 but not pn. sti(w*), and bb but not dor, pn, or au(f).
Dp(l;f)1337 *Dp(l;f)134S
cytology: Dp(l;f)lF4-2A3; 19-20 (Gersh; left break- cytology: Dp(l;f)lC;20; always associated with
point originally given as 2B8-9 by Krivshenko); nucleolus in salivary preparations (Krivshenko);
usually associated with chromocenter in isalivary 1.7 times the length of chromosome 4 at metaphase;
preparations (Krivshenko); 1.4 times the length of has nucleolus organizer (Cooper).
chromosome 4 at metaphase; lacks nucleolus orga- origin: X-ray-induced deletion of most of X euchro-
ganizer (Cooper). matin.
origin: X-ray-induced deletion of most of X euchro- discoverer: Krivsheoko and Cooper, 1954.
matin. genetics: Carries wild-type alleles of y, ®c, sc, svr,
discoverer: Krivshenko and Cooper, 1954. and 66 bat not pn.
Dp(l;f)1346 DpCl;t)U94
cytology: Dp(l;t)lB12-13;20; usually associated cytology: Dp(l;f)lB10~14;20; usually free of nu-
with nucleolus in salivary preparations cleolus in salivary preparations (Krivshenko); 2.7
(Krivshenko); twice the length of chromosome 4 at times the length of chromosome 4 at metaphase;
metaphase; has nucleolus organizer (Cooper). has nucleolus organizer (Cooper).
matin. matin.
discoverer: Krivshenko and Cooper, 1954. discoverer: Krivshenko and Cooper, 1955.
references: Grell, 1964, Genetics 50: 151-66. genetics: Carries wild-type alleles y, ac, sc, svr,
1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32. and bb but not pn.
genetics: Carries wild-type alleles of y, ac, sc, svr, Dp(l;f)U98
and bb but not su(wa), dor, pn, or su(f). cytology: Dp(l;i)lF;20; usually free of nucleolus in
In(l)sc*Lsc*R/Df(l;i)1346 dies. Causes 8.6 per- salivary preparations (Krivshenko); 3.3 times the
cent nondisjunction of chromosome 4 when added length of chromosome 4 at metaphase; has nu-
to normal diploid female genotype. cleolus organizer (Cooper).
Dp(l;0U79 origin: X-ray-induced deletion of most of X euchro-
cytology: Dp(l;f)lC;20; usually associated with matin.
nucleolus in salivary preparations (Krivshenko); discoverer Krivshenko and Cooper, 1955.
2.1 times the length of chromosome 4 at meta- references: Grell, 1964, Genetics 50: 151—66.
phase; has nucleolus organizer (Cooper). 1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32.
origin: X-ray-induced deletion of most of X euchro- genetics: Carries wild-type alleles of y, ac, sc, svr,
matin. suCw*), and bb but not dor, pn, or su(t). Causes 3.2
discoverer: Krivshenko and Cooper, 1955. percent nondisjunction of chromosome 4 when
genetics: Carries wild-type alleles of y, ac, sc, svr, added to normal diploid female genotype.
and bb but not pn. Dp(l;f)lS01
cytology: Dp(l;f)2A;19E4-Fl; usually free of nu-
Dp(J;{)1488 cleolus in salivary preparations (Krivshenko); 4.4
cytology: Dp(l;f)2A;20; frequently associated with times the length of chromosome 4 at metaphase;
nucleolus in salivary preparations (Krivshenko); has nucleolus organizer (Cooper).
2.5 times the length of chromosome 4 at metaphase; origin: X-ray-induced deletion of most of X euchro-
has nucleolus organizer (Cooper). matin.
origin: X-ray-induced deletion of most of X euchro- discoverer Krivshenko and Cooper, 1956.
matin. genetics: Carries wild-type alleles of y, ac, sc, svr,
discoverer: Krivshenko and Cooper, 1955. and bb but not pn.
references: Grell, 1964, Genetics 50: 151-66. Dp(l;f)1512
1964, Proc. Natl. Acad. Sci. U.S. 52: 226-32. cytology. Dp(l;f)lF;19E4-Fl; usually free of nu-
genetics: Carries wild-type alleles of y, ac, sc, svr, cleolus in salivary preparations (Krivshenko); 3.6
au(wa), and bb but not dor, pn, or su(f). Causes times the length of chromosome 4 at metaphase;
3.8 percent nondisjunction of chromosome 4 when has nucleolus organizer (Cooper).
added to normal diploid female genotype. origin: X-ray-induced deletion of most of X euchro-
matin.
*Dp(l;f)1489 discoverer: Krivshenko and Cooper, 1956.
cytology: Dp(l;f)lD;20; usually associated with genetics: Carries wild-type alleles of y, ac, sc, svr,
nucleolus in salivary preparations (Krivshenko); and bb but not pn.
1.8 times the length of chromosome 4 at metaphase. Dp(l;f)1513
Lacks nucleolus organizer (Cooper). cytology: Dp(l;f)lB10-14;20; usually associated
origin: X-ray-induced deletion of most of X euchro- with nucleolus in salivary preparations
matin. (Krivshenko); more than twice the length of chro-
discoverer: Krivshenko and Cooper, 1955. mosome 4 at metaphase; has nucleolus organizer
genetics: Carries wild-type alleles of y, ac, ac, svr, (Cooper).
and bb but not pn. origin; X-ray-induced deletion of most of X euchro-
Dp(l;f)1492 matin.
cytology. Dp(I;f)lB10-12;20; always associated with discoverer: Krivshenko and Cooper, 1956.
nocleolus in salivary preparations (Krivshenko); genetics: Carries wild-type alleles of y, ac, sc, svr,
1.9 tines the length of chromosome 4 at meta- and bb but not pn.
phase; lacks nucleolus organizer (Cooper). Dp(l;f)15U
origin: X-ray-induced deletion of most of X euchro- cytology. Dp(l;f)lB12~13;20; usually associated
auttin. with the nucleolus in salivary preparations
discoverer: Krivshenko and Cooper, 1955. (Krivshenko); 1.9 times the length of chromosome 4
genetics: Carries wild-type alteles of y, ac, ®c, avr, at metaphase; has nucleolus organizer (Cooper).
and fob bat not pn. origin: X-ray-induced deletion of most of X euchro-
dies. matin.
discoverer: Krivshenko and Cooper, 1956. *Dp(l;f)R37

genetics: Carries wild-type alleles of y, ac, sc, svr, cytology: Dp(l;f)lA3-4;16F2-3;19F-20Al.
and bb but not pn. new order: |l6F3 - 20-2OF |.
Dp(l;f)1518 origin: X-ray-induced deletion of most of euchro-
cytology: Dp(l;f)2A4-Bl;20; usually free of nu- matin from R(l)2.
cleolus in salivary preparations (Krivshenko); 3.9 discoverer Pontecorvo.
times the length of chromosome 4 at metaphase; synonym: Del(Xc2)37.
has nucleolus organizer (Cooper), references: Slizynska, 1942, DIS 16: 67.
origin: X-ray-induced deletion of most of X euchro- *Dp(l;f)R38
matin. cytology: Dp(l;f)lA3-4;lF;20.
discoverer: Krivshenko and Cooper, 1956. new order: J1A4 - 1F|2O.2OF - 20Al|.
genetics: Carries wild-type alleles of y, ac, sc, svr, origin: X-ray-induced deletion of most of euchro-
and bb but not pn. matin from R(l)2.
*Dp(l;f)Al: Duplication(l;free) from Austin discoverer Pontecorvo, 1942.
origin: Aneuploid segregant from T(1;3)A1/+. synonym: Del(Xc2)38.
*Dp(l;f)A12 references: 1942, DIS 16: 65.
cytology: Dp(l;f)lB-C;13Bl-S. *Dp(l;f)R4Q
origin: Aneuploid segregant from T(1;2;4)A12/+. cytology: Dp(l;f)lA3-4;lF4-5;2Q.
*Dp(l;f)eq: Duplication(T ;free) from eqvational new order: |lA4 - 1F4|2O-2OF - 20Al|.
producer origin: X-ray-induced deletion of most of euchro-
origin: X-ray-induced deletion of most of euchromatin from R(l)2.
matin from X chromosome carrying eq. discoverer: Pontecorvo.
discoverer: Schultz, 34k4. synonym: Del(Xc2)40.
genetics: Contains y + to pn + , inclusive, and bb+. references: Slizynska, 1942, DIS 16: 67.
Male fertile but rather inviable; has occipital
bristles; eyes rough, wings spread, wing veins *Dp(l;f)R41
thickened. Female has occipital bristles; wings cytology: Dp(l; 1)1A3-4;1F4-5;2O.
straight edged and coarse textured. Female with new order: |lA4 - 1F4|2O-2OF - 20Al|.
two duplications occasionally survives and shows origin: X-ray-induced deletion of most of euchro-
extreme spread wings and rough eyes. matin from R(l)2.
Dp(l;f)R: Duplication(l;free) from Ring X discoverer Pontecorvo.
cytology: Dp(l;f)lA3-4;3A;l9F-20A1. synonym: Del(X<;2)41.
new order: |lA4 - 3A|20-20F - 20Al|. references: Slizynska, 1942, DIS 16: 67.
origin: Spontaneous deletion of most of euchromatin *Dp(l;f)R42
from R(l)2. cytology: Dp(l;i)lA3-4;2A2-3;20.
discoverer Schultz, 35dlO. new order. |lA4 - 2A2|20-20F - 20Al|.
synonym: Dp(l;f)X^, origin: X-ray-induced deletion of most of euchro-
genetics: Covers y to kz but not bb. Variegation for matin from R(l)2.
dor, ac, svr, pn, and kz; decreased as Y's are discoverer Pontecorvo.
added. Variegation of y insensitive to F's. synonym: Del(Xc^)42.
*Dp(l;f)Rl references: Slizynska, 1942, DIS 16: 67.
origin: X-ray-induced deletion of most of euchro- *Dp(l;f)R43
matin from R(l)2. cytology: Dp(l;f)lA3-4;lF4-5;20.
discoverer: Pontecorvo. new order: |lA4 - 1F4|2O.2OF - 20Al|.
synonym: Del(Xc*)l. origin: X-ray-induced deletion of most of euchro-
references: 1942, DIS 16: 65. matin from R(l)2.
*Dp(l;f)R35 discoverer Pontecorvo.
cyto Iogy: Dp(l;t)lA3-4;l 7A4-S;19F-2OA1. synonym: D&l(Xc^)43.
new order. |l7A5 - 20-20F|. references: Slizynska, 1942, DIS 16: 67.
origin: X-ray-induced deletion of most of euchro-
matin from R(l)2. *Dp(l;f)R53d
discoverer: Pontecorvo. cytology: Dp(l;f)lA3-4;lF~2A;20.
synonym: Del(X^)35. new order: |lA4 - 1 F [ 2 ( K 2 0 F - 2QAl|.
references: Slizynska, 1942, DIS 16: 67. origin: X-ray-induced deletion of most of euchro-
*Dp(hW36 matin from R(l)2.
cytology: Dp(l;i)lA3-4;17A4-5;l9F-20Al. discoverer: S. Brown, 1953.
new order. |l7A5 - 20-2OF |, synonym: Del(Xc2)53d.
origin: X-ray-induced deletion of most of euchro- references: 1955, DIS 29: 70.
matin from R(l)2, Brosseau, 1955, DIS 29: 106.
di scoverer Pontecorvo. genetics: Contains wild-type allele® of y, ac, me,
synonym: and mu(®%- covers Dl(l)26®-1, Female tolerates two
reference.: Slizynska, 1942, DIS 16: 67. duplications; male tolerates only one. Fly heal- or
homozygous for y and the duplication shows mo- Dp(2;2)41A

saicism for y. There is probably both variegation cytology: Tandem duplication for material in 41 A.
for y and loss of the duplication. origin: Spontaneous in the In(2L)Cy + In(2R)Cy
*Dp(l;f)RA chromosome of a balanced In(2L)Cy +In(2R)Cy/
cytology: Dp(l;f)lA3-4;lF-2A;20 (Slizynska). M(2)S210 stock.
new order: |lA4 - 1F|2O-2OF - 21A11. discoverer: Schultz, 1945.
origin: X-ray-induced deletion of most of euchro- genetics: Acts as a suppressor of M(2)S2, and
matin from R(l)2. perhaps as a partial suppressor of L. Fly hetero-
discoverer: Pontecorvo. zygous for the duplication appears more stocky
references: 1942, DIS 16: 65. than normal.
*Dp(l;0sc260-27; Duplicathn(l;free) scute *Dp(2;2)bw*i-CyR
cytology: Dp(l;f)lA8-Bl;19F. cytology: Dp(2;2)58A4-Bl;59D.
origin: Aneuploid segregant from T(l;2)sc26°-27/+. origin: Associated with In(2R)bwALCyR.
*Dpd;f)w">3: Duplication(l;free) white-mottled DP(2;2)bwV34kLCyL
cytology: Dp(l;f)3C-D;19-20; breakpoints inferred cytology: Dp(2;2)41;42A2-3.
from genetic data, origin: Aneuploid recombinant from
discoverer Muller, 1925.
references: 1930, J. Genet. 22: 299-334. cytology: Dp(2;2)58A4-Bl;59E2-4.
genetics: w/Dp(l;f)wm3 male has variegated eyes origin: Associated with In(2R)bwVDeiLcyR.
and is sterile; C(1)RM, w/Dp(l;f)w'n3 female has
variegated eyes and is fertile. cytology: Dp(2;2)58A4-Bl;59D6-El.
Xc*: see Dp(l;f)R origin: Associated with In(2R)bwVDe2LcyR.
Dp(2;2)C619
*Dp(l;f)y-sc: Duplication(l;free) for yellow and cytology: Dp(2;2)26A;28E.
scute new order: 21 - 28E|26A — 60.
origin: X-ray-induced deletion of most of X euchro- origin: X ray induced in oocyte.
matin. discoverer: Roberts and Thomas, 1965.
discoverer: Oliver, 32k21. references: Roberts, 1966, Genetics 54: 969-79.
references: 1937, DIS 7: 19. Thomas and Roberts, 1966, Genetics 53: 855—62.
phenotype: Carries wild-type alleles of y and sc but genetics: Homozygous viable. Reduces recombina-
not pn. tion in 2L. Map distance between al and pr reduced
Dp(l;f)z9: Duplication(l;free) zeste from 44.2 to 7.3 in Dp(2;2)C619/+ and to 17.0 in
cytology. Dp(l;f)3E7-Fl; 19-20. Dp(2;2)C619 homozygotes.
origin: X-ray-induced deletion of most of euchro-
matin from z-bearing X chromosome. cytology: Dp(2;2)41B2-Cl;42A2-3.
discoverer Gans. origin: Associated with In(2R)CyLbv?VD°lR.
references: 1953, Bull. Biol. France Belg., Suppl. Dp(2;2)Cyl-bwVD*2*
38: 1-90 (fig.). cytology: Dp(2;2)41A-B;42A2-3.
genetics: Contains z and wild-type alleles of y origin: Associated with In(2R)Cy^bwVD°2R.
through dm. Dp(2;2)Cy*bwV34kR
cytology: Dp(2;2)58A4-Bl;59.
Dp(2;J)C239: Duplication(2;l) Crossover origin: Aneuploid recombinant from
suppressor ln(2R)bwV3** + Cy/+.
cytology: Dp(2;l)7A-B;36C;39E. Dp(2;2)Px*: Dp(2;2) Plexate
origin: Aneuploid segregant from T(l,;2)C239/+. cytology: Dp(2;2)60B;60Dl-2.
Dp(2;l)OR19: DupHcation(2;l) from Oak Ridge origin: Associated with In(2LR)S56f.
cytology: Dp(2;l)20;48E;51F;S7C.
origin: Aneuploid segregant from T(1;2)OR19/+. Dp(2;2)S: Duplication(2;2) Star
cytology: Dp(2;2)21D2-3;21E2-3; tandem repeat.
Dp(2;Y)bw+: Duplication(2;Y) brown-wild type new order: 21A - 21E2|21D3 - 60.
cytology: Dp(2;Y)YL;$8Fl-59A2;6QE3-Fl. origin: Spontaneous as an asymmetrical crossover.
origin: Aneuploid segregant from T(Y;2)bw*Y/+. discoverer: E. B. Lewis, 39i.
Dp&Y)G references: 1941, Proc. Na'l. Acad. Sci. 27: 31-35.
ey tol ogy: Dp(2; Y)36B4-S;40F. 1945, Genetics 30: 137-66.
origin: Aneuploid segregant from T(Y;2)G/+. genetics: Duplicated segment contains the loci of S
*Dp(2;Y)H and ast. ast mutant in both members of the dupli-
cytology: Dp(2;Y)37Bl-2;4QB2-3. cation (+ ast + ast). Duplication appears wild type
origin: Aneaploki segregant from T(Y;2)H/+. when homozygous or when heterozygous with mat.
*Dp(2;Y)R24 Heterozygous with S, it has normal or only slightly
cytology. Dp(2;Y)45A;51E. roughened eyes. Various combinations of S and ast
origin: Aneuploid segregpnt from T(Y;2)R24. alleles have been introduced into the duplication.
CHROMOSOME ABERRATIONS - INVERSIONS 321
Through unequal crossing over, a triplication and a

quintuplication of the region have been synthesized. INVERSIONS
Dp(2;2)S56f
cytology: Dp(2;2)21C6-Dl;22A3-Bl. ln(l)65: Inversion(l) 65
origin: Associated with In(2LR)Px4. cytology: In(l)lC;10B.
Dp(2;2)SM5 origin: X ray induced simultaneously with T(l;3)65
cytology: Dp(2;2)42A2-3}42D;53C;58A4-Bl;58F. in y.
origin: Associated with In(2LR)SM5. discoverer: Lindsley, Edington, and Von Halle.
Dp(2;3)C328: Duplication(2;3) Crossover genetics: Inseparable from y. About 1 percent non-
suppressor disjunction*and 21.8 percent recombination in
cytology: Dp(2;3)55C;58B;80. In(l)65/+ female; 25.9 percent nondisjunction and
origin: Aneuploid segregant from T(2;3)C328/+. 19.7 percent recombination in Jn(l)65/+/Y female
Dp(2;3)dp: Duplication(2;3) dumpy (Grell, 1962, Genetics 47: 1737-54).
cytology: Dp(2;3)34D;41A;80;81. tn(l)94-2A
origin: Aneuploid segregant from T(2;3)dp/+. cytology: In(l)lF-2A;5E-6A (Lindsley).
Dp(2;3)P: Duplication(2;3) Pale origin: Derived by recombination from C(1)94-2A.
cytology: Dp(2;3)58E3-F2;60D14-E2;96B5-Cl. discoverer: Rosenfeld.
origin: Aneuploid segregant from T(2;3)P/+. genetics: Leads to partial stabilization of tandem
Dp(2;3)P32: Duplication(2;3) from Pasadena ring compound X chromosome. Recoverable in de-
cytology: Dp(2;3)41A;42D-E;44C-D;89D7-El. rivative single ring, R(l)9-4. Exists in three inter-
origin: Aneuploid segregant from T(2;3)P32/+. changeable configurations in the C(1)TR (e.g.,
Novitski and Braver, 1954, Genetics 39: 197—209).
Dp(2;f) 1: Duplication(2; free)
cytology: Dp(2;f)21;41 (left breakpoint tentative) *tn(l)272-13
superimposed on ln(2LR)40F;59E. cytology: In(l)lA6-Bl;llA7-8;llF2-12Al;18A4-Bl.
new order: 21J41 - 40F|S9E - 60. new order: 1A1 - 1A6J12A1 - 18A4|llA7 -
Tentative, 1B1|11A8 - 11F2|18B1 - 20.
origin: X-ray-induced derivative of In(2LR)bwV326; origin: X ray induced.
possibly a deletion of most of the long arm. discoverer Demerec, 1940.
discoverer: E. H. Grell, 1959. references: Sutton, 1943, Genetics 28: 213.
genetics: Carries normal alleles of or, sp, bs, Pin, genetics: Mutant for sc and 1(1)272-13 but not ac or
and Px, but not of al or px. SVT. Male lethal.
*ln(l)303-1
Dp(3;J)N264-6: Duplication(3;l) Notch cytology: In(l)2B13-15;7Bl-3,9Dl-3.
cytology- Dp(3;l)3C9-Dl;73E;80C. new order. 1 - 2B13|9D1 - 7B3|2B15 - 7Bl|9D3 -
origin: Aneuploid segregant from T(l;3)N264-6/+. 20.
Dp(3;l)O5: Duplication(3;l) of Oliver origin: X ray induced.
cytology: Dp(3;l)4F2-3;88A-C;92C-D. discoverer Demerec.
origin: Aneuploid segregant from T(1;3)O5/+. synonym: Tp(l)303-l.
Dp(3;1)ry35: Duplication(3;l) rosy references: Hoover, 1938, Z. Induktive
cytology: Dp(3;l)20;87C-E;91B-C. Abstammungs- Vererbungslehre 74: 420—34 (fig.).
origin: Aneuploid segregant from T(l;3)ry3S/+. genetics: Nearly lethal.
Dp(3;3)bxdlOO: Duplication(3;3) bithoraxoid ln(T)481

cytology: Dp(3;3)66C;89B5-6;89E2-3. cytology: In(l)12E-F;14B.
origin: Aneuploid recombinant product from origin: X ray induced simultaneously with
Tp(3)bxdl00/+,
Dp(3;3)bxd"0 discoverer. Lindsley, Edington, and Von Halle,
cytology: Dp(3;3)89E2-3;91C7-Dl;92A2-3. references: 1960, Genetics 45: 1649—70.
origin: Recombinant product from Tp(3)bxdllo/+. ln(J)A99b: lnversion(1) from Austin
cytology: ln(l)W3-El;19D-E.
Dp(3;4)ry + : Duplication(3;4) rosy-wild type origin: X ray induced.
cytology: Dp(3;4)86D2-3;88B;101A-D;101 F. discoverer: Stone,
new order: 88B - 86D3|l01F - 101D; genetics: Viability, fertility, and egg hatch good.
Tentative. H1)AB
origin: X-ray-induced derivative of the 3RD4P ele- cytology: In{l)9F;13Fl-10.
ment of T(3;4)86I> = T(3;4)86D2-3;101F. discoverer: Bodeman.
discoverer E. H. Grell, 1960. references: Stone and Thomas, 1935, Genetica 17:
references: 1962, Z. Vererbungslehre 93: 371-77. 170-84.
genetics: Carries normal alleles of cu, kmr, and ry. genetics: Primary nondisjunction 0.5 percent, sec-
Shows tendency toward somatic elimination. ondary 29.3 percent; recombination 18.2 percent in
322 GENETIC VARIATIONS OF DROSOPHILA MELAHOGASTER
In(l)AB/+ and 26.3 in In(l)AB/+/Y female (Grell, genetics: Reversal of BlBl to wild type; un, vb, t,
1962, Genetics 47: 1737-54). Stone and Thomas lh, and os not affected. Male lethal.
(1935) obtained 14.3 percent recombination in
cytology: In(l)16A2-4;20A2-3.
ln(l)ac3: Inversion(l) achaete origin: X ray induced.
cytology: In(l)lB2-3;lB14-Cl. discoverer: Demerec, 38d.
origin: X ray induced. references: Sutton, 1943, Genetics 28: 97—107.
discoverer: Dubinin, 1929. genetics: Position effect at B.
synonym: In(l)sc10.
references: 1930, Zh. Eksperim. Biol. 6: 300-24. ln(l)BM ': Inversion(l) Bar of duller
1932, J. Genet. 25: 163-81. cytology: In(l)16A2-5;20A3-B (Sutton, 1943,
1933, J. Genet. 27: 447. Genetics 28: 97-107).
genetics: Associated with ac3. origin: X ray induced.
discoverer: Muller, 34e.
cytology: In(l)8C17-Dl;16E2-3 (Hoover). references: 1935, DIS 3: 29.
discoverer: Mackensen. genetics: Position effect at B. Primary nondisjunc-
references: Stone and Thomas, 1935, Genetica 17: tion 0.4 and secondary 18.5 percent; recombination
170-84. 32 percent in In(l)BM1/+ and 35.4 percent in
Hoover, 1938, Z. Inductive Abstammungs- In(l)BM1/+/Y female (Grell, 1962, Genetics 47:
Vererbungslehre 74: 420-34 (fig.). 1737-54).
genetics: Homozygous female sterile and therefore ln(l)B>*2
used as an X chromosome balancer. Inversion de- cytology: In(l)16A2-5;20E (Sutton, 1943, Genetics
parts slightly from wild-type phenotype in that 28: 97-107).
eyes are rounded and slightly bulging. Total re- origin: X ray induced.
combination 3.8 percent in In(l)AM/+ (Stone and discoverer: Muller, 34e.
Thomas, 1935). references: 1935, DIS 3: 29.
ln(J)At: Inversion(l) Attenuated genetics: Position effect at B.
cytology: In(l)16A4-5;18C4-6;20A2-3 superimposed
on In(l)lB3-4;20B-Dl*<lB2-3;20B-DlR + In(l)4D7- *ln(l)Br*y-2: Inversion(l) Bar-reversed
El;llF2-4. cytology: In(l)3F8-4Al;16A2-4; right break in right
new order: 1A - 1B3|2OB - 20A3|l6A5 - section of Dp(l;l)B =Dp(l;l)15F9-16Al;16A7-Bl.
18C4J20A2 - 18C6|l6A4 - 11F4|4E1 - new order: 1 - 3F8|l6A2 - 16Al|l6A7 -
11F2|4D7 - 1B3J20D1 - 20F. 4Al|l6A4 - 2 0 .
origin: X ray induced in In(l)scSiLsc8R+dl-49. origin: X ray induced in Dp(l;l)B.
discoverer: Valencia and Valencia, 1949. discoverer: Bishop, 1940.
synonym: Tp(l)At. references: Sutton, 1943, Genetics 28: 100.
references: 1949, DIS 23: 64. genetics: Reversal of B.
genetics: Associated with At. Male and homozygous
female viable and fertile. cytology: ln(l)15F9-16Al;16A7-Bl;20AS-Bl; right
break between segments of Dp(l;l)B =Dp(l;l)15F9-
*ln(VB263-S: Inversion(l) Bar 16A1;16A7-B1.
cytology: In(l)15F9-16Al;16A7-Bl;17A3-4, Left new order: 1 - 16A7J20A5 - 16Al|20Bl - 20F.
break occurs between repeated regions associated origin: X ray induced in B.
with Dp(l;l)B=Dp(l;l)15F9-16Al;16A7-Bl discoverer: Bishop, 1940.
(Kaufmann and Sutton). references: Sutton, 1943, Genetics 28: 100.
new order: 1 - 16A7|l7A3 - 16A1J17A4 - 20. genetics: B reversion.
origin: X ray induced in B.
discoverer: Demerec, 33k. ln(1)bbDf: Inversion(l) bobbed-Deficiency
references: Sutton, 1943, Genetics 28: 97—107. cytology: ln(l)4D2-3;20B-C;20C-D; deficient for
genetics: B reversed; lethal; un, vb, t, lh, and os° 20C-D; two-thirds normal length at metaphase.
not affected. new order: 1 - 4D2|20B - 4D3|20D - 20F.
*ln(1)B263.24 origin: X ray induced.
cytology: In(l)10C2-Dl;12D2-El;15F9~16Al;16A7-Bl; discoverer: Sivertzev-Dobzhinsky and Dobzhansky,
right breakpoint between first and second segments 31b.
of BlB' triplication, references: 1933, Genetics 18: 173-92.
new order: 1 - 1OC2J16A7 - 12E1J10D1 - Sturtevant and Beadle, 1936, Genetics 21: 554-
12D2|l6Al - 16A7|l6Al - 20. 604.
origin: X ray induced in Dp(l;l)BiBi = Dp(l;l)15F9- genetics: Right breakpoint between tb and rg. De-
16A1;16A7-B1. ficient for bb. In(l)bbDf/+ female produces about
discoverer: Demerec, 34a. 2 percent exceptional sons from four-strand double
synonym: Tp(l)B363-24, exchange within inverted segment. Secondary ex-
references: Sutton, 1943, Genetics 28: 97-107. ceptions about 13 percent.
ln(l)bbDfLCI*: Inversion(l) bobbed-Deficiency Sturtevant and Beadle, 1936, Genetics 21: 554—
Left Cl-Right 604.
cytology: In(l)4D2-3;20B-CL4A5-Bl;17A6-BlR; du- genetics: Left break between ec and bi; right break
plicated for 4B1-D2 and 17B1-20B. between os and fu; 1(1)C associated with left break
origin: Recombinant containing left end of In(l)bbDi (Muller). About 0.35 percent primary and 37 per-
and right end of In(l)Cl. cent secondary exceptions. Total recombination
references: Sturtevant and Beadle, 1936, Genetics about 1 percent.
21: 554-604. other information: In(l)Cl, sc 1(1)C & v si B is the
genetics: Duplicated for bi, rb, fu, and car but not C1B chromosome, described in the section on bal-
ec, rg, f, os, or 66. Survives as small male with ancers.
less convex outer wing margins than normal and ln(l)CH-bbDfR: Inversion(l) CI-Left bobbed-
usually one or more notches at wing tips; sterile, Deficiency Right
with collapsed testes. Heterozygous female fertile, cytology: In(l)4A5-Bl;17A6-B1^4D2-3;20C-DR; de-
with slightly narrowed wings. ficient for 4B1-D2 and 17B1-20C.
ln(l)bbt>fLY4R: Inversion(l) bobbed-Deficiency origin: Recombinant containing left end of In(l)Cl
Left yellow-4 Right and right end of In(l)bbDt.
cytology: ln(l)4D2-3;20B-CLlA8-Bl;18A3-4R; du- references: Sturtevant and Beadle, 1936, Genetics
plicated for 1B1-4D2 and 18A4-20B. 21: 554-604.
origin: Recombinant containing left end of/nCIJ&fr0* genetics: Deficient for bi, rb, fu, car, and 66 but not
and right end of In(l)y4. ec, tg, f, or os. Both bi and fu lethal when hetero-
references: Sturtevant and Beadle, 1936, Genetics zygous for/n(7j)C/I'66z>'-R. Heterozygous female
21: 554-604. extreme Minute \M(1)4BC +M(l)n], with abnormal
genetics: Duplicated for ac through rb and fu through wing shape; ovaries normal but female sterile.
car. Heterozygous female has stubby posterior ver- Male lethal.
ticals and disarranged scute liars; outer wing
margin less convex than normal; fair viability and ln(\)Cll-y4R: Inversion(T) Cl-Left yellow-4 Right
fertility. Enhances expression of heterozygous B. cytology: In(l)4A5-Bl;17A6-Bl^lA8-Bl;18A3-4R;
Male lethal. duplicated for 1B1-4A5, deficient for 17B1-18A3.
origin: Recombinant containing left end otln(l)Cl
*ln(1)Br: Inversion(T) Bridged and right end of In(l)y4.
origin: X ray induced. references: Sturtevant and Beadle, 1936, Genetics
discoverer: Muller, 2713. 21: 554-604.
references: 1935, DIS 3: 29. genetics: Duplicated for ac through ec; deficient for
genetics: Associated with dominant mutant, Br. fu but not /, v6, os, or car. Heterozygous female
Crossing over suppressed to right of v, about has irregular acrostichal rows and wings smaller
normal to left. and with less-convex posterior margin than normal.
*ln(l)C18: Inversion(l) Crossover suppressor Enhances expression of heterozygous B. Male
cytology: In(l)3F;17Al-6. lethal.
origin: X ray induced. *ln0)cf3a2: Inversion(l) cut
discoverer: Roberts, 1964. cytology: In(l)7B2-Cl; 19-20.
genetics: Eliminates sc-f recombination. Male origin: X ray induced.
viable. discoverer: Hannah, 47g.
ln(1)CU6 genetics: cr affected but not cm, sn, or oc. Male
cytology: In(l)lF;14A, lethal.
origin: X ray induced. *ln(1)ct3bl
discoverer: Roberts, 1965. cytology: Jn(l)3A4-Bl;7B2-5. May be a duplication
genetics: Eliminates sc-/recombination. Male for 3A3-4 or insertion of material from another
fertile. chromosome.
ln(l)C206 origin: X ray induced,
cytology: In(I)8F;llA;16A. discoverer: Hannah, 47g.
new order: 1 - 8FJ16A - 11A|8F - HA|l6A - 20. genetics: cr affected but not pa, w, cm, m, or oc.
origin: X ray induced. Male lethal.
discoverer. Roberts, 1965.
genetics: Eleven percent recombination between sc cytology: In(l)4E2-3;7B2-4 (Hannah).
and f. Male lethal. origin: X ray induced,
ln(l)CI: Inversion(l) Cl discoverer: De Frank, 47g.
cytology. In(l)4A5-Bl;17A6-Bl (Hoover, 1938, Z. genetics: ct affected but not r6, ex, rg, cm, , or
Induktive Abstatmmmgs- Vererbuagslehre 74: 429). oc. Male lethal.
origin: Spontaneous in a sc t3 v si B chromosome.
discoverer: Muller, 20j. cytol ogy: In(l)7B2-3;l 9-20.
references: 1928, Genetics 13: 279—357. origin: X my induced.
Gershenson, 1935, J. Genet. 30: 115—25. discoverer: Hannah, 47g.
genetics: ct affected but not cm, sn, or oc. Male ln(l)Dl: Inversion(l) from deoxycytidine
lethal. cytology: In(l)13B;16A.
origin: Induced by tritiated deoxycytidine.
cytology: ln(l)7B2-3;20; position of heterochromatic discoverer: Kaplan, 1965.
breakpoint with respect to centromere unknown. references: 1966, DIS 41: 59.
origin: X ray induced in R(l)2. genetics: Male lethal.
discoverer Hannah, 14a3. ln(l)dl-49: Inversion(l) delta-49
genetics: ct affected but not y, ac, sc, cm, sn, or cytology: In(l)4D7-El;llF2-4 [Painter; Hoover,
oc. Male lethal. 1938, Z. Induktive Abstammungs- Vererbungslehre
74: 420-34 (fig.)].
cytology: In(l)3D2-5;7B2-4. discoverer: Muller, 26k.
origin: X ray induced, references: Muller and Stone, 1930, Anat. Record
discoverer: Hannah, 47g. 47: 393-94.
genetics: ct affected but not an, sn, or oc. Male Stone and Thomas, 1935, Genetica 17: 170-84.
lethal. Sturtevant and Beadle, 1936, Genetics 21: 554-
604.
cytology: In(l)4Bl-4;7B4-Cl + In(l)10D5-6;20B-C. genetics: Left break between rb and cv; right be-
origin: X ray induced. tween tw and g. Measures of recombination vary
discoverer: Muller, Valencia, and Valencia, 1946-53. from 5.5 percent (Grell, 1962, Genetics 47: 1737-
references: Valencia, 1966, DIS 41: 58. 54) to about 15 percent (Sturtevant and Beadle,
genetics: Associated with ct43aH1. 1936); secondary exceptions from 33 percent (Grell,
*ln(l)ct268-13 1962) to 44 percent (Sturtevant and Beadle, 1936).
cytology: In(l)2E3-Fl;2F2-3;7B2-3;7B4-5;l9A4- other information: Used as a balancer either with
5;19A6-B1 superimposed on R(1)1A3-4;19F-2OA1. markers y Hw m2 g4 or y w lza with Hw and lz8
2F1-2, 7B3-4, and 19A5-6 missing. sterilizing homozygous female.
new order: |lA4 - 2E3|7B2 - 2F3|l9A4 -
7B5|19B1 - 20-20F - 20Al|. *ln(1)drp: Inversion(l) droop wings
origin: X ray induced in R(l)2. cytology: In(l)12B;20B.
discoverer: Demerec, 34f. origin: Spontaneous from hi.
references: Hoover, 1937, Genetics 22: 634—40 discoverer Ives, 48f.
(fig-). synonym: In(l)hil; Inversion(l) droop.
1938, Z. Induktive Abstammungs- Vererbungslehre references: 1949, DIS 23: 58.
74: 420-34 (fig.). genetics: Associated with mutant droop wings. Male
genetics: Deficient for ct but not scp or sn. Male viable.
lethal. *ln(l)dta: Inversion(l) delta wing
cytology: In(l)6B2-3;15E7-F2.
cytology: In(l)7B2-3;7B4-5;llD8-9; 7B3-4 missing. origin: Induced by triethylenemelamine (CB. 1246).
new order. 1A - 7B2|llD8 - 7B5J11D9 - 20. discoverer: Fahmy, 1953.
discoverer: Hoover, 1938. genetics: Associated with dta. Female sterile.
references: 1938, Z. Induktive Abstammungs- ln(1)e(bx): Inversion(l) enhancer of bithorax
Vererbungslehre 74: 420-34 (fig.), cytology: In(l)3A;4F.
genetics: Deficient for cr but not scp or sn. Male origin: Gamma ray induced.
lethal. discoverer: £. B. Lewis, 53b.
cytology: In(l)6Fll-7Al;7B5-6;10Bll-12. 7A1-B5 genetics: Associated with e(bx).
missing. h(l)EH: Inversion(l) Entire
new order: 1 - 6Fll|lOBll - 7B6|lOB12 - 20. cytology: In(l)lA;20;20B-C. At prophase, distal end
origin: X ray induced, carries a single heterochromatic segment about
discoverer: Demerec, 35g. equal in size to chromosome 4; proximally it
references: Hoover, 1938, Z. Induktive Abstam- carries a very short heterochromatic segment and
mungs- Vererbungslehre 74: 420—34 (fig.). as a second arm two larger heterochromatic seg-
genetics: Deficient for ct but not cm, scp, or sn. ments (Lindsley).
Male lethal. new order: 20 - 1A|2OC - 20F-20.
Tentative,
cytology: In(l)3D6'El;7B3-5. origin: Spontaneous opening out of R(l)l, y.
origin: X ray induced. discoverer Novitski.
discoverer. Hoover, 35j. references: 1949, DIS 23: 94-95.
references: 1938, Z. Induktive Abstammungs- Lindsley, 1958, Z. Vererbungslehre 89: 103-22.
Vererbungslehre 74: 420-34 (fig.). genetics: Entire chromosome, including l(lXfl+, and
genetics: ct affected but not cm, scp, or sn. Male y, inverted. Carries mutant alleles of 66 at each
lethal. end, which acting together produce 66+phenotype.
ln(l)EN2 ln(l)FM6
cytology: In(l)lA3-4;19F-20Al;20. Inferred from cytology: In(l)15D-E;20A-B superimposed on
origin, ln(l)lB2-3;20B-Dl + In(l)3C;4E-F + In(l)4D7-
new order: 20 - 1A3|2OA1 - 2OF«2O. El;llF2-4.
Tentative. new order: 1A - 1 B 2 | 2 0 B | 1 5 E - 20A|l5D -
origin: Spontaneous opening of R(l)2, y + . 11F4|4E|3C - 4D7|llF2 - 4F|3C - 1B3|2OD1 -
discoverer: Muller. 20F.
references: 1956, DIS 30: 140-41. origin: X ray induced in In(l)FM4, y3ld sc8 dm B.
genetics: Entire chromosome inverted like In(l)EN discoverer: R. F. Grell, 55i.
but carries y +rather than y. references: Grell and Lewis, 1956, DIS 30: 70.
*ln(l)EN2B: Inversion(l) Entire 2 of Bender genetics: Male viable and fertile. Female sterile
cytology: In(l)lA3-4;19F~20Al;20. Inferred from owing to dm.
origin, other information: Used as first chromosome bal-
new order: 20 - 1A3|2OA1 - 20F-20. ancer, described as FM6 in the section on bal-
Tentative. ancers.
origin: Spontaneous opening of R(l)2, y v. ln(l)g17Ba6: Inversion(l) garnet
discoverer: M- A Bender, 55e6. cytology: In(l)12B14-15;19F superimposed on
references: 1955, DIS 29: 69. In(l)lB3-4;20B-DlLlB2-3;20B-DlK + In(l)4D7-
*ln(J)exr: Inversion(l) extra venation El;llF2-4.
cytology: In(l)12E8-10;15Dl-3. new order: 1A - 1B3|2OB - 19F|l2B15 -
origin: Induced by triethylenemelamine (CB. 1246). 19F|l2B14 - 11F4|4E1 - 11F2|4D7 - 1B3|2OD1 -
discoverer: Fahmy, 1952. 20F.
references: 1958, DIS 32: 70. origin: X ray induced in In(l)scsi LscSR+ dl-49.
genetics: Affects exr. discoverer: Muller, Valencia, and Valencia, 1946-53.
In(1)f257.4: Inversion(l) forked references: Valencia, 1966, DIS 41: 58.
cytology: In(l)15F2-16Al;16D2-El. genetics: Associated with gl7B»6,
origin: X ray induced, ln(l)gw: Inversion(l) garnet-wild
discoverer: Demerec, 33j. cytology: Breakpoints unknown.
genetics: ( affected. origin: X ray induced.
discoverer: Muller.
ln(l)FM3: Inversion(l) First Multiple references: 1946, DIS 20: 67.
cytology: In(l)3E-F;16A-B;19F-20B; superimposed Chovnick, 1958, DIS 32: 88.
on In(l)lB2-3;20B-Dl + In(l)4D7-El;llF2-4. 1961, Genetics 46: 493-507 (fig.).
new order: 1A - lB2|20B|l6B - 19F|3F - genetics: Associated with gw.
4D7|llF2 - 4 E l | l l F 4 - 16A|3E - 1B3|2OD1 - ln(1)gx: Inversion(l) garnet from X irradiation
2 OF. cytology: In( 1)12; 19-20.
origin: X ray induced in In(l)sc8+dl-49, origin: X ray induced.
y31d S8 cfo, B. discoverer: Muller.
discoverer: R. F. Grell, 1954. references: 1946, DIS 20: 67.
references: Mislove and Lewis, 1954, DIS 28: 77. genetics; Mutant for g.
genetics: Mutant for two lethals, one allelic to In(l)hil: see ln(l)drp
l(l)Jl and therefore covered by y + F and the other *ln(l)hi2: lnversion(1) from high
covered by BSY; both In(l)FM3/y+Y/BsY and cytology*. In(l)lF;20.
In(l)FM3/BsY y+ males survive. The treated chro- origin: Spontaneous in hi.
mosome carried y31d, but ln(l)FM3/y variegates discoverer: Ives.
for yellow bristles. references: Hinton, Ives, and Evans, 1952, Evolu-
other information: Used as a first chromosome bal- tion 6: 19-28.
ancer, described as FM3 in the section on bal- genetics: Male lethal.
ancers. *lnO)hi3
ln(l)FM4 cytology: ln(l)4D;20.
cytology. In(l)3C,4E-F superimposed on ln(l)lB2- origin: Spontaneous in hi.
3;2OB-D1 + In(l)4D7-El;llF2-4. discoverer: Ives,
new order: 1A - 1B2|2OB - 11F4|4E|3C - references: Hinton, Ives, and Evans, 1952, Evolu-
4D7|llF2 - 4F|3C - 1B3|2OD1 - 20F. tion 6: 19-28.
origin: X ray induced in In(l)scs+d1-49, genetics: Male lethal.
y31d SC8 dm B. *ln(1)hi4
discoverer R. F. Grell, 1954. cytology: ln(l)4C;20.
references: Mislove and Lewis, 1954, DIS 28: 77. origin: Spontaneous in hi.
genetics: Male viable and fertile. di scoverer: Ives.
other information: Used as a first chromosome bal- references: Hinton, Ives, and Evans, 1952, Evolu-
ancer, described as FM4 in the section on bal- tion 6: 19—28.
ancers. genetics: Male lethal.
*ln(l)hi5 references: 1937, DIS 8: 11.

cytology: In(l)lF;20. genetics: Associated with Hv.
origin: Spontaneous in hi. ln(l)Hw2: Inversion(l) Hairy wing
discoverer: Ives. cytology: In(l)lA2-3;lA8-Bl;lB2-3.
references: Hinton, Ives, and Evans, 1952, Evolu- new order: 1A1 - lA2|lB2 - lA3|lBl - 20.
tion 6: 19-28. origin: Spontaneous derivative of Dp(l;l)Hw =
genetics: Male lethal. Dp(l;l)lA8-Bl;lB2-3.
*ln(1)hi7 discoverer: Nichols-Skoog. 35a9.
cytology: In(l)12E;20. genetics: Associated with Hw2.
origin: Spontaneous in hi.
discoverer: Ives. *ln(1)K2: Inversion(i) of Krivshenko
references: Hinton, Ives, and Evans, 1952, Evolu- cytology: In(l)6A;9A-B.
tion 6: 19-28. origin: Spontaneous.
genetics: Male lethal. discoverer: Krivshenko, 54c24.
*ln(l)hi8 references: 1956, DIS 30: 75.
cytology: In(l)3C;20. genetics: Homozygous viable.
origin: Spontaneous in hi. *ln(l)t-272-13: Inversion(l) lethal
discoverer. Ives. cytology: In(l)lA6-Bl;llA7-8;llF2-12Al;18A4-Bl.
references: Hinton, Ives, and Evans, 1952, Evolu- new order: 1A1 - lA6|l2Al - 18A4|llA7 -
tion 6: 19-28. l B l | l l A 8 - HF2|l8Bl - 20.
genetics: Male lethal. origin: X ray induced.
*ln(l)hi9 discoverer: Demerec, 1940.
cytology: In(l)8F;20. synonym: Tp( 1)1-272-13.
origin: Spontaneous in hi. references: Sutton, 1943, Genetics 28: 210—17.
discoverer: Ives. genetics: Associated with 1(1)272-13. sc affected
references: Hinton, Ives, and Evans, 1952, Evolu- but not y, ac, or svr.
tion 6: 19-28. In(l)l-v59: Inversion(l) lethal-variegated
genetics: Male lethal. cytology: In( 1)3-4; 19-20.
cytology: In(l)4E2-3;8Al-2. discoverer: Lindsley, Edington, and Von Halle.
origin: Spontaneous in hi. references: 1960, Genetics 45: 1649-70.
discoverer Ives. genetics: Variegated for a lethal; male fertile.
references: Hinton, Ives, and Evans, 1952, Evolu- In(l)l-vl32
tion 6: 19-28. cytology: In(l)4E;19-20.
*ln(l)hm references: 1960, Genetics 45: 1649—70.
cytology: Jn(l)5C;7B;20A-F. genetics: Variegated for a lethal; male fertile.
new order: 1 - 5C|7E - 20A|7E - 5CJ20F.
origin: Recovered among progeny of Florida high. cytology: In(l)5-6;19-20.
discoverer. Ives. origin: X ray induced.
synonym: Tp(l)hill. discoverer: Lindsley, Edington, and Von Halle.
references: Hinton, Ives, and Evans, 1952, Evolu- references: I960, Genetics 45: 1649-70.
tion 6: 19-28. genetics: Variegated for a lethal; male fertile.
genetics: Male lethal. In(l)l-v227
*ln(l)hn2 cytology: ln(l)l-2;19-20.
cytology: ln(l)lC3;20. origin: X ray induced.
origin: Spontaneous in hi. discoverer: Lindsley, Edington, and Von Halle.
discoverer: Ives. references: I960, Genetics 45: 1649-70.
references: Hinton, Ives, and Evans, 1952, Evolu- genetics: Variegated for a lethal; male fertile.
tion 6: 19-28. In(l)l-v231
genetics: Male lethal. cytology: ln(l)lC-D;19-20.
*ln(l)ha3 origin: X ray induced.
cytology: In(l)4E;20. discoverer: Lindsley, Edington, and Von Halle.
origin: Spontaneous in hi. references: I960, Genetics 45: 1649-70.
discoverer: Ives. genetics: Variegated for a lethal; male fertile.
references: Hinton, Ives, and Evans, 1952, Evolu-
tion 6: 19-28. Ind)l-zwl931: Inversionfl) lethat-zeste white
genetics: Male lethal. cytology: ln(l)3A;6.
*IMDHv: hvcrsion(l) Hooked veins origin: X ray induced.
cytology: Breakpoints unknown. discoverer: Judd, 62g31.
discoverer: Tanaka, 3S®4. genetics: Mutant for l(l)zwl.
*ln(l)lzA
cytology: Jn(l)3A8-Bl;13. cytology: In(l)3E;3F;9E;9F-10A; inferred from
origin: X ray induced. Mackensen's figure; bands in 3E-F and 9E-F
discoverer: Judd, 62bl2. missing.
genetics: Mutant for I(l)zw3. new order: 1 - 3EJ9E - 3FJ10A1 - 20.
*ln(l)lzsB: Inversion(l) lozenge-spectacled of synonym: Df(l)Del lz A.
Bishop references: Mackensen, 1935, J. Heredity 26: 163—
cytology: Jn(l)8;20 (Green). 74 (fig.).
origin: X ray induced; genetics: Mutant or deficient for v but not lz or ras.
discoverer: Bishop. No clue to reason for lz appearing in name.
references: Oliver, 1947, Texas Univ. Publ. 4720: ln(l)mK; Inversion(T) miniature of Krivshenko
167-84. cytology: In(l)10E;20B. In mitotic chromosomes,
genetics: Associated with lzsB. right breakpoint is near juncture of heterochromatic
*ln(l)lzl: Inversion(l) lozenge elements hC and hD to the left of the nucleolus
cytology: In(l)8D;20D (Hannah). organizer but to the right of right breakpoint of
origin: X ray induced. In(l)sc< (Cooper, 1959, Chromosoma 10: 535-88).
discoverer: Green and Green. origin: X ray induced.
references: 1956, Z. Induktive Abstammungs- discoverer: Krivshenko, 5513.
Vererbungslehre 87: 708—21. synonym: IntflyK-.
genetics: Mutant for spectacled-like allele of lz. references: 1956, DIS 30: 75.
*ln(l)lz2 genetics: Variegated for m.
cytology: In(l)8D;20D (Hannah). *ln(l)N21*: Inversion!!) Notch
origin: X ray induced. cytology: In(l)3C;20; position of right breakpoint
discoverer: Green and Green. with respect to centromere of ring not determined.
references: 1956, Z. Induktive Abstammungs- origin: X ray induced in R(l)2.
Vererbungslehre 87: 708—21. discoverer: Barigozzi.
genetics: Mutant for a spectacled-like allele of lz. references: 1942, Rev. Biol. (Perugia) 34: 59—72.
*ln(l)lz3 genetics: Variegates for w and ec but not pn, dm, or
cytology: In(l)4D;8E2-3 (Hannah). cv. Seems to carry intermediate allele of N.
origin: X ray induced. *tn(l)N264.7
discoverer: Green and Green. cytology: In(l)3C6-7;3C8-9;8C5-7; 3C7-8 missing
references: 1956, Z. Induktive Abstammungs- (Hoover).
Vererbungslehre 87: 708—21. new order: 1 - 3C6|8C5 - 3C9J8C7 - 20.
genetics: Mutant for a spectacled-like allele of lz. origin: X ray induced,
*ln(l)lz4 discoverer: Demerec, 33k.
cytology: In(l)8A2-Bl;8D (Hannah). genetics: Deficient for fa, &pl, and fan but not w,
origin: X ray induced. rst, or dm.
discoverer: Green and Green.
references: 1956, Z. Induktive Abstammungs- cytology. In(l)lB6-7;lB10-ll;3C7-8; 1B7-10 missing
Vererbungslehre 87: 708—21. (Hoover).
genetics: Mutant for a spectacled-like allele of lz. new order: 1A1 - 1B6J3C7 - lBll|3C8 - 20.
*ln(l)lzS origin: X ray induced,
cytology: ln(l)8D;18F2-19Al (Hannah). discoverer: Demerec, 37f.
origin: X ray induced. genetics: Deficient for fa but not sc, svr, tw, bt, kz,
discoverer: Green and Green. w, rst, dm, or ec.
references: 1956, Z. Induktive Abstammungs- *ln(l)N264.52
Vererbungslehre 87: 708-21. cytology: In(l)3C3-5;20B2-Cl.
genetics: Mutant for a spectacled-like allele of lz. origin: X ray induced,
discoverer: Demerec, 38a.
cytology: In(l)8D;9B12-Cl (Hannah). genetics: Variegates for rst, fa, dm, ec, and bi but
origin: X ray induced. not w, peb, or rb.
discoverer: Green and Green.
references: 1956, Z. Induktive Abstatnmungs- cytology: ln(l)3C9-ll;20D2-El (Hoover),
Vererbungslehre 87: 708—21. origin: X ray induced.
genetics: Mutant for a spectacled-like allele of lz. discoverer: Demerec, 38d.
*ln(Vlz7 references: 1941, Proc. Intern, Congr. Genet., 7th.
cytology: In(l)8D;20D (Hannah). pp. 09-103.
origin: X ray induced. genetics: N and rst mutant but not w, fa, dm, or ec.
discoverer Green and Green. *in(l)N264-71
references: 1956, Z. Induktive Abstammungs- cytology: In(l)3C6-7;2QD-E (Sutton).
Verartnmgslehre 87: 708-21. origin: X ray induced.
genetics: Mutant for a spectacled-like allele of lz, discoverer: Demerec, 38k.
328 GENETIC VARIATIONS OF DROSOPHILA MELAHOGASTER
references: 1941, Proc. Intern. Congr. Genet., 7th. genetics: Associated with Pub. Called inversion
pp. 99-103. because of reduction in crossing over; less than 1
genetics: Carries mutant allele of N and normal percent recombination with spl and about 10 per-
alleles of w, rst, dm, and ec. cent with B.
ln(l)N264.84 *ln(l)rK: Inversion(l) rudimentary of Krivshenko
cytology: ln(l)3C6-7;20A-B (Sutton). cytology: Proximal break in heterochromatin.
origin: X ray induced. discoverer: Krivshenko.
discoverer: Demerec, 39c. references: Agol, 1936, DIS 5: 7.
references: 1941, Proc. Intern. Congr. Genet., 7th. genetics: Mutant for r.
pp. 99-103. ln(l)rb">48aH5: Inversion(l) ruby-mottled
genetics: Variegates for fa and dm but not w, rst, or cytology: In(l)3E3-4;llA7-8;20F superimposed on
bi. In(l)lB3-4;20B-DlLlB2-3;20B-DlR + In(l)4D7-
*ln(l)N264.108 El;llF2-4.
cytology: In(l)3C3-5;3E7-8;20A4-5; 3C5-E7 missing new order: 1A - 1B3|2OB - 11F4|4E1 - 11A7|3E3 -
(Sutton). 1B3|2OD1 - 20F-(3E4 - 4D7|llF2 - 11A8)|2OF.
new order: 1 - 3C3|20A4 - 3E8|20A5 - 20F. origin: X ray induced in In(l)scsl Lsc8R+dl-49.
origin: X ray induced. discoverer: Muller, Valencia, and Valencia, 1946-53.
discoverer: Demerec, 40a. references: Valencia, 1966, DIS 41: 58.
genetics: Deficient for spl, dm, and M(1)3E but not genetics: Associated with rbm4SoH5t
for w or ec. *ln(l)rg?: Inversion(l) rugose
*ln(1)N264.J12 cytology: In(l)4E;7A (J. I. Valencia).
cytology: ln(l)3C6-7;3F5-6 (Sutton). origin: X ray induced.
origin: X ray induced. discoverer: Cantor, 46d20.
discoverer: Demerec, 40b. genetics: Mutant for rg.
genetics: Carries normal alleles of w, dm, and ec. *ln(1)rgp: Inversion(l) rugose from P3 2
*h(l)N264.JU cytology: In(l)3C;4E (Darby).
cytology: ln(l)2C8-10;3C7-9 (Sutton). origin: Induced by P32,
origin: X ray induced. discoverer: Bate man.
discoverer: Sutton, 40e. references: 1951, DIS 25: 77-78.
genetics: Carries normal alleles of pn, w, rst, and genetics: Mutant for rg.
dm. ln(l)rst3: Inversion(l) roughest
ln(J)NP: Inversion(l) Notch from P*2 cytology. In(l)3C3-4;20B. Right breakpoint about
cytology: In(l)3C;8E (Darby). one-fourth the distance between the heterochro-
origin: Induced by P32. matic-euchromatic junction and the centromere, ap-
discoverer: Bateraan, 1950. proximately between heterochromatic segments hC
references: 1950, DIS 24: 55. andhD (Cooper, 1959, Chromosoma 10: 535—88).
1951, DIS 25: 77. origin: X ray induced.
genetics: Carries normal allele of w. discoverer: Gruneberg, 33116.
*ln(l)Nel.A: Inversion(l) of Nel-A 1935, J. Genet. 31: 163-84 (fig.).
cytology: In(l)12A;18D. 1937, J. Genet. 34: 169-89.
origin: Spontaneous. genetics: Left breakpoint between w and rst; right
discoverer: Nel. breakpoint to the left of bb [Emmens, 1937, J.
*ln(l)Nel.B Genet. 34: 191-202 (fig.); Kaufmann, 1942,
cytology: In(l)llA;12F. Genetics 27: 537-49 (fig.)]. Mutant for rst and in
origin: Spontaneous. X/0 male variegates for w (Gersh, 1963, DIS 37:
discoverer: Nel. 81). Precise reinversion of Jn(l)rst3 accompanied
*ln(l)ney: Inversion(T) narrow eye by reversion of phenotype reported to occur sponta-
cytology: In(l)10A;16D. neously (Grunberg, 1934) and after X irradiation of
origin: X ray induced. oocytes (Novitski, 1961, Genetics 46: 711—17) but
discoverer: Becker, 1950. not after irradiation of sperm (Kaufmann, 1942).
references: 1952, DIS 26: 69. *ln(l)rstl: Inversion(i) roughestlike
genetics: Associated with ney. cytology: Breakpoints unknown,
ln(l)pdf: Inversion(l) podfoot origin: X ray induced,
cytology: Jn(l)16B;19F-20A. discoverer: Oliver, 29d3.
discoverer: Welshons, 57h6. genetics: Associated with rstl.
references: I960, DIS 34: 54. fn(l)S: Inversion(l) of Sinitskaya
genetics: Associated with pdt. cytology: In(l)6Al-3;10F10-llAl (Slizynski, 1948,
*ln(l)Pub: lnv»rsion(1) Pub DIS 22: 77).
discoverer: P. Farnsworth. origin: X ray induced simultaneously with ln(l)scsi.
references: Lefevre, 1954, DIS 28: 75. discoverer: Sinitskaya.
references: Muller and Prokofyeva, 1934, Dokl. which Peacock interpreted as the result of non-
Akad. Nauk SSSR, n.s. 4: 74-83. random orientation of the first meiotic division
other information: In(l)scsl+S used as a crossover with respect to the postulated functional pole of
suppressor in certain balancers, e.g., Base. the primary spermatocyte. Irregularities in meiotic
ln(1)sc4: Inversion(l) scute behavior of In(l)sc4Ijsc8R in the male are affected
cytology: In(l)lB3-4;19F-20Cl (Muller and by the Y chromosome present (Peacock, 1965) and
Prokofyeva, 1934, Dokl. Akad. Nauk SSSR n.s. 4: the temperature at which meiosis occurs (Zim-
74-83; Prokofyeva-Belgovskaya, 1937, Izv. Akad. mering, 1963, Genetics 48: 133-38).
Nauk SSSR, Ser. Biol. 2: 393-426; Schultz and In(l)sc4Lsc8R/Y/Y male gives quite regular seg-
Redfield, 1951, Cold Spring Harbor Synp. Quant. regation of the two Y's and low recovery of the X.
Biol. 16: 175—97). In mitotic chromosomes, the ln(l)sc4Lsc9R: Inversion(l) scute-4 Left scute-9
right break is to the right of and near the euchro- Right
matic-heterochromatic juncture in the heterochro- cytology: In(l)lB3-4;19F-20Cl^lB2-3;18B8-9R; left
matic segment termed hD (Cooper, 1959, Chromo- breakpoint data inconsistent with genetic observa-
soma 10: 535-88). tions. Duplicated for 18B9-19F.
origin: X ray induced in a y chromosome. origin: Recombinant containing left end of In(l)sc4
discoverer: Agol, 1928. and right end of In(l)sc9.
references: 1929, Zh. Eksperim. Biol. 5: 86—101. discoverer: Muller.
1931, Genetics 16: 254-66. references: 1935, Genetica 17: 237-52.
Serebrovksy and Dubinin, 1930, J. Heredity 21: genetics: Deficient for l(l)sc. Duplicated for loci
259-65. right of 18B9 including car+ but not bb+. Male
Sturtevant and Beadle, 1936, Genetics 21: 554— lethal but viable in presence of Dp(l;2)sci 9 =
604. Dp(l;2)lBl-2;lB4-7;25-26.
Muller and Raffel, 1940, Genetics 25: 541-83. /n(7)sc^ L sc L8R : Inversionfl) scute-4 Left scute
genetics: Mutant at sc; also carries y. Left break to of Levy 8 Right
the right of sc and to the left of l(l)sc inferred cytology: In(l)lB3-4;19F-20ClLlB3-4;20B-CR. Mi-
from observations that In(l)sc8Ijsc4R is deficient totic chromosomes deficient for the proximal one-
for sc (Sturtevant and Beadle, 1936) and third of hD, all of hC, and the distal half of hB
In(l)sc4Lsc9R is male lethal except in the pres- (inferred from Cooper, 1959, Chromosome 10: 535—
ence of Dp(l;2)sc19 (Muller, 1935, Genetica 17: 88).
247). Right break in the proximal heterochromatin origin: Recombinant containing left end of In(l)sc4
to the left of bb inferred from observation that and right end of Jnfi^sc^*.
In(l)sc4Lsc8R is deficient for bb (Gershenson, references: Muller, Raffel, Gershenson, and
1933, J. Genet. 28: 297-313; Sturtevant and Prokofyeva-Belgovskaya, 1937, Genetics 22: 87—
Beadle, 1936). In(l)sc*/+ female produces about 6 93.
percent exceptional males from four-strand double genetics: Deficient for bb and the nucleolus or-
exchange. Secondary exceptions about 4 percent. ganizer.
ln(l)sc*Lsc81*: Inversion(l) scute-4 Left scute-8 ln(l)sc4Lsc51R: tnversion(l) scute-4 Left scute
Right of Sinitskaya I Right
cytology: In(l)lB3-4;19F-20ClLlB2-3;20B-DlR; cytology: In(l)lB3-4;19F-20ClLlB3-4;20B-DlR; de-
duplicated for 1B3, mitotic chromosomes deficient ficient for proximal third of hD, all of hC and hB,
for the proximal third of hD, all of hC and hB, and and the distal majority of hA (Cooper, 1959, Chro-
the distal majority of hA (Cooper, 1959, Chromo- mosoma 10: 535—88). About 0.6 the length of a
soma 10: 525-88). About 0.6 the length of a normal normal X at metaphase.
X at metaphase. origin: Recombinant containing left end of In(l}sc4
origin: Recombinant containing left end of In(l)sc4 and right end of In(l)scSi.
and right end of In(l)sc8. references: Muller, Raffel, Gershenson, and
discoverer: Gershenson. Prokofyeva-Belgovskaya, 1937, Genetics 22: 87—
references: 1933, J. Genet. 28: 297-313. 93.
1933, Biol. Zh. (Moscow) 2: 145-59, 419-24. genetics: Deficient for bb. Behavior in meiosis of
genetics: Duplicated for the sc locus, carrying both the male like that of In(l)t}C4L'Sc8R.
4L 4
ac4 and s c 8 ; deficient for the bb locus and the y ^: lnv&rsion(1) scute-4 Left yellow-4
nucleolus organizer [i.e., Dl(l)bbG]. Shown by Right
Ritossa and Spiegelmann (1965, Proc. Natl. Acad. cytology: In(l)lB3-4;l9F-20ClL1AS-B1;1SA3-4R;
Sci. U.S. 53: 737-45) to be deficient for all the duplicated for 1B1-3 and 18B1-19F.
DNA that is complementary to ribosomal RNA origin: Recombinant containing left end of In(l)&c4
present in a haploid chromosome set. In the male, and right end of In(I)y4.
In(l)Bc4LscSR frequently fails to pair with the Y references: Sturtevant and Beadle, 1936, Gen#tics
and when it does the unpaired X and Y usually 21: 554-604.
proceed to the same pole (Peacock, 1965, Genetics genetics: Duplicated for the loci of ac, «c, car, and
51: 573—83). Furthermore, reciprocal meiotic M(J)n; either deficient for y oc carries y4. Both
products are not recovered with equal frequency. male and female look normal.
In(l)sc7 genetics: Carries 1(1)] 1 +, y+ (or y31d), and a c + dis-

cytology: In(l)lB4-6;5D3-6 (Schultz). tally and 1(1)J1+, y, and ac+ proximally on long
origin: X ray induced in a wa chromosome. arm. Carries heterochromatic short arm of In(l)EN.
discoverer: Dubinin, 1929. Carries 66 + in distal heterochromatin derived from
references: 1930, Zh. Eksperitn. Biol. 6: 300—24. In(l)sc8 and a mild allele of 66 in proximal hetero-
Serebrovsky and Dubinin, 1930, J. Heredity 21: chromatin derived from In(l)EN.
259-65. ln(l)sc8Lsc4R: Inversion(i) scute-8 Left scute-4
Dubinin, 1933, J. Genet. 27: 443-64. Right
Sturtevant and Beadle, 1936, Genetics 21: 554- cytology: In(l)lB2-3;20B-DlLlB3-4;19F-20ClR; de-
604. ficient for 1B3 and duplicated for proximal part of
genetics: Mutant for sc. Normal disjunction and 33 hD, all of hC and hB, and the distal majority of
percent recombination in In(l)sc7/+ female; 26 hA. About 1.4 times the length of a normal X at
percent secondary nondisjunction and 27 percent metaphase.
recombination in In(l)sc?/+/Y female (Grell, 1962, origin: Recombinant containing left end of In(l)sc8
Genetics 47: 1737—54). wa removable from the in- and right end of In(l)sc4,
version by double crossing over in triploid. references: Sturtevant and Beadle, 1936, Genetics
In(1)sc8 21: 554-604.
cytology: In(l)lB2-3;20B-Dl (Muller and Prokofyeva, genetics: Deficient for sc; duplicated for 66 + and
1934, Dokl, Akad. Nauk SSSR n.s. 4: 74-83; the nucleolus organizer. In(l)scaLsc4R/ + female
Prokofyeva-Belgovskaya, 1937, Izv. Akad. Nauk often has crippled legs. Male survives rarely and
SSSR. Ser. Biol. 2: 393-426; Schultz and Redfield, is extreme sc.
1951, Cold Spring Harbor Symp. Quant. Biol. 16: ln(l)sc8LscLSR: Inversionfl) scute-8 Left scute
185). Mitotic chromosomes show break in proximal of Levy 8 Right
heterochromatin extremely close to the centromere cytology: In(l)lB2-3;20B-Dl^lB3-4;20B-CR; defi-
in proximal part of element hA (Cooper, 1959, cient for 1B3 and mitotic chromosomes duplicated
Chromosoma 10: 535—88). for the proximal half of hB and the distal majority
origin: X ray induced. of hA (inferred from Cooper, 1959, Chromosoma 10:
discoverer: Sidorov, 1929. 535-88).
references: 1931, Zh. Eksperim. Biol. 7: 28—40. origin: Recombinant containing left end of In(l)scs
1936, Biol. Zh. (Moscow) 5: 1-26. and right end of In(l)scLa.
Noujdin, 1935, Zool. Zh. 14: 317-52. references: Muller, Raffel, Gershenson, and
Patterson and Stone, 1935, Genetics 20: 172—78. Prokofyeva-Belgovskaya, 1937, Genetics 22: 8 7 -
Sturtevant and Beadle, 1936, Genetics 21: 554— 93.
604. genetics: Deficient for sc.
genetics: Mutant for sc and shows a Hw effect; var- ln(l)sc8Lscs1R: Inversionfl) scute-8 Left scute
iegates for ac, y, and probably 1(1)]I (Hess, 1962, of Sinitskaya 7 Right
Verhandel. Deut. Zool. Ges., Zool. Anz., Suppl. cytology: In(l)lB2'3;20B-Dl^lB3-4;20B-DlR; defi-
26; 87-92) in X/0 male. Left break between ac cient for 1B3.
and sc because induced deficiencies for the ter- origin: Recombinant containing left end of ln(l)sc8
minal uninverted portion of In(l)scs are deficient and right end of In(l)scsi.
for y and ac but not sc* (Patterson and Stone, genetics: Deficient for sc. A few extreme sc males
1935) and because In(l)sc8Lsc4R is deficient for survive.
sc (Sturtevant and Beadle, 1936). Right break be- ln(l)sc*Ly3p*: lnversion(l) scute-8 Left yellow-3
tween 66 and centromere because deficiencies for of Patterson Right
terminal genes are frequently deficient for bb cytology: In(l)lB2-3;20B-DlLlA;20R; duplicated for
(Patterson, 1933, Genetics 18: 32—52) as is 1A-B2.
In(l)ac4LscaR (Gershenson, 1933, J. Genet. 28: origin: Recombinant containing left end of In(l)sc8
297-313; Sturtevant and Beadle, 1936). ln(l)ac*/+ and right end of In(l)y3P.
female produces about 3 percent exceptional sons references: Muller, 1935, J. Heredity 26: 469-78.
from four-strand double crossing over within the in- genetics: Duplicated for y and ac loci; not deficient
version and about 8.7 percent recombination. for 66. Male viable.
In(l)nc8/-¥/Y female produces 19 percent sec- ln(l)sc8Ly4R: Inversionfl) scute-8 Left yellow-4
ondary nondisjunction and 12 percent recombina- Right
tion (Sturtevant and Beadle, 1936; Grell, 1962, cytology. In(l)lB2-3;20B-Dl^lA8-Bl;18A3-4R; du-
Genetics 47: 1737-54). plicated for 1B1-2 and 18A4-20B.
8
ÊNR: Inversiond) scuto-S Left Entire origin: Recombinant containing left end of In(l)sc8
Right and right end of In(l)y4.
cytology: In{l)lB2-3;20B-DlLlA;20;20B-CR. references: Sturtevant and Beadle, 1936, Genetics
origin: Reccfflbinant containing left end of In(l)sc8 21: 554-604.
and right end of In(l)EN. genetics: Duplicated for ac, car, M(l)n, and 66.
references: Lixtdsiey, 1958, Z. Vererbungslehre 89: Both male and female survive and show Hw effect
103-22. of In(l) sc*.
In(l)sc9 ln(1)scL^: Inversion(i) scute of Levy

cytology: In(l)lB2-3;18B8-9 (J. I. Valencia). Left cytology: In(l)lB3-4;20B-C; inferred from genetic
breakpoint irreconcilable with Muller's genetic data. In mitotic chromosomes, right break is in
evidence (1935, Genetica 17: 237-52) that the left center of heterochromatic segment hB (Cooper,
break of In(l)sc* is to the right of that of In(l)sc4. 1959, Chromosoma 10: 535-88) to right of nu-
origin: X ray induced. cleolus organizer.
discoverer: Levit, 1929. discoverer: Levy, 1932.
references: 1930, Arch. Entwicklungsmech. Organ. references: Muller, Raffel, Gershenson, and
122: 770-83. Prokofyeva-Belgovskaya, 1937, Genetics 22: 8 7 -
Norton and Valencia, 1965, DIS 40: 40. 93.
genetics: Mutant for sc. Left break to right of sc Muller and Raffel, 1938, Genetics 23: 160.
and 1(4)sc, inferred from observation that Raffel and Muller, 1940, Genetics 25: 541-83.
In(l)sc4Lsc9R is lethal in male unless Dp(l;2)sc1^ genetics: Mutant for sc. Left break between sc and
is present (Muller, 1935). Right break right of sby, l(l)sc, probably based on viability of reciprocal
smd, and coc and left of car, as shown by the de- recombinants with In(l)sc4 and In(l)scsl (Raffel
ficiency for sby, smd, and coc of In(l)y4Lsc9R and Muller, 1940). Right break to right of 66 be-
(Norton and Valencia, 1965). cause In(l)sc4LscLSR deficient for 66 (Muller,
In(l)sc* o: see In(l)ac3 Raffel, Gershenson, and Prokofyeva-Belgovskaya,
ln(l)sc29 1937).
cytology: In(l)lB,13A2-5 (Raffel) . ln(l)scL6Lsc*R: Inversion(l) scute of Levy 8
discoverer: Agol, 1930. Left scute-4 Right
genetics: Mutant at sc. Left break to right of 1(1 )sc cytology: In(l)lB3-4;20B-CLlB3-4;19F-20ClR. Mi-
(Muller). totic chromosomes duplicated for proximal third of
*ln(1)scS2c hD, all of hC, and the distal half of hB (inferred
origin: Spontaneous. from Cooper, 1959, Chromosoma 10: 535—88).
discoverer: Green, 52c. origin: Recombinant containing left end of In(l)scL8
references: 1952, DIS 26: 63. and right end of In(l)sc4.
genetics: Mutant for sc and su(s); i.e., su(s)s3c. references: Muller, Raffel, Gershenson, and
Inversion inferred from failure of sc$2c ô recom- Prokofyeva-Belgovskaya, 1937, Genetics 22: 87—
bine with ras or v. 93.
*ln(l)sc90 genetics: Duplicated for 66 and the nucleolus or-
cytology: In(l)lB4-7;1D2-E1; inferred from Goldat's ganizer.
fig. 2.
origin: X-ray-induced derivative of set. ln(1)scL8Lsc8R: Inversion(l) scute of Levy 8
discoverer: Goldat. Left scute-8 Right
references: 1936, Biol. Zh. (Moscow) 5: 803—12. cytology: ln(l)lB3-4;20B-CLlB2-3;20B-DlR; dupli-
genetics: Mutant for sc. cated for 1B3 and mitotic chromosomes deficient
for proximal half of hB and distal majority of hA
cytology: In(l)lB2-3;llD3-8. (inferred from Cooper, 1959, Chromosoma 10: 535—
origin: X ray induced. 88).
discoverer: Sutton, 39b. origin: Recombinant containing left end of In(l)scL8
references: 1943, Genetics 28: 210—17. and right end of In(l)sc8.
genetics: Mutant for sc but not y, ac, or svr. references: Muller, Raffel, Gershenson, and
ln(l)sc260-22 Prokofyeva-Belgovskaya, 1937, Genetics 22: 87—
cytology: In(l)lB2-3;lE2-3. 93.
origin: X ray induced. genetics: Duplicated for sc but not 66. Survives as
discoverer: Sutton, 39f. X/0 male and homozygous female.
genetics: Mutant for sc but not y, ac, or svr. In(l)scl-8LSCS1R: lnversion(l) scute of Levy 8
*ln(1)scA: Inversion(l) scute of Agol Left scute of Sinitskaya 7
discoverer: Agol. Right
references: 1936, DIS 5: 7. cytology: In(l)lB3-4;20B-C^lB3-4;20B'DlR; mitotic
genetics: Mutant for sc; semilethal. Genetically, chromosome deficient for proximal half of hB and
appears to extend from sc to near r (54.5). distal majority of hA (inferred from Cooper, 1959,
ln(l)scJ1: Inversion(l) scute of Jacobs-Muller Chromosoma 10: 535—88).
cytology: ln(l)lA4-5;lB4-5. origin: Recombinant containing left end of In(l)scLS
discoverer: Jacobs-Muller. and right end of fnfl>c 5 ^.
references: Muller, 1932, Proc. Intern. Congr. references: Muller, Raffel, Gershenson, and
Genet., 6th. Vol. 1: 225. Prokofyeva-Belgovskaya, 1937, Genetics 22: 87—
Muller, Prokofyeva, and Raffel, 1935, Nature 135: 93.
253-55. genetics: Homozygous viable and fertile. Does not
genetics: Mutant for sc and 1(1 )J1. affect expression of variegation of
ln(l)scs1: Inversion(l) scute of Sinitskaya genetics: Mutant for sc. Left break between ac and
cytology: In(l)lB3-4;20B-Dl; inferred from genetic sc and right break between bb and the centromere;
identity of left break with that of ln(l)sc* and of determined by the viability and bb + phenotype of
right break with that of In(l)sc8. In mitotic chro- reciprocal recombinants with In(l)sc8. Muller and
mosomes, right break is in proximal part of most and Valencia (1947) presume the right break to be
proximal heterochromatic segment, hA (Cooper, to the right of that of In(l)sc8 because of the more
1959, Chromosoma 10: 535-88). extreme ac and sc phenotype of In(l)scV2.
origin: X ray induced simultaneously with In(l)S. *ln(J)sd2: Inversion(i) scalloped
discoverer Sinitskaya, 34c. origin: X ray induced.
references: Muller and Prokofyeva, 1934, Dolk. discoverer: Panshin, 33g7.
Akad. Nauk SSSR 4: 74-83. references: 1935, DIS 3: 28.
Muller and Raffel, 1938, Genetics 23: 160. genetics: Mutant for sd. Crossing over inhibited.
Raffel and Muller, 1940, Genetics 25: 541-83. ln(J)sdS8d
Crew and Lamy, 1940, J. Genet. 39: 273-83. origin: Gamma ray induced.
genetics: Mutant for sc. Left break between sc and discoverer: Ives, 58dl4.
lfl^c, probably based on the viability of reciprocal references: 1961, DIS 35: 46.
recombinants with In(l)sc4 and In(l)scLS (Raffel genetics: Mutant for sd. Not a translocation geneti-
and Muller, 1940). Right break to right of bb be- cally; reduces recombination between ras and f by
cause In(l)sc4LscSiR is deficient for bb (Muller, 80 percent.
Raffel, Gershenson, and Prokofyeva-Belgovskaya, ln(J)sdx: Inversion(l) spreadex
1937, Genetics 22: 87-93). origin: X ray induced.
ln(1)scsllLsc4R: Inversion(l) scute of Sinitskaya discoverer: Muller.
1 Left scufe-4 Right references: 1965, DIS 40: 35.
cytology: In(l)lB3-4;20B-Dl^lB3-4;l9F-20C1R; genetics: Associated with sdx.
duplicated for proximal part oihD, all of hC and *ln(l)stx: Inversion(l) streakex
hB, and distal majority of hA. About 1.4 times the origin: X ray induced.
length of a normal X at metaphase. discoverer: Muller, 26k30.
origin: Recombinant containing left end of ln(l)scsl references: 1935, DIS 3: 30.
and right end of ln(l)sc4. genetics: Associated with stx.
genetics: Duplicated for bb+ and the nucleolus or- ln(l)sx: Inversion(l) sexcombless
ganizer. Carries twice the amount of DNA that is cytology: In(l)llD4-6;14B8-9 + In(l)llE2-6;15E2-4.
complementary to Drosophila ribosomal RNA and Is new order: 1 - HD4|l4B9 - 15E2|llE2 -
found in a normal haploid chromosome set (Ritossa 11D6|14B8 - HE6|l5E4 - 20.
and Spiegelmann, 1965, Proc. Natl. Acad. Sci. U.S. origin: X ray induced.
53: 737-45). discoverer: Muller, 261.
ln(l)scs1Lsc8R: fnversion(l) scute of Sinitskaya references: Mukherjee, 1963, DIS 38: 62 (fig.),
7 Left scute-8 Right genetics: Associated with mutant sx, which is male
cytology: In(l)lB3-4;20B-DlLlB2-3;20B-DlR; du- sterile.
plicated for 1B3. *ln(l)Thl: Inversion(l) Thymidine
origin: Recombinant containing left end of Jn(l)scsi cytology. In(l)12C;16C t
and right end of In(l)sc8. origin: Induced by ingested H3-thymidine.
genetics: Duplicated for sc. discoverer: Kaplan.
ln(l)scSlLscL8R: InvetshnfT) scute of Sinitskaya genetics: Male lethal.
1 Left scute of Levy 8 Right ln(l)vao: Inversion(l) varied outspread
cytology. In(l)lB3-4;20B-Dl^lB3-4;20B-CR; mitotic cytology: In(l)18C5-6;l9E7-8.
chromosomes duplicated for proximal half of hB origin: Induced by triethylenemelamine (CB. 1246).
and distal majority of hA (inferred from Cooper, discoverer: Fahmy, 1953.
1959, Chromosoma 10: 535—88). references: 1959, DIS 33: 94.
origin; Recombinant containing left end of In(l)scsi genetics: Mutant for vao; variegated for an eye
and right end of In(l)scL8. color, possibly car. Male sterile.
references: Muller, Raffel, Gershenson, and *ln(1)w2S8-S2: Inversion(l) white
Prokofyeva-Belgovskaya, 1937, Genetics 22: 8 7 - cytology: In(l)3C7-9;8Ell-8Fl (Sutton).
tn(l)scv2: Inversion(l) scute of Valencia discoverer: Demerec, 40a.
cytology: In(l)lB2-3;20B-F; inferred from genetic genetics: Mutant for w and rst but not for spl, Iz,
identity with In(l)sc8. Mitotic X has break at right dvr, or tip.
end of proximal beterochromatic element hA very *ln(1)wG; Inversion(l) white of Goldschmidt
close to centromere (Cooper, 1959, Chromosoma cytology: In(l)3C;3D-E (Kodani).
10: 535-88). origin: X ray induced in In(l)yo = In(l)lA;lC3-4.
origin: Gamma ray induced. discoverer: Goldschmidt.
discoverer: J. I. Valencia, 23h46. references: 1945, Univ. Calif. (Berkeley) Publ.
references; Mailer and Valencia, 1947, DIS 21: 70. Zool. 49: 522.
m4: Inversion(l) white-mottled new order: 20 - 3C5|20Al|lA4 - 3C3|20F«20.

cytofogy: In(l)3Cl-2;20A (Sutton). Right breakpoint Tentative.
in mitotic chromosomes is to the left of 66 in hC or origin: X-ray-induced derivative of R(l)2 = R(1)1A3-
hD (Cooper, 1959, Chromosoma 10: 535-88). 4;19F-2OA1 opened in inverted order.
origin: X ray induced. discoverer: M. A Bender, 55b28.
discoverer: Muller, 1929. references: 1955, DIS 29: 69.
references: 1930, J. Genet. 22: 299-334. genetics: Variegated for w.
genetics: Variegated for w. Left break to the left of ln(l)w<»J; Inversion(l) white-mottled of Jonsson
w. cytology: In(l)3C2-3;20.
ln(l)w»4Lrst3R: lnver$ion(l) white-mottled 4 origin: Associated with T(l;4)w™J = T(1;4)3C2-
Left roughest-3 Right 3;20;102C.
cytology: In(l)3Cl-2;20AL3C3-4;20BR deficient for
3C1-2. ln(1)wlt>Jl-rst3R: Inversion(T) white-mottled of
origin: Recombinant containing left end of In(l)wm4 Jonsson Left roughest-3 Right
and right end of In(l)rst3. cytology: In(l)3C2-3:20L3C3-4;20BR; deficient for
genetics: Deficient for w and 1(1)3C3 but not rst 3C3.
(Lefevre and Wilkins, 1966, Genetics 53: 175-87). origin: Recombinant carrying left end of In(l)wmJ,
ln(l)wm4l-wm-iR: Inversion(l) white-mottled 4 which is part of T(l;4)wmJ, and right end of
Left white-mottled of Jons son In(l)rst3.
Right references: Lefevre, 1963, DIS 37: 49—50.
cytology: In(l)3Cl-2;20AL3C2-3;20R; deficient for Lefevre and Wilkins, 1966, Genetics 53: 175-87.
3C2 and for an undetermined portion, including the genetics: Deficient for 1(1)3C3 but not w.
centromere, of the base of the X.
new order: 1 - 3 C l | 2 0 A - 3C3|lO2C - 101A. ln(l)wmMc: Inversian(l) white-mottled of McLean
origin: Recombinant containing left end of In(l)wm4 cytology: In(l)3Cl-2;20A-C; inferred from genetic
and right end of In(l)wmJ, which is part of data.
T(1;4)W"J. origin: X ray induced.
references: Lefevre, 1963, DIS 37: 49-50. discoverer. McLean.
Lefevre and Wilkins, 1966, Genetics 53: 175-87. references: Muller, 1946, DIS 20: 68.
genetics: Deficient for white; male viable and there- genetics: Variegates for w and rst. Complementary
fore not deficient for l(l)3C3. Also deficient for single recombinants between In(l)wm4 =In(l)3Cl-
proximal heterochromatin, probably including 66. 2;20A and In(l)W"Mc a r e viable, fertile, and 66 + .
ln(l)w»Slb Left breakpoints therefore identical and right
cytology: In(l)3Cl-2;20; right break proximal to the breakpoints on the same side of 66.
nucleolus organizer (Gersh).
origin: X ray induced. ln(l)w*C: Inversion(l) white-variegated of
discoverer: Baker, 51bl9. Catcheside
genetics: Variegated for w and rst. Recombinant cytology: ln(l)3C 1-2; 19-20 superimposed on
carrying left end of In(l)wmSib and right end of the R(1)1A3-4;19F-2OA1.
4-centric element of T(l;4)wmJ = T(1;4)3C2- new order: |lA4 - 3 C 1 J 1 9 - 3C2|20«20F - 20Al|.
3;20;102C is white eyed and male viable, indi- origin: X ray induced in R(l)2.
cating that In(l)wmS1*>, like In(l)W»4 ~In(l)3Cl- discoverer Catcheside.
2;20A, is broken between 3C1 and 2 (Gersh). references: Hinton, 1955, Genetics 40: 952—61.
ln(J)wm53i genetics: Variegates for w, rst, spl, and N but not y.
cytology: In(l)lA;3C3-5;20;20B-C;20C-F. Inferred X/Y male viability reduced; X/Y/Y male more
from origin. viable. Characterized by variable degree of insta-
new order: 20 - 3C5|20C|lA - 3C3|20F-20. bility manifested by production of gynandromorphs,
Tentative. X/0 males, and dominant lethals. An extreme ex-
origin: X ray induced in In(l)EN = In(l)lA;20;20B-C. ample gave 140 females, 106 gynandromorphs, 181
discoverer: M. A Bender, 53j. X/0 males, and 868 dominant lethals among 1295
references: 1955, DIS 29: 69. putative ring/rod zygotes. Small ring-shaped dupli-
genetics: Variegated for w. cations are generated infrequently (analysis by
Hinton, 1955). Behavior of rod derivatives of
cytology: In(l)3C3-5;20D. In(l)wvC (Hinton, 1957, Genetics 42: 55-65) sug-
origin: Neutron induced. gests generation of dicentrics through sister-strand
discoverer Mickey, 5413. fusion rattier than exchange. Fusion postulated to
references: 1963, DIS 38: 29. occur in heterochromatin of the 3C1J19 reunion
genetics: Variegated for w, point of In(I)wvC. Mitotic abnormalities in
ln(lhm5Sb cleavage of In(l)wvC embryos described (Hinton,
cytology: In(l)lA3-4;3C3-5;20;19F-20Al;20Al-F. 1959, Genetics 44: 923-31). Chromosome tends to
Inferred from origin. Appears as a rod in mets- become stable in stocks. Viability and fertility
phase. correlated with stability.
ln(l)y3P: Inversion(l) yellow-3 of Patterson genetics: Deficient for ac, car, M(l)n, and bb but
cytology: In(l)lA;20. Reported as In(l)lBl-2;20 by not svr; either deficient for y or carries y4. Heter-
Muller and Prokofyeva (1935), but this is contra- ozygous female Minute and poorly viable but
dictory to subsequent observations placing y in fertile. Male lethal.
1A. ln(l)y4Lsc9R: Inversion(l) yellow-4 Left scvte-9
origin: X ray induced. Right
discoverer: Patterson, 31e25. cytology: In(l)lA8-Bl;l8A3-4LlB2-3;l8B8-9*;
references: Muller, 1935, Genetica 17: 237-52. deficient for 1B1-2 and 18A4-B8.
Muller and Prokofyeva, 1935, Proc. Natl. Acad. origin: Recombinant containing left end of In(l)y4
Sci. U.S. 21: 16-26. and right end of In(l)sc9.
Sidorov, 1936, Biol. Zh. (Moscow) 5: 3-26. references: Norton and Valencia, 1965, DIS 40: 40.
genetics: Variegated for y and, to a lesser extent, genetics: Deficient for ac, sby, smd, and coc but
for Hw. Genetic breaks between 1(1)J1 and y and not 1(1)J1, fu, hdp, bkl, obi, crk, ton, bk, Oil, or
between bb and centromere. Inversion slightly pph. Either deficient for y or carries y4. Male
longer than In(l)sc8. lethal.
ln(l)y3Pl-sC8R: Inversion(l) yellow-3 of *ln(l)yS
Patterson Left scute-8 Right cytology: In(l)lA-B;14D (Muller and Raffel).
cytology: In(l)lA;20LlB2-3;20B-Dl^; deficient for discoverer: Patterson.
1A-B2. genetics: Mutant for y. Recessive lethal associated
origin: Recombinant containing left end of In(l)y3P with right breakpoint.
and right end of In(l)scs. *ln(l)yG: Inversion(l) yellow of Goldschmidt
references: Muller, 1935, J. Heredity 26: 469-78. cytology: In(l)lA;lC3-4 (Kodani).
genetics: Deficient for y and ac but not 1(1)J1, sc, origin: Spontaneous.
or bb. Male viable. discoverer Goldschmidt.
WVy* synonym: In(l)yPx bt.
cytology: In(l)lA8-Bl;l8A3-4 (Norton and Valencia, genetics: Mutant for y.
1965, DIS40: 40). *ln(l)z2: Inversion(l) zeste
origin: X ray induced. cytology: Dp(l;l)2C10-Dl;4D2-4;18F-19A.
discoverer. Serebrovsky. new order: 1 - 4D2|l8F - 2Dl|l9A - 20.
references: Dubinin and Friesen, 1932, Biol. Zentr. origin: X ray induced.
52: 147-62. discoverer: Gans.
Sturtevant and Beadle, 1936, Genetics 21: 554— references: 1953, Bull. Biol. France Belg., Suppl.
604. 38: 1-90.
genetics: Mutant for y. Right break between 1(1 )J1 JnjflJK; seeln(l)mK
and ac; left break between fu on left and sby, smd,
and coc on right; shown by deficiency of ln(1LR)l-vl39: Inversion(lLR) lethal-variegated
In(l)y*L8C9R for ac, sby, smd, and coc but not cytology: In(lLR)3C6-7.
1(1)] 1 or fu (Norton and Valencia, 1965). In(l)y4/+ origin: X ray induced.
female produces about 2 percent exceptional sons discoverer Lindsley, Edington, and Von Halle.
from four-strand double exchange in the inverted references: 1960, Genetics 45: 1649—70.
regions; In(l)y4/+/Y female produces about 7 per- Gersh, 1965, Genetics 51: 477-80 (fig.).
cent secondary exceptions (Sturtevant and Beadle, genetics: Variegated for w, rst, and a lethal; re-
1936). quires two Y chromosomes for survival; X/Y/Y
ln(J)y4t-sc4R: Inversion(l) yellow-4 Left scute-4 male fertile. Recombinant carrying left end of the
Right 4DXp element of TXl;4)w™5 = T(l;4)3C3-4;101Fl-2
cytology: Jn(l)lA8-Bl;18A3-4LlB3-4;19F-20ClR; and right end of In(lLR)l-vl39 variegates for w but
deficient for 1B1-3 and 18A4-19F. not for rat or the lethal.
origin: Recombinant containing left end of In(l)y4
and right end of In(l)sc4. *ln(JLR)sc260.25: Inversion(lLR) scute
references: Sturtevant and Beadle, 1936, Genetics cytology: Jn(lLR)lB2-3.
21: 554-604. origin: X ray induced.
genetics: Deficient for ac, sc, car, and M(l)n but discoverer. Sutton, 39k.
not pn, rat, or o*. Heterozygous female Minute and synonym: Tp(l)4sc^^0-25,
poorly viable but fertile. Male lethal. references: 1940, Genetics 25: 628—35 (fig.).
/ttfJiriLscSR; Inversiond) yellow-4 Left scute-8 genetics: Mutant for sc; variegated for y and ac, but
Right not svr. Genetic tests indicate loci of 1(1}J 1, y
cytology: In(l)lA8-Bl;18A3-4L1B2-3;2OB-Dl **; and ac are located at the base of X to the right of
deficient for 1B1-2 and 18A4-20B. 64. Sutton judged it to be a transposition of
origins Recombitmnt containing left end of ln(l)y4 1A1-B2 into the proximal heterochromatin, but
and right end of lrt(ljsc*. since this requires three breaks with one to the
references: Sturtevant and Beadle, 1936, Genetics left of 1A1, a pericentric inversion is deemed more
21: 554-604. probable. Recombination between
and a normal sequence yields Df(l)sc260~2S = references: 1955, Genetics 40: 5—23.
Df(l)lB2-3 and Dp(l;l)sc2 60-25 = Dp(l;l)lB2-3. genetics: Associated with bwR67.
The deficiency is deficient for 1(1)]1, y, and ac *ln(2)bw*73
(Sutton, 1940). cytology: In(2)40F-41A;59E4-Fl.
ln(lLR)sc^1: Inversion(lLR) scute of Valencia origin: X ray induced.
cytology: In(lLR)lA8-C3; inferred from genetic re- discoverer: Slatis.
sults. references: 1955, Genetics 40: 5—23.
origin: Gamma ray induced. genetics: Associated with bwR73,
discoverer: J. I. Valencia, 46h23. *1n(2)bw*7 9
synonym: Jnp(l)scvl (Inp symbolizes a pericentric cytology: In(2)40F-41A;59F2-3.
inversion). origin: X ray induced.
references: Muller and Valencia, 1947, DIS 21: 69— discoverer: Slatis, 50g26.
70. references: 1955, Genetics 40: 5—23.
genetics: Mutant for ac and sc. A single exchange genetics: Associated with bwR7$.
between In(lLR)scvl and a normal X chromosome ln(2)C56: lnversion(2) Crossover suppressor
produces one recombinant with the left end of cytology: In(2)40-41;59B; position of left breakpoint
In(lLR)scvl that is deficient for the tip of X, in relation to centromere not determined.
Df(l)scvi, and one with the right end of origin: X-ray induced.
In(lLR)scv* that is duplicated for the tip of X, discoverer: Roberts, 1964.
Dp(l;l)scvl. Left break between ac and M(l)Bld genetics: Homozygous lethal. Recombination be-
based on observation that Dp(l;l)scvi is dupli- tween b and sp strongly reduced.
cated for ac, and Df(l)scVi is deficient for ac but ln(2)C113
not M(l)Bld. Right break in XR. Dp(l;l)scvi cytology: In(2)40-41;46D; position of left break with
carrying y in normal position and y+ in duplicated respect to centromere not determined.
region provides an excellent marker system for origin: X ray induced.
right end of the X. discoverer: Roberts, 1965.
genetics: Homozygous lethal. Recombination be-
*ln(2)bwR1s: lnversion(2) brown-Rearranged tween b and sp reduced.
cytology: ln(2)40F-41A;59E4-Fl. ln(2)CU2
origin: X ray induced. cytology: In(2)36B-C;40-41; position of right break-
discoverer: Slatis. point with respect to centromere not determined.
references: 1955, Genetics 40: 5—23. May contain a T(2;3).
genetics: Associated with bwR18. origin: X ray induced.
*ln(2)bw*3* disoveren Roberts, 1965.
cytology: In(2)40F-41A;59Dll-El. genetics: Homozygous viable. Recombination be-
origin: X ray induced. tween al and b reduced rather sharply.
discoverer: Slatis. H2K224
references: 1955, Genetics 40: 5—23. cytology: ln(2)25E;40-41; position of right break-
genetics: Associated with bwR3S. point with respect to centromere not determined,
*ln(2)bw*45 discoverer: Roberts, 1965.
cytology: In(2)40F-41A;59E3-4. genetics: Homozygous lethal. Recombination be-
origin: X-ray-induced derivative of bw. tween al and b virtually eliminated.
discoverer: Slatis. ln(2)C282
references: 1955, Genetics 40: 5-23. cytology: In(2)31E;40-41; position of right b*e»k-
genetics: Associated with bwR4S. point with respect to centromere not determined.
cytology: In(2)40'41;59Dll-El. discoverer Roberts, 1965.
origin: X ray induced. genetics: Homozygous viable. Recombination be-
discoverer: Slatis. tween al and b strongly reduced.
genetics: Associated with bwR47. *ln(2L)SJd
*ln(2)bw*56 cytology: In(2L)25A;29F.
cytology: In(2)40F-41A;59D-E. origin: Neutron induced.
origin: X ray induced. discoverer: Mickey, 53d4.
di scoverer: Slatis. references: 1963, DIS 38: 29.
references: 1955, Genetics 40: 5-23. other information: Eye color mottled.
genetics: Associated with bwRS6. tn(2L)A
cytology: In(2L)26A;33E.
cytology: In(2)4QF-41A;59E4-Fl. origin: Naturally occurring inversion.
origin: X ray induced. discoverer: Oshima and Watanabe.
discoverer: Slatis. references: 1965, DIS 40: BB.
*ln(2L)ast™2: lnversion(2L) asteroid-reverted ln(2L)dpolvR: lnvershn(2L) dumpy-oblique lethal

cytology: In(2L)21E2-3;31. vortex Ruffled
origin: X ray induced inas(. cytology: In(2L)25A;25B3-4.
discoverer: E. B. Lewis, 1942. origin: X ray induced.
references: 1945, Genetics 30: 158. discoverer: Schultz, 33a25.
genetics: Partial reversion of ast. genetics: Mutant at dp. Homozygous lethal.
ln(2L)C123: lnversion(2L) Crossover *ln(2L)ho40: lnversion(2L) heldout
suppressor cytology: In(2L)21D4-El;22E2-3.
cytology: In(2L)23D-E;38C;39A. origin: X ray induced.
new order: 21 - 23D|39A - 38C|23E - 38C|39A - discoverer: E. B. Lewis, 1940.
60. synonym: In(2L)ho.
discoverer: Roberts, 1965. genetics: Mutant for ho but not S or ast. Homozy-
genetics: Homozygous viable. Recombination re- gous male sterile.
duced in 2L. ln(2L)K
ln(2L)C127 cytology: In(2L)22D;26B.
cytology: In(2L)23C;32A. discoverer Oshima and Wa tana be.
discoverer Roberts, 1965. *ln(2L)lt'»2: lnversion(2L) light-mottled
genetics: Homozygous viable. Recombination be- cytology: In(2L)22F-23A;40B-F.
tween al and b virtually eliminated. origin: X ray induced.
ln(2L)C236 discoverer: Hessler, 1957.
cytology: In(2L)22B;25F. references: 1958, Genetics 43: 395-403.
origin: X ray induced. genetics: Variegated for It.
discoverer: Roberts, 1965. *ln(2L)lt">20
genetics: Homozygous lethal. Recombination be- cytology: In(2L)32C;40B-F.
tween al and breduced. origin: X ray induced.
ln(2L)C263 discoverer: Hessler, 1957.
cytology: In(2L)24C;2SF;26F; 25F-26F missing. references: 1958, Genetics 43: 395—403.
new order: 21 - 24C|25F - 24C|26F - 60. genetics: Variegated for It.
origin: X ray induced. *ln(2L)lt">26
discoverer: Roberts, 1965. cytology: In(2L)27C;40B-F.
genetics: In(2L)C263/SMl and In(2L)C263/SM5 fe- origin: X ray induced,
males nearly sterile. Recombination reduced in discoverer: Hessler, 1957.
2L. Homozygous lethal. references: 1958, Genetics 43: 395—403.
genetics: Variegated for It.
ln(2L)Cy: lnversion(2L) Curly *ln(2L)Ml: tnversion(2L) of Mourad
cytology: In(2L)22Dl-2;33F5-34Al. cytology: In(2L)38E;40F.
origin: Naturally occurring inversion. origin: Spontaneous.
discoverer: Ward, 2If. discoverer Mourad and Mallah.
references: 1923, Genetics 8: 276—300. references: 1960, Evolution 14: 166-70.
Sturtevant, 1931, Carnegie Inst. Wash. Publ. No.
421: 20. cytology: In(2L)2lF;33A.
genetics: Exists with and without Cy. Homozygous origin: Spontaneous.
viable without Cy. Crossing over in In(2L)Cy/+ discoverer Mourad and Mallah.
heterozygote greatly reduced in 2L. references: 1960, Evolution 14: 166—70.
other information: The combination of In(2L)Cy + ln(2L)NS: lnversion(2L) from Nova Scotia
ln(2R)Cy often used to balance chromosome 2. cytology: In(2L)23E2-3;35Fl-2 (Bridges and Li in
Balancers usually carry a dominant such as Cy, Morgan, Bridges, and Schultz, 1936, Carnegie Inst.
S2, Bl, or L4 and one or more of the following: Wash. Year Book 35: 292).
a/2, dptvl, dplvl, E(S), b, pr, ltf, cn origin: Naturally occurring inversion.
discoverer Sturtevant, 13i.
tn(2L)Cyl-t*: lnversion(2L) Curly-Left t-Right synonym: CIIL; C2L.
cytology: In(2L^2Dl-2;33F5-34A1^22D3- references: Sturtevant, 1919, Carnegie Inst. Wash.
EI;34A8-9R. Deficient for 22D2 and 34A1-8. Publ. No. 278: 305-41.
origin: Recombinant carrying left end of In{2L)Cy genetics: Crossing over in 2L greatly reduced; none
and right end of In(2L)t. between S and b; 0.3 percent between b and pr.
discoverer: Bridges. ln(2L)t: lnversion(2L) t
references: Morgan, Bridges, and Schultz, 1937, cytology. In(2L)22D3-El;34A8-9 (Bridges and Li in
Carnegie lust. Wash. Year Book 36: 300—1. Morgan, Bridges, and Schultz, 1936, Carnegie Inst.
genetics: Acts as suppressor of S; usually carries Wash. Year Book 35: 292).
Cy. origin: Naturally occurring inversion.
discoverer: Bridges, 21a30. new order: 21 - 40F|55E - 51F|57E - 55E|57E -

synonym: C(2;3); C(2L)T; C(2L)HR. 59D8|51F - 40F|59D9 - 60.
references: Sturtevant, 1931, Carnegie Inst. Wash. origin: X-ray-induced derivative of bw.
Publ. No. 421: 20. discoverer: Slatis, 48kl6.
other information: Found in many natural popula- references: 1955, Genetics 40: 5—23.
tions (e.g., Waiters, 1944, Texas Univ. Publ. genetics: Associated with bw&3.
4445: 129-74; Oshima and Watanabe, 1965, DIS *ln(2LR)bw*20
40: 88). cytology: In(2LR)40D;59D5-6.
ln(2L)Tg: lnversion(2L) Tegula origin: X ray induced.
cytology: In(2L)2l6;22F. discoverer: Slatis.
discoverer: E. B. Lewis, 1962. genetics: Associated with bwR20t
references: Mora, 1963, DIS 38: 32. *ln(2LR)bw*5S
genetics: Associated with Tg. cytology: In(2LR)24Bl-D;42B + In(2R)40F'
41A;59D4-5.
*ln(2LR)40d new order: 21 - 24D|42E - 41A|59D4 -42E|24E1 -
cytology: In(2LR)26D;41A-B. 40F|59D5 - 60.
discoverer: T. Hinton and Atwood, 40d. discoverer: Slatis, 50d23.
references: Demerec, Kaufmann, Sutton, and Fano, references: 1955, Genetics 40: 5—23.
1941, Carnegie Inst. Wash. Year Book 40: 225- genetics: Associated with bw^SS.
34. ln(2LR)bwV1: lnversion(2LR) brown-Variegated
Hinton, 1942, DIS 16: 48. cytology: In(2LR)21C8-Dl;60Dl-2 +
genetics: Variegated for a dominant dark eye color In(2LR)40F;59D4-El.
and irregular facets; more extreme at low tempera- new order. 21A - 21C8|6OD1 - 59El|40F -
ture. Homozygous lethal. Certain stocks con- 59D4|40F - 21Dl|60D2 - 60F.
taining In(2LR)40d fail to grow on media lacking origin: X ray induced.
RNA or adenine (Hinton, Ellis, and Noyes, 1951, discoverer: Muller, 1929.
Proc. Natl. Acad. Sci. U.S. 37: 293-99). This was synonym: Ins(2LR)Pm: Inversion(2LR) Plwn.
true at pH 7.0 but not at pH 5.0 (Ellis, 1959, references: 1930, J. Genet. 22: 299-334 (fig.).
Physiol. Zool. 32: 29-39). Glass, 1934, J. Genet. 28: 69-112 (fig.).
ln(2LR)102 1934, Am. Naturalist 68: 107-14.
cytology: In(2LR)26A;51C + In(2R)41;57A. Bridges, 1937, Cytologia (Tokyo), Fujii Jub. Vol.
new order: 21 - 26A|51C - 41 |57A - 51C|26A - 2: 745-55.
41 |57A - 60. genetics: Mutant for ds; variegated for It, bw, mi,
origin: X ray induced in dsw sp3. and abb. al* arose after origin. Double crossovers
discoverer: R. F. Grell, 53k. in 2L but not 2R fairly frequent. Single exchange
references: Kramer and Lewis, 1956, J. Heredity 47: in region 21D1-40F of 2L between In(2LR)bwv*
132-36. and a normal sequence produces a recombinant
Grell and Lewis, 1956, DIS 30: 71. carrying left end of normal chromosome 2, which is
other information: Useful as a balancer. duplicated for 21A1-C8 and deficient for 60D2-F5.
*ln(2LR)aM60: lnversion(2LR) arc of Meyer Heterozygote for this recombinant poorly viable,
cytology: Breakpoints unknown. fertile, brown-Variegated, Minute, and dwarf with
origin: X ray induced. pebbled arc wings; deficient for locus of M(2)c.
discoverer: Meyer, 6Of. Reciprocal recombinant deficient for 21A1-C8 and
references: 1963, DIS 37: 50. duplicated for 60D2-F5; heterozygote poorly viable,
genetics: Associated with aM6°. fertile, bw\ Minute giant; deficient for al and
*ln(2LR)alM60: lnversion(2LR) aristaless of Meyer M(2)21Cl-2.
discoverer Meyer, 60f. origin: X ray induced.
references: 1963, DIS 37: 50. discoverer: Van Atta.
genetics: Mutant for a/. Homozygous lethal. Inver- references: 1932, Genetics 17: 637-59.
sion inferred from crossing over inhibition in 2L genetics: Variegated for bw. Breaks most probably
and 2R. just to the left of centromere and near bw.
*ln(2LR)ah: lnversion(2LR) aristaless-variegated ln(2LR)kwV30kl
cytology: In(2LR)21B-Cl,41. origin: X ray induced.
origin: X ray induced. discoverer Van Atta.
discoverer: E. B. Lewis, 1940. references: 1932, Genetics 17: 637-59.
references: 1945, Genetics 30: 137—66. genetics: Variegated for aw. Breaks most likely
genetics: Variegated for at. Homozygous lethal. just to the left of centromere and near hw.
ln(2LR)bwR3: lnversion(2LR) brown-Rearranged
cy tol ogy: In(2LR)40F;SlF;55E;57E;58DS-9. cytology: lti(2LJR)40F;S9E.

discoverer: Dobzhansky, 32g6. genetics: Variegated for It.
synonym: In(2LR)Pm3: Inversion(2LR) Pltxm-2. *ln(2LR)lt>*33
references: Schultz and Dobzhansky, 1934, Genetics cytology: In(2LR)40B-F;58E.
Schultz, 1936, Proc. Natl. Acad. Sci. U.S. 22: 27- discoverer: Hessler, 1957.
33. references: 1958, Genetics 43: 395-403.
genetics: Variegated for bw. genetics: Variegated for It.
ln(2LR)C251: lnvershn(2LR) Crossover ln(2LR)0: lnversion(2LR) of Oster
suppressor cytology: In(2LR)30E-F;50C10-Dl superimposed on
cytology: In(2LR)36F;57B. In(2L)22D 1-2 ;33F 5-34 Al + In(2R)42A2-3;58A4-Bl
origin: X ray induced, (Lindsley).
discoverer Roberts, 1965. new order: 20 - 22Dl|33F5 - 3OF|5OD1 -
genetics: Homozygous viable. Recombination re- 58A4|42A2 - 34Al|22D2 - 30E|50C10 -
duced in 2R. 42A3|58B1 - 60.
ln(2LR)D origin: X ray induced in In(2L)Cy +In(2R)Cy,
cytology: In(2LR)36F;49B. Cy dplvIpr ct\2.
discoverer: Oshima and Watanabe. discoverer: Oster.
In(2LR)dp: see T(2;3)dp other information: Used as a balancer for chromo-
ln(2LR)Gla: lnversion(2LR) Glazed some 2, described as CyO in the section on bal-
cytology: In(2LR)27D;51E superimposed on ancers .
In(2L)22Dl-El;33F4-34A9 (Morgan, Bridges, and In(2LR)Pnt: see In(2LR)bWvi
Schultz, 1936, Carnegie Inst. Wash. Year Book 35: In(2LR)Pm»: see In(2LR)bwV3 2g
293; Grell and Grell, 1962, DIS 36: 71). *ln(2LR)pxS2g: InversionQLR) plexus
new order: 21 - 22Dl|33F5 - 27D|51E - origin: X ray induced in en crs.
34A9|22E1 - 27D|51E - 60. discoverer: Iyengar and Meyer, 52g.
origin: X ray induced in chromosome containing references: 1956, DIS 30: 73.
In(2L)Cy = In(2L)22Dl-2;33F5-34Al or In(2L)t = Meyer, 1956, DIS 30: 81.
In(2L)22D3-El;34A8- 9. 1958, DIS 32: 83.
genetics: Associated with Gla. Effective crossover genetics: Mutant for px. Pericentric inversion with
suppressor; no single or double crossovers re- breakpoints between dp and 6 and between px and
covered to the left of c (Alexander, 1952, Texas sp. Homozygous female fertile but male sterile.
Univ. Publ. 5204: 219-26). Male genitalia rotated. Sterility factor not a lie lie
ln(2LR)lt"3: lnversion(2LR) light-mottled to a6 and not covered by duplication in bw+Y, as
cytology: In(2LR)40B-F;60D. is crs, the male sterility factor present in original
origin: X ray induced. chromosome.
discoverer. Hessler, 1957. ln(2LR)Px4: lnversion(2LR) Plexate
references: 1958, Genetics 43: 395—403. cytology: In(2LR)22A3-Bl;60B-C^;21 C8-
genetics: Variegated for It. D1;6OD1-2R + In(2R)42A2-3;58A4-Bl; deficient for
*ln(2LR)lt'«9 60B-D1 and duplicated for 21D1-22A3.
cytology: In(2LR)40B-F;S6E. new order: 21A - 22A3|60B - 58B1 |42A3 -
origin: X ray induced. 58A4|42A2 - 21Dl|60D2 - 60F.
discoverer. Hessler, 1957. origin: Synthetic. This chromosome is a recurrent
references: 1958, Genetics 43: 395-403. product of recombination in region 33F-40F be-
genetics: Variegated for It. tween In(2LR)21C8-Dl;60Dl-2 from ln(2LR)bwV1
ln(2LR)ltmi2 \jn(2LR)2lC8-Dl;60Dl-2 + In(2LR)40F;59D4-El]
cytology: In(2LR)40B-F;60D. and ln(2LR)22A3-Bl;60B-C from SMI [jn(2L)Cy =
origin: X ray induced. In(2L)22Dl-2;33F5-34Al + In(2LR)22A3-Bl;60B-C +
discoverer: Hessler, 1957. In(2R)Cy =In(2R)42A2-3;S8A4-Bl]. Recombinant
references: 1958, Genetics 43: 395—403. carries tip of 2L and In(2R)Cy from SMI and tip of
genetics: Variegated for It. 2R and most of 2L from In(2LR)bwVi. The recip-
*ln(2LR)lt'*22 rocal recombinant is In{2LR)S56f.
cytology: In(2LR)40B-F;S9D. discoverer: Thompson.
origin: X ray induced. references: Burdick, 1956, DIS 30: 69.
discoverer: Hessler, 1957. genetics: Deficient for bs, 6a, Pin, Px, and probably
references: 1958, Genetics 43: 395-403. sp; duplicated for S.
genetics: Variegated for It. ln(2LR)Rev: Inversion(2LR) Revolute
*ln(2LR)lt>»2S cytology: In(2LR)40F;52Dl0-El (Bridges and Li in
cytology: ln(2LR)40B'F;57C-D. Morgan, Bridges, and Schultz, 1936, Carnegie Inst.
origin: X ray induced. Wash. Year Book 35: 293).
discoverer: Dobzhansky, 31b5. other information: Used as a balancer for chromo-

genetics: Variegated for It and .Rev. some 2, described as SMI in the section on bal-
ln(2LR)Rev&: lnversion(2LR) Revolute of Bridges ancers.
cytology. In(2LR)40;52C-E (E. B. Lewis). ln(2LR)SM5
origin: Spontaneous. cytology: In(2L)21D2-3;36C + In(2L)29C-E;40F +
discoverer: Bridges, 36e22. In(2R)42D;53C;58F superimposed on In(2L)22Dl-
synonym: In(2LR)Rvd. 2;33F5-34A1 + In(2LR)22A3-Bl;60B-C +
references: Morgan, Bridges, and Schultz, 1936, In(2R)42A2-3;58A4-Bl. Duplicated for regions
Carnegie Inst. Wash. Year Book 35: 293. 42A3-D and 58B1-F.
genetics: Mutant or variegated for Rev. new order: 21A - 21D2|36C - 4 0 F | 2 9 C -
22D2|34A1 - 36C|21D3 - 22A3|60B - 58Bl|42A3 -
ln(2LR)S56f: lnversion(2LR) Star 42D|42D - 42A3|58B1 - 58F|53C - 42D|53C -
cytology: In(2L)22Dl-2;33F5-34Al + In(2LR)21C8- 58A4|42A2 - 40F|29E - 33F5|22D1 - 22Bl|60C -
Dl;60Dl-2L22A3-Bl;60B-CR + ln(2LR)40F;59D4- 60F.
El; deficient for 21D1-22A3 and duplicated for origin: X ray induced in several steps in
60B-D1. In(2LR)SMl.
new order: 21A - 21C8|60Dl - 59El|40F - discoverer: R. F. Grell, 1955.
59D4J40F - 34Al|22Dl - 33F5|22D1 - 22Bl|60C - references: Mislove and Lewis, 1955, DIS 29: 75.
60F. genetics: Variegated for It owing to In(2L)29C-
origin: Synthetic. This chromosome is a recurrent E;40F. In(l)SM5/M(2)l lethal (C. Hinton); probably
product of recombination in region 33F-40F be- related to break in 58F.
tween In(2LR)21C8-Dl;60Dl-2 from In(2LR)bwvl other information: Excellent balancer for all of chro-
\jn(2LR)21C8-Dl;60Dl-2 + In(2LR)40F;59D4-El] mosome 2, described as SM5 in the section on bal-
and In(2LR)22A3-Bl;60B-C from SMI \jn(2L)Cy = ancers.
In(2L)22Dl-2;33FS-34Al + In(2LR)22A3-Bl ;60B-C + ln(2LR)U: lnversion(2LR) Upturned
In(2R)Cy =In(2R)42A2-3;58A4-Bl\ Recombinant origin: X ray induced.
carries the tip of 2L and In(2LR)40F;59D4-El from discoverer Ball, 32a27.
In(2LR)bwVl and the tip of 2R and In(2L)Cy from references: 1935, DIS 3: 17.
SMI. Reciprocal recombinant is In(2LR)Px4. genetics: Associated with U.
discoverer: Thompson.
references: Burdick, 1956, DIS 30: 69. ln(2R)41-47
genetics: Deficient for S; duplicated for Px. cytology: In(2R)41A;47A.
origin: X ray induced simultaneously with
ln(2LR)S325 T(l;2)B*>d.
cytology: In(2LR)21D2-3;21D3-E2;21E2-3;41. discoverer: Bridges.
new order. 21A - 21E2|41 - 21D3|41 - 60. references: Morgan, Bridges, and Schultz, 1936,
Tentative. Carnegie Inst. Wash. Year Book 35: 291.
origin: X ray induced in Dp(2;2)S = Dp(2;2)21D2- genetics: Probably not separable from T(l;2)Bbd =
3;21E2-3. T(l;2)16Al-2;48C2-3.
discoverer E. B. Lewis. *ln(2R)bw^: lnversion(2R) brown-Auburn
genetics: Break in 2L either in or between dupli- cytology: In(2R)41;59D.
cated segments of Dp(2;2)S. origin: X ray induced.
*ln(2LR)SK: lnversion(2LR) Star of Krivshenko discoverer: Dubinin.
cytology: Breakpoints near ends of 2L and 2R. synonym: In(2R)PmD1.
discoverer: Krivshenko. references: 1936, Biol. Zh. (Moscow) 5: 851-66,
references: 1936, DIS 5: 8. genetics: Variegated for bw and mi; variegation for
genetics: Associated with SK. bw dominant to bw. Dubinin claims brown-Varie-
gated effect exists at both ends of the inversion.
ln(2LR)SMh lnversion(2LR) Second Multiple other information: Ninety-one secondary rearrange-
cytology: In(2LR)22A3-Bl;60B-C superimposed on ments derived from irradiation of In(2R)bwA
In(2L)22Dl-2;33F5-34Al + In(2R)42A2-3;58A4-Bl. analyzed by Dubinin.
new order 21 - 22A3|&0B - 58B1 [42A3 - *ln(2R)bwALCyR; lnversion(2R) brown-Auburn
58A4|42A2 - 34Al]22D2 - 33F5J22D1 - Left Curly-Right
22B1 |6OC - 60F. cytology: ln(2R)4t;59DL42A2-3;$8A4~BlR; deficient
origin: X ray induced in In(2L)Cy + In(2R)Cy. for 41-42A2 and duplicated for 58B1-59D.
discoverer R. F. Grell, 1953. origin: Recombinant carrying left end of
references: 1953, DIS 27: 58. In{2R)bwA and r%ht end of In(2R)Cy.
genetics: The pericentric inversion, In(2LR)22A3- references: Dubinin, 1936, Biol. Zh. (Moscow) 5:
B1;6QB-C, enhances balancing power of In(2L)Cy + 851-66.
In(2R)Cy since it causes the single crossover be- genetics: Variegated for bw; Minute, presumably
tween the two Cy inversions to yield complemen- owing to deficiency for M(2)S2. Wings divergent
tary products that are dominant lethal. with incised inner margins.
*ln(2R)bwR32: Inversion(lR) brown-Rearranged references: 1963, DIS 38: 29.

cytology: ln(2R)41A;59D. genetics: Variegated for bw.
origin: X ray induced. *ln(2R)bwV54b
discoverer: Slatis. cytology: In(2R)41A;60D9-ll (seems unlikely that
references: 1955, Genetics 40: 5—23. right break at 60D9-11; perhaps at 59D9-11).
genetics: Associated with bwR32. origin: Neutron induced.
*ln(2R)bw*33 discoverer: Mickey, 54bl2.
cytology: In(2R)41;59D~E. references: 1963, DIS 38: 29.
origin: X ray induced. genetics: Variegated for bw.
discoverer: Slatis. *ln(2R)bwV54c
references: 1955, Genetics 40: 5—23. cytology: In(2R)41;59El.
genetics: Associated with bwR33. origin: Neutron induced.
*ln(2R)bwV2: lnversion(2R) brown-Variegated discoverer: Yanders, 54c5.
origin: X ray induced, references: Mickey, 1963, DIS 38: 29.
discoverer: Harris, 1929. genetics: Variegated for bw.
references: Muller, 1930, J. Genet. 22: 299—34. /n(2RJiwVD»': lnversion(2R) brown-Variegated of
Glass, 1933, J. Genet. 28: 69-112. Demerec
1934, Am. Naturalist 68: 107-14. cytology: In(2R)41B2-Cl;59E2-4 [Bridges, 1937,
genetics: Variegated for bw. Linkage data indicate Cytologia (Tokyo), Fujii Jub. Vol. 2: 745-55].
that one break is just to the right of centromere. origin: X ray induced.
*ln(2R)bwV7 discoverer: Demerec, 33i28.
origin: X ray induced. genetics: Variegated for bw.
discoverer: Winchester, 1932. ln(2R)bwVD°U-CyR: lnversion(2R) brown-
references: 1938, DIS 9: 23. Variegated of Demerec 1
Glass, 1939, DIS 12: 47. Left Curly-Right
geneti cs: Variegated for bw. One break at bw and cytology: In(2R)41B2-Cl;59E2-4L42A2-3;58A4-BlR;
another to the right of spindle attachment of 2R. deficient for 41C1-42A2 and duplicated for 58B1-
Gives viable recombinants with In(2R)Cy = 59E2.
ln(2R)42A2-3; S8A4-B1. origin: Recombinant carrying left end of
ln(2R)bwV3Okio In(2R)bwVDel and right end of In(2R)Cy.
origin: X ray induced. geneti cs: Deficient for M(2)S2 but not rl or M(2)p;
discoverer. Van Atta, 30klO. duplicated for M(2)l, bw, and mi.
genetics: Variegated for bw. cytology: In(2R)41A-B;59D6-El [Bridges, 1937,
ln(2R)bwV34k Cytologia (Tokyo), Fujii Jub. Vol. 2: 745-55].
cytology: In(2R)41;59E superimposed on In(2R)42A2- origin: X ray induced.
3;S8A4-B1. discoverer: Demerec, 33J14.
new order 21 - 41 |59E - 58Bl|42A3 - genetics: Variegated for bw. Left break to the right
58A4|42A2 - 4l|59E - 60. of rf and right break between bw and mi.
origin: X ray induced in In(2R)Cy.
discoverer: Oliver, 34k22. ln(2R)bwVD°2l-CyR; lnversion(2R) brown-
references: 1937, DIS 7: 19. Variegated of Demerec 2
genetics: Variegated for bw. Recombination in re- Left Curly-Right
gion 43A3-58A4 between In(2R)bwV34k+Cy and a cytology: In(2R)41A-B;59D6-El L42A2-3;58A4-B1R;
normal sequence produces reciprocal duplication- deficient for 41B-42A2 and duplicated for 58B1-
deficiency types: D%2R)bwV34*LcyL + 59D6.
Dp(2R)CyRbwV34kR = Df(2R)41;42A2*3 + origin: Recombinant carrying left end of
Dp(2R)58A4-Bl;59 (i.e., 21 - 41J59E - ln(2R)bwVD*2 and right end of In(2R)Cy.
58BI |42A3 - 60) and Dp(2R)bwV34kLCyL + genetics: Duplicated for M(2)l and bw but not mi;
D%2R)CyRbwV34kR *=Dp(2R)41;42A2-3 + deficient for M(2)S2 but not rl or M(2)p.
Df(2R)58A4-Bl;59 (i.e., 21 - 58A4J42A2 - 4l|59 - *ln(2R)bwVI; ln(2R)brown-Varlegated of Ives
60). cytology: In(2R)41A;59D.
*ln(2R)bwV40b origin: Spontaneous.
cytology: In(2R)41A-B;59D-E, discoverer Ives, 38113.
discoverer: T. Hinton, 40b. genetics: Associated with bwVI.
references: Atwood, 1942, DIS 16: 47. ln(2R)C72; lnversion(2R) Crossover suppressor
genetics: Variegated for bw. cytology: In(2R)50E;57F;60D.
new order: 21 - 5 O E | S 7 F - 60D|57F - 5OE|6OD -
cytology. In(2R)41A-B;59D4-9. 60F.
origin: Gamma ray induced, origin: X ray induced.
discoverer: Mickey, 54®6. discoverer. Roberts and D. Stewart, 1964.
genetics: Homozygous viable. Recombination be- synonym: C(2)R.

tween b and sp sharply reduced. references: Sturtevant, 1919, Carnegie Inst. Wash.
in(2R)C129 Publ. No. 278: 305-41.
cytology: In(2R)43F;56E. 1931, Carnegie Inst. Wash. Publ. No. 421: 1-27.
origin: X ray induced. genetics: Crossing over reduced to about 1.5 per-
discoverer: Roberts, 1965- cent between centromere and inversion and to
genetics: Homozygous lethal. Recombination be- about 0.1 percent between inversion and tip of
tween b and sp reduced. chromosome.
ln(2R)Cy: lnversion(2R) Curly other information: Found in many natural populations
cytology: In(2R)42A2-3;58A4-Bl (Bridges and Li in (e.g., Warters, 1944, Texas Univ. Publ. 4445: 129—
Morgan, Bridges, and Schultz, 1936, Carnegie Inst. 174; Oshima and Watanabe, 1965, DIS 40: 88).
Wash. Year Book 35: 292). In(2R)PmDi: see In(2R)bw*
origin: Spontaneous. ln(2R)PuK: lnversion(2R) Punch of Krivshenko
discoverer: L. Ward, 21f. cytology: In(2R)41;57E-F,
Sturtevant, 1931, Carnegie Inst. Wash. Publ. No. discoverer: Krivshenko, 53k24.
421: 20. synonym: In(2R)PmK.
Graubard, 1932, Genetics 17: 81-105. references: 1954, DIS 28: 75.
genetics: Left breakpoint between ap and pk. Homo- genetics: Associated with PuK (Rowan). PuK/Pu2
zygous viable and fertile. Crossing over in 2R is lethal.
strongly reduced. Carries en 2 in most laboratory ln(2R)vgu: lnversion(2R) vestigial-Ultra
stocks, cytology: In(2R)49Cl-2;50Cl-2 (Ratty and Lindsley,
other information: Used in combination with 1964, DIS 38: 30).
In(2L)Cy as a balancer for chromosome 2. origin: Gamma ray induced.
h(2R)CyLbwV>eiR: lnversion(2R) Curly-Left discoverer: Ives, 55131.
brown-Variegated of references: 1956, DIS 30: 72-73.
Demerec 7 Right genetics: Associated with vgu. Homozygous lethal.
cytology: ln(2R)42A2-3;58A4-B1^41B2-Cl;59E2-4R;
duplicated for 41C1-42A2 and deficient for 58B1- ln(3)C41: lnversion(3) Crossover suppressor
59E2. cytology: In(3)80-8l;91E-F; position of left break-
origin: Recombinant carrying left end of In(2R)Cy point with respect to centromere not determined.
and right end of In(2R)bwVD*l. origin: X ray induced.
genetics: Deficient for M(2)l, bw, and mi, duplicated discoverer: Roberts, 1964.
for M(2)S2 but not rl or M(2)p. genetics: Homozygous lethal. Recombination be-
ln(2R)Cy'-bwy'>*2R: foyersion(2R) Curly-Left tween st and ca reduced.
brown-Variegated of ln(3)C229
Demerec 2 Right cytology: In(3)67B;80-81; position of right break-
cytology: ln(2R)42A2-3;58A4-BlL41A-B;59D6-ElR; point with respect to centromere not determined.
duplicated for 41B-42A2 and deficient for 58B1- origin: X ray induced.
59D6. discoverer: Roberts, 1965.
origin: Recombinant carrying left end of ln(2R)Cy genetics: Homozygous lethal. Recombination be-
and right end of In(2R)bwVD<>2. tween ve and st sharply reduced.
geneti cs: Deficient for M(2)l and bw but not mi; ln(3)C289
duplicated for M(2yS2 but not rl or M(2)p. cytology: In(3)80-8l;93E; position of left breakpoint
ln(2R)G: lnversion(2R) Gallup with respect to centromere not determined. May
cytology: In(2R)50E;54D (T. Hint on). also contain a T(2;3).
discoverer: Ives. discoverer: Roberts, 1965.
references: 1957, DIS 31: 83. genetics: Homozygous lethal. Recombination be-
genetics: Associated with N-2G. Crossing over in tween st and ca reduced.
2R reduced to about 13 percent. *ln(3)Pl00.48; lnvcrsion(3) pink
ln(2R)M: lnversion(2R) of Mourad cytology: In(3)80-81;85A6~Bl; position of left break-
cytol ogy: In(2R)S4 Fl -55A1; 58 F-59A. point with respect to centromere not determined.
discoverer: Mourad and Mallah. discoverer: Alexander.
references: 1960, Evolution 14: 166—70. references: Ward and Alexander, 1957, Genetics 42:
ln(2R)HS: lnversion(2R) from Nova Scof/a 42-54.
cytology: In(2R)52A2-Bl;56F9-13 (Bridges and Li in genetics: Mutant for p.
Morgan, Bridges, and Schultz, 1936, Carnegie Inst. *ln(3)pi 00.88
Wash. Year Book 35: 292-3). cytology: In(3)80-81;94Dll-El; position of left
origin: Naturally occurring inversion. breakpoint with respect to centromere not deter-
discoverer: Sturtevant, 13i. mined.
origin: X ray induced. genetics: Homozygous lethal. Recombination be-

discoverer: Alexander, tween ve and st sharply reduced.
references: Ward and Alexander, 1957, Genetics 42: ln(3L)C299
42-54. cytology: In(3L)63C;80.
genetics: Mutant for p. origin: X ray induced.
discoverer: Roberts, 1965.
*ln(3L)l00.307 genetics: Homozygous lethal. Recombination be-
cytology: In(3L)62E2-4;64C2-4. tween ve and at virtually eliminated.
origin: X ray induced simultaneously with $100,307% ln(3L)C302
discoverer: Alexander. cytology: In(3L)63A;71A.
references: Ward and Alexander, 1957, Genetics 42: origin: X ray induced,
42-54. discoverer: Roberts, 1965.
*tn(3L)100r2 genetics: Homozygous lethal. Recombination be-
cytology: In(3L)76A4-Bl;80 superimposed on tween ve and st virtually eliminated.
Dp(l;3)3B4-Cl;4B4-5;80. ln(3L)D: lnversion(3L) Dichaete
new order: 61 - 76A4|(3C1 - 4B4)|80 - 76Bl|80 - cytology: In(3L)69D3-El;70C13-Dl (Bridges).
100. origin: Spontaneous.
origin: X ray induced in Dp(l;3)N264-l00m discoverer: Bridges, 15a3.
discoverer: Gersh, 1959. references: Morgan, Bridges, and Schultz, 1937,
synonym: N264-100r2. Carnegie Inst. Wash. Year Book 36: 301.
references: 1959, Genetics 44: 163-72. genetics: Associated with D.
genetics: Selected because white variegation darker ln(3L)hlO0.12: hversion(3L) hairy
than Dp(l;3yN26*-100. Removes duplication from cytology: In(3L)61A2-3;66D.
centromere region. origin: X ray induced.
ln(3L)100r8 discoverer: Alexander.
cytology: ln(3L)73Fl-74Al;80 superimposed on references: Ward and Alexander, 1957, Genetics 42:
Dp(l;3)3B4-Cl;4B4-5;80. 42-54.
new order: 61 - 73Fl|(3Cl - 4B4)|80 - 74Al|80 - genetics: Mutant for h.
100. *ln(3L)h100.239
origin: X ray induced in Dp(l;3)N264-100. cytology: In(3L)66Dll-12;80C,
discoverer: Gersh, 1959. origin: X ray induced.
genetics: Selected as a partial reversion of white discoverer: Alexander.
mottling in Dp(l;3)N264-10<>. Removes duplication references: Ward and Alexander, 1957, Genetics 42:
from centromere region. 42-54.
*h(3L)100rn genetics: Mutant for h.
cytology: ln(3L)65Al-Bl;80 superimposed on *ln(3L)M: lnversion(3L) of Mourad
Dp(l;3)3B4-Cl;4B4-5;80. cytology: In(3L)66D;71D.
new order, 61 - 65Al|(3Cl - 4B4)|80 - 65Bl|80 - origin: Spontaneous.
100. discoverer: Mourad and Mallah.
origin: X ray induced in DtfltfW264~l00- synonym: ln(3L)F =In(3L)66C;71B of Oshima and
discoverer: Gersh, 1959. Watanabe (1965, DIS 40: 88) probably the same.
references: 1959, Genetics 44: 163—72. references: 1960, Evolution 14: 166—70.
genetics: Selected as a partial reversion from white- ln(3L)P; lnversion(3L) of Payne
mottled. Removes duplication from centromere re- cytology: In(3L)63C;72El-2 (Bridges and Li in
gion. Morgan, Bridges, and Schultz, 1937, Carnegie Inst.
*ln(3L)Apt: lnversion(3L) Apart Wash. Year Book 36: 301).
cytology: Breakpoints unknown. origin: Naturally occurring inversion.
origin: X ray induced. discoverer: Payne, 17g.
discoverer: Belgovsky, 34e23. references: 1918, Indiana Univ. Studies 5 No. 36:
references: 1935, DIS 3: 27. 1-45.
genetics: Associated with Apt. 1924, Genetics 9: 327-42.
Sturtevant, 1931, Carnegie Inst. Wash. Publ. No.
*ln(3L)Bit: lnversion(3L) Bitten 421: 18.
cytology: Breakpoints unknown. genetics: Homozygous viable, although it often con-
origin: X ray induced. tains lethals of independent origin.
dlscovere*: Lefevre, 48g5. other information: Often associated with ln(3R}P.
references: 1949, DIS 23: 5E. Much used as a balancer for 3L. Allows only about
genetics: Associated with Bit. 0.02 percent crossing over between m and st. Bal-
ln(3L)C90: lnversion(3L) Crossover suppressor ancers contain recessive lethals or Af e. Balancers
cytology: tn(3L)62B;$0C. for all of chromosome 3 made by combining Jn(3L)P
origin; X ray induced, with Jn(3R)P or ln{3R)C. Found in many wild pop-
discoverer: Roberts and D. Stewart, 1964. ulations (e.g., Waiters, 1944, Texas Univ. Publ.
4445: 129—174; Oshima and Watanabe, 1965, DIS genetics: Partial reversion of white-mottled to wild
40: 88). type. Duplication not removed from region of cen-
ln(3L)pers: lnversion(3L) persimmon tromere.
cytology: In(3L)63C2-5;73B2-5. ln(3LR)100r27
origin: X ray induced. cytology: In(3LR)80;96B3-5 superimposed on
discoverer: Demerec, 3712. Dp(l;3)3B4-Cl;4B4-5;80.
references: 1941, OIS 14: 40. new order: 61 - 8O|(3C1 - 4B4)|96B3 - 80|96B5 -
genetics: Associated with pers. 100.
origin: X ray induced in Dp(l;3)N264-XO0,
*ln(3LJ$pr: lnversion(3L) Spread discoverer: Gersh.
cytology: Breakpoints unknown. references: 1959, Genetics 44: 163-72.
origin: X ray induced. genetics: Almost complete reversion of the white-
discoverer: Oliver, 32k21. mottled to wild type. Break between duplication
references: 1935, DIS 4: 15. and centromere.
genetics: Associated with Spr.
ln(3LR)C35: lnversion(3LR) Crossover suppressor
*ln(3L)th'00.293: lnvershn(3L) thread cytology: In(3LR)64B;89E.
cytology: In(3L)72A2-Bl;76A4-Bl;79A4-Bl. origin: X ray induced.
new order: 61 - 72A2[79A4 - 76Bl|72Bl - discoverer: Roberts, 1964.
76A4|79B1 - 100. genetics: Homozygous lethal. Recombination prac-
origin: X ray induced. tically eliminated between ve and st and reduced
discoverer: Alexander, between s< and ca.
synonym: Tp(3)(h100-293. ln(3LR)C1J7
references: Ward and Alexander, 1957, Genetics 42: cytology: In(3LR)64D;89B.
42-54. origin: X ray induced.
genetics: Mutant for th. discoverer: Roberts, 1965.
genetics: Homozygous lethal. Recombination prac-
ln(3LR)65;S3 tically eliminated between ve and s£.
origin: X ray induced simultaneously with, but sub- h(3LR)C165
sequently separated from, T(2;3)Sbv. cytology: In(3LR)64C;83C.
discoverer: E. B. Lewis, 1948. origin: X ray induced.
references: 1956, DIS 30: 76-77. disocverer: Roberts, 1965.
ln(3LR)100rl genetics: Homozygous viable. Recombination be-
cytology: In(3LR)80;96Bl-3 superimposed on tween ve and st virtually eliminated.
Dp(l;3)3B4-Cl;4B4-5;80.
new order 61 - 8O|(3C1 - 4B4)|96B1 - 80[96B3 - ln(3LR)C17S
100. cytology: ln(3LR)65C;95E.
origin: X ray induced in Dp(l;3yN264-100. origin: X ray induced.
discoverer: Gersh. discoverer: Roberts, 1965.
references: 1959, Genetics 44: 163—72. genetics: Homozygous lethal. Recombination prac-
genetics: Selected as an almost complete reversion tically eliminated between ve and st and between
to wild type of the white-mottled effect of st and ca.
Dp(l;3)N26'*-^00. Duplication removed from region ln(3LR)C190
of the centromere. cytology: In(3LR)69F;89D.
*ln(3LR)100r3 origin: X ray induced,
cytology: In(3LR)80;86Cl-Dl superimposed on discoverer: Roberts, 1965.
Dp(l;3)3B4'Cl;4B4-5;80. genetics: Homozygous lethal. Recombination re-
new order: 61 - 8O|86C1 - 8OJ(3C1 - 4B4)J86D1 - duced between ve and st and between st and ca.
100. ln(3LR)C269
origin: X ray induced in Dp(l;3yN264-10Q, cytology: In(3LR)78C;98F.
discoverer: Gersh. origin: X ray induced.
references: 1959, Genetics 44: 163—72. discoverer: Roberts, 1965.
genetics: Does not remove duplication from region genetics: Homozygous viable. Recombination be-
of centromere. tween st and ca virtually eliminated.
*ln(3LR)100r7 h(3LR)C334
cytology: In(3LR)8Q;99B-Cl superimposed on cytology- ln(3LR)67B;88D;91F.
Dpfl; 3)3 B4- Cl;4B4-5;80. new order: 61 - 67E|88D - 67EJ91F - 88D|91F -
r»«w order: 61 - 80|99B - 8OJ(3C1 - 4B4)|99€1 - 100.
©rifin: X ray induced in Dp(t;3pf264~10®> discoverer: Roberts, 1965.
S s«©v••m-er. Germfe, genetics: Hoosozygous lethal. Recombination re-
1959, Genetics 44: 163—72. duced between ve and at and between &t and ca.
ln(3LR)CxD: lnversion(3LR) Crossover references: 1963, DIS 38: 29.

Suppressor Dichaete genetics: Mutant or deficient for M(3)54c and st.
cytology: In(3LR)71F;85C +In(3LR)80;84A;93F ln(3LR)P88
superimposed on In(3L)69D3-El;70C13-Dl. cytology: In(3LR)61A;89C-D; deficient for bands in
new order: 61A - 69D3|70C13 - 69El|70Dl - 89C-D.
71F|85C - 84A|80 - 84A|93F - 85C|71F - origin: X ray induced,
80|93F - 100 (Bridges). discoverer: E. B. Lewis, 55h.
origin: X ray induced in In(3L)D. genetics: Deficient for ss but not bx.
discoverer: Oliver. ln(3LR)sep: lnversion(3LR) separated
synonym: CxD; Dcx. cytology: In(3LR)65E;85E (Lewis, 1951, DIS 25:
references: Glass, 1933, J. Genet. 28: 70. 108-9).
Federova, 1937, Dokl. Acad. Nauk SSSR 14: 135- discoverer: Muller.
38. genetics: Mutant for sep. Also carries ri and pp,
genetics: Carries D (separates from other inversions which can be removed only with great difficulty.
with frequency of 0.2 percent). Crossing over
strongly reduced in chromosome 3 except distal ln(3LR)mi: lnversion(3LR) Third Multiple
half of 3L; virtually no crossing over between st cytology: In(3L)63C;72El-2 + In(3LR)69E;9lC +
and e. In(3R)89B;97D.
other information: Name easily confused with what new order: 61 - 63C|72E1 - 69E|91C - 97D|89B -
has been called C(3)x, which appears to be 72E2|63C - 69E|91C - 89B|97D - 100.
In(3L)P +In(3R)P (Lewis, 1956, DIS 30: 130). origin: Derived from T(2;3JMe/ri, presumably by a
ln(3LR)DcxF: lnversion(3LR) Dichaete crossover double crossover with exchanges in regions 72E2-
suppressor of Federova 80 and 81-89B, which replaced the T(2;3) break-
cytology: In(3L)62;67 superimposed on In(3L)69D3- point in 3 with ri.
El;70C13-Dl + In(3LR)71F;85C + discoverer: E. B. Lewis.
In(3LR)80;84A;93F. From Federova's drawings references: 1949, DIS 23: 92.
(1937), there appears to be an inversion from about 1953, DIS 27: 58.
62 to 67 in addition to a complex rearrangement, genetics: Carries Me, ri, and sbd1.
presumably In(3LR)CxD. other information: Used as a balancer for chromo-
new order: 61 - 62J67 - 62|67 - 69D3|70C13 - some 3, described as TM1 in the section on bal-
69El|70Dl - 71F|85C - 84A|80 - 84A|93F - ancers.
85C|71F - 80|93F - 100. ln(3LR)TM3
origin: X ray induced in In(3LR)CxD. cytology: Jn(3LR)71C;94D-F + In(3LR)76C;93A +
discoverer: Fedorova. In(3LR)79E;100C superimposed on XD3P from
synonym: In(3LR)CxF; DcxF; CxF.D. T(l;3)lA8-Bl;61Al-2 + In(3LR)65E;85E +
references: 1937, Dokl. Acad. Nauk SSSR 14: 135- In(3R)92Dl-El;100F2-3.
38. new order: 1A1 - 1A8|61A2 - 65E|85E -
genetics: Carries I>. Crossing over strongly in- 79E|l00C - 1OOF2|92D1 - 85E|65E - 7lC|94D -
hibited throughout chromosome 3. 93A|76C - 71C|94F - 1OOC|79E - 76C|93A -
*ln(3LR)Hh lnversion(3LR) Hirsute 92El|l00F3 - 100F5.
cytology: In(3LR)71A;91F. origin: Induced by repeated irradiation of the XD3P
origin: X ray induced. element of T(l;3)sc260-2 0i which carried
discoverer. Bishop, 1939. bi(3LR)sep + In(3R)C, y+ ri p sep bx34e e «.
genetics: Associated with Hi. discoverer E. B. Lewis.
*ln(3LR)K: Inversion(JLR) of Krivshenko references: Mitchell, 1958, Cold Spring Harbor
cytology: ln(3LR)6lC6-7;100A-B; only the left end Symp. Quant. Biol. 23: 279-90.
recovered. Lewis, 1960, DIS 34: 51.
new order: 100F - 100B|61C7 - 100. other information: Used as a balancer for chromo-
origin: X ray induced in oocytes. some 3, described as TM3 in the section on bal-
genetics: Result of a pericentric inversion followed ancers.
by an exchange or of a trans location between 3L of
one chroma tid and 31? of its sister or homolog. cytology: In(3LR)74;94 superimposed on
ln(3LR)M-54c: lnversion(3LR) Minute-54c In(3L)63C;72El-2 + In(3LR)61A;89CD +
cytology: In(3L)73A9-1O;75D7-E1 + In(3LR)61C2- ln(3R)92Dl-El;tOOF2-3.
3;80C4-5;93B4-5;100B8-9. new order: 61AJ89C - 74|94 - 100F2|92Dl -
new mdm: 61A - 61C2|93B5 - lQOBSlSOCS - 89D|61A - 63C|72E1 - 63CJ72E2 - 74|94 -
93B4J80C4 - 75El|73A10 - 75D7|73A9 - | - | - 92E1J100F3 - 100F5.
61C3J100B9 - 100F. Also carries ®n inversion origin: X ray induced in ln(3L)P + In(3LR)P88 +
with unspecified breakpoints in the region between In{3R)C, bx^4® e.
61C3 and 73A9. discoverer E. B. Lewis and F. Bacher, 66i.
©rigim Neutron induced. SJienetics: Homoanygote lethal. Deficiency for •* but
discoverer: Mickey, 54clO. not bx associated with In(3LR)P88-
other information: Used as a balancer for chromo- ln(3R)C

some 3. Described as TM6 in section on balancers. cytology: In(3R)92Dl-El;100F2-3 (Bridges and Li in
ln(3LR)Ubxl0i: !nversion(3LR) Ultrabithorax Morgan, Bridges, and Schultz, 1937, Carnegie Inst.
cytology: In(3LR)80;89D9-El. Wash. Year Book 36: 301).
origin: X ray induced. origin: Naturally occurring inversion.
discoverer: E. B. Lewis, 1947. discoverer: Sturtevant, 13f.
references: 1949, DIS 23: 59. synonym: C3; CIIIRE; In(3R)E.
genetics: Mutant for Ubx. references: 1913, Science 37: 990-92.
ln(3LR)Ubxi30 1917, Proc. Natl. Acad. Sci. U.S. 3: 555-58.
cytology: In(3LR)61A- C; 74;89D-E; 93B;96A. 1926, Biol. Zentr. 46: 697-702.
new order: 61A|96A - 93B|89D - 74|61C - 1931, Carnegie Inst. Wash, Publ. No. 421: 1-27.
74|89E - 93B|96A - 100. Muller, 1918, Genetics 3: 422-99.
origin: X ray induced in e s . genetics: Homozygous viable. Crossing over in 3R
discoverer: E. B. Lewis. reduced to 1 percent between centromere and ss, to
references: 1952, Proc. Natl. Acad. Sci. U.S. 38: 0.2 percent between ss and e; no crossovers be-
955-60. tween e and tip of 3R recovered except for rare
1952, DIS 26: 66. doubles within inversion.
genetics: Mutant for Ubx; homozygous lethal. Also other information: First inversion demonstrated
carries e°. genetically (Sturtevant, 1926). Used as a balancer
other information: A useful balancer for chromosome for the region from Dl to 3R tip. Balancers contain
3, described as TM2 in the section on balancers. Sb, e, l(3)a, or l(3)e. Balancer for all of chromo-
ln(3LR)UbxA some 3 made by combining with In(3L)P. Found in
origin: X ray induced. wild populations (e.g., Oshima and Watanabe,
discoverer: Schalet, 1959. 1965, DIS 40: 88).
references: 1960, DIS 34: 53, 55.
genetics: Mutant for Ubx; homozygous lethal. One ln(3R)C133: lnversion(3R) Crossover suppressor
breakpoint in 3L between h and st and another left cytology: In(3R)93F;97C-D2.
of e (probably at Ubx). origin: X ray induced,
*ln(3R)300.96 genetics: Homozygote rarely survives. Recombina-
cytology: In(3R)89F2-90Al;99B2-4. tion between st and ca sharply reduced.
origin: X ray induced simultaneously with e300.96. ln(3R)C208
discoverer: Alexander. cytology: In(3R)91B;96B.
references: Ward and Alexander, 1957, Genetics 42: origin: X ray induced.
42-54. discoverer: Roberts, 1965.
genetics: Carries an independent mutant for e. genetics: Homozygous viable with wings held at 45°
Homozygous viable but male sterile. angle from body axis.
ln(3R)cav; lnversion(3R) claret-variegated
ln(3R)Antp&: lnversion(3R) Antennapedia of cytology: In(3R)81F;99C-E.
Bacon origin: X ray induced.
cytology: In(3R)84A;85E, discoverer: E. B. Lewis.
origin: X ray induced. genetics: Variegates for ca.
discoverer: Bacon, 50g. ln(3R)Cyd: lnversion(3R) Curlyoid
references: Lewis, 1956, DIS 30: 76. discoverer: Jollos.
genetics: Mutant for Antp. references: Curry, 1939, DIS 12: 46.
genetics: Associated with Cyd.
*ln(3R)AntpLC: lnversion(3R) Antennapedia o/ other information: May be In(3R)P.
Le Calvez ln(3R)DlB: lnversion(3R) Delta-Barish
cytology: In(3R)84A5-6;92A5-6. cytology: In(3R)90A;91A (Schultz).
origin: Neutron induced. discoverer Schultz, 1933.
discoverer: Le Calvez. genetics: Mutant for Dl.
references: 1948, Bull. Biol. France Belg. 82: 97— *ln(3R)elO0.265: hversion(3R) ebony
113 (fig.), cytology. ln(3R)93B5-6;95E.
genetics: Associated with AntpLC. origin: X ray induced.
discoverer Alexander.
ln(3R)AntpR: lnversion(3R) Antennapedia of references: Ward and Alexander, 1957, Genetics 42:
Rappaport 42-54.
cytology: In(3R)83F;86C (Ben-Zeev). genetics: Mutant for e; homozygous viable.
origin: X ray induced. In(3R)E: see In(3R)C
discoverer: Rappaport, 1963. ln(3R)hp: lnversion(3R) humped
references: Falfc, 1964, DB 39: 60. cytology: Breakpoints unknown.
genetics: Associated with AntpR. origin: Spontaneous.
discoverer: Bridges, 31a22. *ln(3R)pl00.290: inversion(3R) pink

genetics: Associated with hp. cytology: In(3R)85B3-4;85D12-15.
ln(3R)Hu: lnversion(3R) Humeral origin: X ray induced.
cytology: In(3R)84B2-3;84F2-3;86B4-Cl. discoverer: Alexander.
new order: 61 - 84B2|84F2 - 84B3|86B4 - references: Ward and Alexander, 1957, Genetics 42:
84F3|86C1 - 100. 42-54.
origin: X ray induced, genetics: Mutant for p.
discoverer: Ruch, 1931. ln(3R)P: lnversion(3R) of Payne
genetics: Associated with Hu. cytology: In(3R)89C2-3;96A18-19 [Bridges and
,ln(3R)J Bridges, 1938, Genetics 23: 111-14 (fig.)J.
cytology: In(3R)96E;98F. origin: Widespread in natural populations.
origin: Naturally occurring inversion. discoverer: Payne, 17g.
discoverer: Oshima and Watanabe. references: 1918, Indiana Univ. Studies 5, No. 36:
references: 1965, DIS 40: 88. 1-45.
*ln(3R)K: Inversion(3R) of Kodani 1924, Genetics 9: 327-42.
cytology: In(3R)86Fl-87Al;96Fll-97Al. Sturtevant, 1931, Carnegie Inst. Wash. Publ. No.
origin: Spontaneous. 421: 1-27.
discoverer: Kodani. genetics: Crossing over reduced in heterozygous
*ln(3R)M: lnversion(3R) of Mourad female to 1 percent between p and sr; none be-
cytology: Jn(3R)86F,-100E. tween sr and to; 0.5 percent between to and ca.
origin: Spontaneous. other information; Widespread in laboratory stocks
discoverer: Mourad and Mallah. and is part of the balancers, LVM and C(3)x. Also
references: 1960, Evolution 14: 166—70. found in many wild populations (e.g., Warters,
ln(3R)Mo: lnversion(3R) from Missouri 1944, Texas Univ. Publ. 4445: 129-74; Oshima
cytology: In(3R)93D;98F2-3 (Bridges and Li in and Watanabe, 1965, DIS 40: 88).
Morgan, Bridges, and Schultz, 1936, Carnegie Inst. *ln(3R)sr*2: lnversion(3R) stripe
Wash. Year Book 35: 293). cytology: ln(3R)90Dl-El;93B-E.
origin: Naturally occurring inversion. origin: X ray induced.
discoverer Sturtevant, 1924. discoverer Alexander, 1959.
references: 1931, Carnegie Inst. Wash. Publ. No. references: 1960, Genetics 45: 1019-22.
421: 6-7. genetics: Mutant for sr.
genetics: Crossing over reduced in heterozygote to
about 5 percent between centromere and sr and 0.3 *ln(3R)su(pr): lnversion(3R) suppressor of purple
percent between sr and ca. cytology: Breakpoints unknown,
other information: Pound in natural populations origin: Spontaneous.
(e.g., Waiters, 1944, Texas Univ. Publ. 4445: discoverer: Stern, 27c2.
129-74; Oshima and Watanabe, 1965, DIS 40: 88). synonym: su^-pr.
ln(3R)Msc: lnversion(3R) Multiple sex comb references: 1929, Z. Induktive Abstammungs-
cytology: ln(3R)84B;84F. Vererbungslehre 52: 373-89.
origin: Spontaneous, 1934, DIS 1: 35.
discoverer. Tokunaga, 64a. genetics: Associated with su(pr).
references: 1966, DIS 41: 57. In(3R)Vna: see Tp(3)Vno
genetics: Associated with Msc. *ln(3R)W: lnversion(3R) of barters
ln(3R)Na: lnversion(3R) from Naples cytology: In(3R)86B;92F.
cytology: In(3R)86F2>3;96Flt-97Al;97A2-5. 97A1-2 origin: Naturally occurring inversion.
missing. discoverer: Warters.
new order: 61 - 86F2J96F11 - 86F3J97A5 - 100. references: 1944, Texas Univ. Publ. 4445: 129-74.
origin: Spontaneous. lnp(l)&cvi": see In(lLR)scVi
discoverer: Carfagna and Nicoletti, 1960. In&( ) : see In( )
references: 1963, DIS 38: 32. Ins(2LR)Pm: see
genetics: Carries a lethal, which may be separable
from inversion or the deficiency for 97A1-2 may be RINGS
the lethal.
other information: Breakpoints similar to those of R(l)h Ring(l)
Im(3R)K »In(3R)86Fl-$7Al;96FlI~97Al and may cytology: R(1)1A;2OB-C; salivary chromosomes
be the same. show deficiency for roost of 1A and a duplication
*ln(3R)Nel-D: lnYmsion(3R) of Nel for 20C-D [Schultz and Catcheside, 1937, J. Genet.
cytology: tn{3R)86D;97A. 35: 315-20 (fig.)]. Ring shaped in
origin: Spontaneous in natural population. new order: |lA - 20-20F - 20CJ.
discoverer: Nel. origin: Spontaneous from C(1)RM, y female.
other information: Possibly the same as In(3R)K « discoverer: L. V. Morgan, 1922.
In(3R)S6 F1-87A1;96FU'97AJ. synonym: X°; X&.
CHROMOSOME ABERRATIONS - RINGS 347
references: 1926, Proc. Natl. Acad. Sci. U.S. 12: a small segment, the centric constriction, a small
180-81. segment.
1933, Genetics 18: 250-83. origin: Regular product of exchange in C(l)TMBs9-4.
genetics: Carries y. Male and homozygous female discoverer: Lindsley and Sandier, 1963.
have reduced viability; X/0 male lethal (Schultz, references: 1965, Genetics 51: 223—45 (fig.).
1941, Proc. Intern. Congr. Genet., 7th. pp. 2 5 7 - genetics: Carries y. R(l)9-4/0 male viable. Based
62). Somewhat unstable, tending to be eliminated on origin, R(l)9-4 euchromatically but not hetero-
during mitosis. Shows about five times as much chromatically indentical with R(l)l.
somatic crossing over as rod X (Brown, Walen, and R(J)63
Brosseau, 1962, Genetics 47: 1573—79). Crossing cytology: Ring shaped in mitotic figures. Early pro-
over reduced in ring/rod heterozygote; only double phase shows heterochromatic constitution, pro-
crossovers recovered. Exceptional males result ceeding from the normally proximal euchromatin,
from four-strand double crossing over in R(l)l/+ across the centromere to the normally distal eu-
female. chromatin, to be as follows: two large segments
other information: Tends to open out into a rod separated by an ill-defined constriction, a constric-
[e.g., In(l)EN\ spontaneously in stock. tion, a small segment, the centric constriction, a
R0)2 small segment.
cytology: R(1)1A3-4;19F-2OA1; salivary chromo- origin: Regular product of exchange in C(1)TM2.
somes deficient for 1A1-3 and duplicated for all of discoverer: Lindsley and Sandier, 63g.
region 20 [Schultz and Catcheside, 1937, J. Genet. references: 1965, Genetics 51: 223—45 (fig.).
35: 315—20 (fig.)]. Ring shaped in metaphase. genetics: Carries y. R(l)63/0 male survives. Based
new order: J1A4 - 20-20F - 20Al|. on origin, R(l)63 is euchromatically but not hetero-
origin: Spontaneous as a detachment of C(1)RM, y + . chromatically identical with R(l)l.
discoverer: Beadle, 34b (ring nature discovered by R(J)94-2A1
Boche). cytology: R(1)1A;1F-2A;5E-6A;17F-18A;2O; dupli-
synonym: X°2. cated for 1A-F and 18A-20.
genetics: Carries y+. More viable than R(l)l; X/0 new order |lA - 5 E | l F - 1A|2O-2O - 6A|l8A - 20|.
male survives. Ordinarily, ring elimination less origin: Spontaneous product of C(1)94-2A. Possibly
than 1 percent (Battacharya, 1950, Proc. Roy. Soc. a product of breakage of double second-anaphase
Edinburgh, B 64: 199—215; Braver and Blount, bridge formed by exchange between the arms of the
1950, Genetics 35: 98), but nearly 20 percent of compound.
the first progeny of 11-day-old females crossed to discoverer: Armentrout, 1964.
ring-bearing males are gynandromorphs (Hannah, *R(1)CJ
1955, Z. Induktive Abstammungs- Vererbungslehre cytology: Ring shaped in mitotic figures.
86: 600—21). Crossing over reduced in ring/rod origin: Spontaneous derivative of In(l)sc8LENR;
heterozygote; only double crossovers recovered. arose by recombination between distal heterochro-
Exceptional males result from four-strand double matic segment of In(l)sc8 and heterochromatic
exchange in R(l)2/+ female. short arm of In(l)EN.
other information: Ring may open out spontaneously discoverer: Lindsley, 1950.
in stock; e.g., ln(l)EN2. references: 1958, Z. Vererbungslehre: 89: 103—22.
genetics: Carries y. On basis of origin, R(1)C1 is
euchromatically identical with R(l)l, but it must
cytology: Ring shaped in mitotic figures. Early pro- be different heterochromatically since R(l)Cl/0
phase shows heterochromatic constitution, pro- male viable.
ceeding from normally proximal euchromatin, across R(l)l-v459
the centromere to the normally distal euchromatin, cytology: R(1)3D-F.
to be as follows: a large segment, a well-defined origin: Associated with T(l;2;3)l-v459.
constriction, a large segment, a constriction, a R(l)y*: RingCI) yellow
small segment, the centric constriction, a small cytology. R(l)lA8-Bl;18A3-4; deficient for 1A and
segment. duplicated for 18—20.
origin: Regular product of exchange in C(1)TMBS9-1. new order | l B l - 20-20 - 18A4|.
discoverer: Lindsley and Sandier, 1963. origin: Regular product of exchange within inversion
references: 1965, Genetics 51: 223-45 (fig.). in C(1)RM heterozygous for In(l)y4 * In(l)lA8-
genetics: Carries y. R(l)9-l/0 male survives. On Bl;18A3-4.
basis of origin, R(l)9-1 is euchromatically but not discoverer Sturtevant and Beadle,
heterochromatically identical with R(l)l. references: 1936, Genetics 21: 554—604.
R(l)9-4 Novitski and Sandier, 1956, Genetics 41: 194-206.
cytology: Ring shaped in mitotic figures. Early pro- genetics: Deficient for l(l)J 1, duplicated for car-bb.
phase shows heterochromatic constitution, pro- Heterozygous female survives; male lethal, owing
ceeding from normally proximal euchromatin, across to deficiency for 1(1}] 1.
the centromere to the normally distal euchromatin R(Y): see Y Derivatives in Special Chromosomes
to be as follows: a large segment, a constriction, section
genetics: Male viable and fertile with or without a

TRANSLOCATIONS free F.
*T(1;Y)9
*T(1;?)sc260-23: Translocationd;?) scute cytology: T(1;Y)2C;19F;YS.
cytology: T(l;?)lB2-3; position of second break not new order: 1A - 2C|YSP - Y^;
determined. 20 - 19F|2C - 19F|Y SD .
origin: X ray induced, origin: X ray induced in y/y*Y sperm.
discoverer, button, 1939. discoverer: Nicoletti.
references: 1943, Genetics 28: 210—17. references: Nicoletti and Lindsley, 1960, Genetics
genetics: Mutant for sc but not y or svr. 45: 1705-22.
1960, DIS 34: 95-97.
*T(1;Y)1 genetics: Male viable but sterile.
cytology: T(1;Y)16F;YL.
origin: X ray induced in y + F. T(l;Y)10
discoverer: Nicoletti. cytology: T(1;Y)3E;YL.
references: Nicoletti and Lindsley, 1960, Genetics origin: X ray induced in y/y+Y.
45: 1705-22. discoverer: Nicoletti.
1960, DIS 34: 95-97. references: Nicoletti and Lindsley, 1960, Genetics
genetics: Male viable but sterile. 45: 1705-22.
T(1;Y)2 1960, DIS 34: 95-97.
cytology: T(1;Y)5E;11F;19F;YS. genetics: Male viable; fertile with a free F but
new order: 1 - 5E|YSP - YL; sterile without.
20 - 19F|llF - 5E|l9F - 11F|Y SD .
origin: X ray induced in y+Y. cytology: T(1;Y)19F;YS.
discoverer: Nicoletti. origin: X ray induced in y+Y.
references: Nicoletti and Lindsley, 1960, Genetics discoverer: Nicoletti.
45: 1705-22. references: Nicoletti and Lindsley, 1960, Genetics
1960, DIS 34: 95-97. 45: 1705-22.
genetics: Male viable and fertile with or without a 1960, DIS 34: 95-97.
free F. genetics: Male viable; fertile with a free F but
T(1;Y}3 stecile without.
cytology: T(1;Y)3E;YS.
origin: X ray induced in y/y+Y sperm. *T(J;Y)U
discoverer: Nicoletti. cytology: T(1;Y)7D;YL.
references: Nicoletti and Lindsley, 1960, Genetics origin: X ray induced in y+Y.
genetics: Male lethal. 45: 1705-22.
T(1;Y)4 1960, DIS 34: 95-97.
cytology: T(l;Y)llA;YL. genetics: Male lethal.
origin: X ray induced in y+Y. T(1;Y)U
discoverer Nicoletti. cytology: T(1;Y)19F;YS.
references: Nicoletti and Lindsley, 1960, Genetics origin: X ray induced in y+Y.
genetics: Male viable and fertile with or without a 45: 1705-22.
free F. I960, DIS 34: 95-97.
T(1;Y)6 genetics: Male viable; fertile with a free F but
cytology: T(1;Y}11D;YS. sterile without.
origin: X ray induced in y+Y.
discoverer: Nicoletti. *T(1;Y)15
references: Nicoletti and Lindsley, 1960, Genetics cytology: T(1;Y)14F;Y^.
45: 1705-22. origin: X ray induced in y+Y.
I960, DIS 34: 95-97. discoverer: Nicoletti.
genetics: Male lethal. references: Nicoletti and Lindsley, 1960, Genetics
*T(1;Y)8 45: 1705-22.
cytology: T(1;Y)4B;YL. 1960, DIS 34: 95-97.
origin: X ray induced in y*Y. genetics: Male lethal.
discoverer: Nicoletti. T(1;Y)J6
references: Nicoletti and Lindsley, 1960, Genetics cytology. T(1;Y)4C;YL.
45: 1706-22. origin: X ray induced in y + F.
, DIS 34: 95-97. discoverer: Nicoletti.
CHROMOSOME ABERRATIONS - TRANSLOCATIONS 349
references: Nicoletti and Lindsley, 1960, Genetics genetics: Male viable; fertile with a free Y but
45: 1705-22. sterile without.
1960, DIS 34: 95-97. T(l;Y)102
genetics: Male lethal. cytology: T(1;Y)7D;Y^.
*T(1;Y)18 origin: X ray induced in BSY.
cytology. T(1;Y)19F;YS. discoverer: Nicoletti.
origin: X ray induced in y+Y. references: Nicoletti and Lindsley, 1960, Genetics
discoverer: Nicoletti. 45: 1705-22.
references: Nicoletti and Lindsley, 1960, Genetics I960, DIS 34: 95-97.
45: 1705-22. genetics: Male viable; fertile with a free Y but
1960, DIS 34: 95-97. sterile without.
genetics: Male viable; fertile with a free Y but
sterile without. T(l;Y)103
T(1;Y)19 cytology: T(1;Y)19F;YS.
L
cytology: T(1;Y)17A;Y . origin: X ray induced in BSY.
origin: X ray induced in y+Y. discoverer: Nicoletti.
discoverer: Nicoletti. references: Nicoletti and Lindsley, 1960, Genetics
references: Nicoletti and Lindsley, 1960, Genetics 45: 1705-22.
45: 1705-22. 1960, DIS 34: 95-97.
1960, DIS 34: 95-97. genetics: Male lethal.
genetics: Male viable; fertile with a free Y but T(1;Y)W4
sterile without. cytology: T(1;Y)3D;YL.
T(l;Y)20 origin: X ray induced in y/BsY sperm,
cytology: T(1;Y)11A,YL. discoverer Nicoletti.
origin: X ray induced in y + Y. references: Nicoletti and Lindsley, 1960, Genetics
references: Nicoletti and Lindsley, 1960, Genetics 1960, DIS 34: 95-97.
45: 1705-22. genetics: Male viable and fertile with or without a
1960, DIS 34: 95-97. free Y.
genetics: Male viable but sterile.
T(J;Y)2J T(l;Y)105
cytology: T(1;Y)1F;Y^. cytology: T(1;Y)19F;YS.
origin: X ray induced in y/y+Y sperm. origin: X ray induced in BSY.
discoverer: Nicoletti. discoverer Nicoletti.
references: Nicoletti and Lindsley, I960, Genetics references: Nicoletti and Lindsley, 1960, Genetics
45: 1705-22. 45: 1705-22.
1960, DIS 34: 95-97. 1960, DIS 34: 95-97.
genetics: Male viable but sterile. genetics: Male viable; fertile with a free Y but
T(1;Y)22 sterile without.
cytology: T(1;Y)19E;YS. T(l;Y)106
origin: X ray induced in y*Y. cytology: T(1;Y)16A;YL.
discoverer: Nicoletti. origin: X ray induced in BSY.
references: Nicoletti and Lindsley, I960, Genetics discoverer Nicoletti.
1960, DIS 34: 95-97. 45: 1705-22.
genetics: Male viable; fertile with a free Y but 1960, DIS 34: 95-97,
sterile without. genetics: Male viable; fertile with a free Y but
*T(1;Y)WO sterile without.
cytology: T(1;Y)13F;YS.
origin: X ray induced in BSY. T(l;Y)107
discoverer: Nicoletti. cytology: T(l;Y)3C;Y^,
references: Nicoletti and Lindsley, 1960, Genetics origin: X ray induced in y/BsY sperm.
genetics: Male lethal. 45: 1705-22.
T(1;Y)1OJ 1960, DIS 34: 95-97.
cytology: T(1;Y)19E;YS. genetics: Male viable; fertile with a free F but
origin: X ray induced in BSY. sterile without.
discoverer: Nicoletti. T(l;Y)108
references: Nicoletti and Lindsley, 1960, Genetics cytology: T(1;Y)5D;YL.
45: 1705-22. origin: X ray indticed in BSY.
1960, DIS 34: 95-97. discoverer: Nicoletti.
references: Nicoletti and Lindsley, 1960, Genetics genetics: Male viable; fertile with a free Y but
45: 1705-22. sterile without.
1960, DIS 34: 95-97.
genetics: Male lethal. cytology: T(1;Y)19F;YS.
tt7;Y)I77 origin: X ray induced in BSY.
cytology: T(1;Y)3C;YL. discoverer: Nicoletti.
origin: X ray induced in y/BsY sperm. references: Nicoletti and Lindsley, 1960, Genetics
references: Nicoletti and Lindsley, 1960, Genetics 1960, DIS &4: 95-97.
45: 1705-22. genetics: Male viable; fertile with a free Y but
1960, DIS 34: 95-97. sterile without.
genetics: Male lethal. Variegates for w and Af.
cytology: T(1;Y)15A;YL. cytology: T(1;Y)17E;YS.

origin: X ray induced in BSY. origin: X ray induced in BSY.
discoverer: Nicoletti. discoverer: Nicoletti.
references: Nicoletti and Lindsley, 1960, Genetics references: Nicoletti and Lindsley, 1960, Genetics
45: 1705-22. 45: 1705-22.
1960, DIS 34: 95-97. 1960, DIS 34: 95-97.
genetics: Male lethal. genetics: Male lethal.
*T(1;Y)1U
cytology: T(l;Y)20A;Ys. cytology: T(l;Y)20A;Ys.
origin: X ray induced in BSY. origin: X ray induced in BSY.
45: 1705-22. 45: 1705-22.
1960, DIS 34: 95-97. 1960, DIS 34: 95-97.
genetics: Male viable; fertile with a free Y but genetics: Male viable; fertile with a free Y but
sterile without. sterile without.
T(1;Y)1U
cytology: T(1;Y)3C;Y^, cytology: T(1;Y)19F;YS.
origin: X ray induced in y/BsY sperm. origin: X ray induced in BSY.
references: Nicoletti and Lindsley, I960, Genetics references: Nicoletti and Lindsley, 1960, Genetics
45: 1705-22 (fig.). 45: 1705-22.
1960, DIS 34: 95-97. 1960, DIS 34: 95-97.
genetics: Male lethal. genetics: Male viable; fertile with a free Y but
sterile without.
cytology: T(1;Y)8F;YS. U7;Y)724
origin: X ray induced in cytology: T(1;Y)9F;Y^.
discoverer. Nicoletti. origin: X ray induced in BSY.
references: Nicoletti and Lindsley, 1960, Genetics discoverer: Nicoletti,
I960, DIS 34: 95-97. 45: 1705-22.
genetics: Male lethal. 1960, DIS 34: 95-97.
*T(1;Y)117 genetics: Male viable; fertile with a free Y but
cytology: T(1;Y)17A;YL. sterile without.
origin: X ray induced in BSY.
discoverer: Nicoletti. W;Y)125
references: Nicoletti and Lindsley, I960, Genetics cytology: T(1;Y)15D;YL.
45: 1705-22. origin: X ray induced in BSY.
1960, DIS 34: 95-97. discoverer: Nicoletti.
genetics: Male viable; fertile with a free Y but references: Nicoletti and Lindsley, 1960, Genetics
sterile without. 45: 1705-22.
Tif7;YJ77« 1960, DIS 34: 95-97.
cytology: T(1;Y)16E;YL. genetics: Male viable; fertile with or without a free
origin: X ray induced in BSY. y.
discoverer: Nicoletti.
references; Nicoletti and Linda ley, I960, Genetics cytology: T(1;Y)3C;YL.
45: 1705-22. origin: X ray induced in y/BsY sperm.
I960, DIS 34: 95-97. discoverer Nicoletti.
45: 1705-22. 45: 1705-22.
I960, DIS 34: 95-97. 1960, DIS 34: 95-97.
genetics: Male lethal. genetics: Male viable; fertile with a free Y but
cytology: T(1;Y)11A;YL. T(l;Y)J40
origin: X ray induced in BSY. cytology: T(1;Y)3C;YL.
discoverer: Nicoletti. origin: X ray induced in y/BsY sperm,
references: Nicoletti and Lindsley, 1960, Genetics discoverer Nicoletti.
1960, DIS 34: 95-97. 45: 1705-22.
free Y. genetics: Male viable and fertile with or without a
free Y.
cytology: T(1;Y)6E;YS.
origin: X ray induced in BSY. cytology: T(1;Y)19B;YS.
45: 1705-22 (fig.). references: Nicoletti and Lindsley, 1960, Genetics
1960, DIS 34: 95-97. 45: 1705-22.
free Y. genetics: Male viable; fertile with a free Y but
cytology: T(1;Y)19F;YS. T(1;Y)U2
origin: X ray induced in BSY. cytology: T(1;Y)13E;YL.
1960, DIS 34: 95-97. 45: 1705-22 (fig.).
genetics: Male viable; fertile with a free Y but 1960, DIS 34: 95-97.
sterile without. genetics: Male viable; fertile with a free Y but
cytology: T(l;Y)19E;Ys. T(1;Y)145
origin: X ray induced in BSY. cytology: T(1;Y)11B;YS.
references: Nicoletti and Lindsley, I960, Genetics discoverer: Nicoletti.
1960, DIS 34: 95-97. 45: 1705-22.
genetics: Male viable; fertile with a free Y but 1960, DIS 34: 95-97.
sterile without. genetics: Male viable but sterile.
*T(1;Y)135 T(1;Y)U7
cytology: T(1;Y)18C;YL. cytology: T(1;Y)8F;YS.
origin: X ray induced in BSY. origin: X ray induced in fl^F.
discoverer Nicoletti. discoverer Nicoletti.
45: 1705-22. 45: 1705-22.
1960, DIS 34: 95-97. 1960, DIS 34: 95-97.
genetics: Male viable; fertile with a free Y but genetics: Male viable and fertile with or without a
sterile without. free F.
T(1;Y)137 *T(1;Y)U8
cytology. T(1;Y)19F;YS. cytology: T(1;Y)2D;YL.
origin: X ray induced in BSY. origin: X ray induced in y/BsY sperm,
discoverer: Nicoletti. discoverer Nicoletti.
45: 1705-22. 45: 1705-22.
1960, DIS 34: 95-97. 1960, DIS 34: 95-97.
genetics: Male viable; fertile with a free Y but genetics: Male viable and fertile with or without a
sterile without. free Y.
*T(1;Y)139 T(1;Y)U9
cytology: T(l;Y)20A;Ys. cytology: TXl;Yy6E;Y^.
origin: X ray induced in origin: X ray induced in BSY.
discoverer Nicoletti. di scoverer Nic oletti.
45: 1705-22. 45: 1705-22.
1960, DIS 34: 95-97. 1960, DIS 34: 95-97.
genetics: Male viable and fertile with or without a genetics: Male lethal.
free Y. T(J;Y)159
T(l;Y)150 cytology: T(1;Y)18A;Y^.
cytology: T(1;Y)3F;YS. origin: X ray induced in BSY.
origin: X ray induced in y/BsY sperm. discoverer: Nicoletti.
references: Nicoletti and Lindsley, 1960, Genetics 45: 1705-22.
45: 1705-22. 1960, DIS 34: 95-97.
1960, DIS 34: 95-97. genetics: Male viable and fertile with or without a
genetics: Male viable but sterile. free Y.
T0;Y)151 T(J;Y)164
cytology: T(1;Y)19F;YS. cytology: T(1;Y)3C;YL.
origin: X ray induced in BSY. origin: X ray induced in y/BsY sperm.
45: 1705-22. 45: 1705-22.
1960, DIS 34: 95-97. 1960, DIS 34: 95-97.
genetics: Male viable and fertile with or without a genetics: Male lethal.
free Y. *T(1;Y)169
W;Y)152 cytology: T(1;Y)11D;YS.
cytology: T(1;Y)13A;YL. origin: X ray induced in B^Y.
origin: X ray induced in BSY. discoverer: Nicoletti.
references: Nicoletti and Lindsley, 1960, Genetics 45: 1705-22 (fig.).
45: 1705-22. 1960, DIS 34: 95-97.
1960, DIS 34: 95-97. genetics: Male viable and fertile with or without a
genetics: Male viable; fertile with a free Y but free Y.
sterile without.
T(l;Y)240
T(1;Y)15S cytology: T(1;Y)14A;YL.
cytology: T(1;Y)7B;YS. origin: X ray induced in BSY.
origin: X ray induced in B^. discoverer: Nicoletti.
references: Nicoletti and Lindsley, 1960, Genetics 45: 1705-22 (fig.).
45: 1705-22. 1960, DIS 34: 95-97.
1960, DIS 34: 95-97. genetics: Male viable; fertile with a free Y but
genetics: Male lethal. sterile without.
T(1;Y)156 *T(1;Y)290
cytology: T(1;Y)7D;YL. cytology: T(1;Y)1A;2OA;YS.
origin: X ray induced in B^F. new order: 1A|YSP _ Y*<;
di sc overer: Nicole tti. 20F - 20A|lA - 20A|Y SD .
references: Nicoletti and Lindsley, 1960, Genetics origin: X ray induced in y/Bty sperm.
genetics: Male viable; fertile with a free Y but 45: 1705-22.
sterile without. 1960, DIS 34: 95-97.
*T(1;Y)157 genetics: Male lethal.
cytology: T(1;Y)14F;YL.
origin: X ray induced in B®Y. *T(l;Y;2)7
discoverer: Nicoletti. cytology: T(1;Y)14F;YS + T(Y;2)Y^;36C
references: Nicoletti and Lindsley, I960, Genetics new order: 1 — 14F|Y SP - Y LP J36C - 21;
45: 1705-22. 20 - 14F|Y SD ;
1960, DIS 34: 95-97. | - 60.
genetics: Male viable; fertile with a free Y but origin: X ray induced in
sterile without. discoverer: Nicoletti.
*T(l;Y)m references: Nicoletti and Lindsley, 1960, Genetics
cytology: T(1;Y)11A;YL. 45: 1705-22.
origin: X ray induced in BSY. 1960, DIS 34: 95-97.
discoverer Nicoletti. genetics: Male lethal.
T(1;Y;2)17 references: Nicoletti and Lindsley, 1960, Genetics

cytology: T(1;Y;2)7B;YL;39. 45: 1705-22.
new order: 1 - 7B|39 - 60; I960, DIS 34: 95-97.
20- 7B|YLD; genetics: Male lethal.
yS _ Y L P |39 - 21. *T(1;Y;2)16O
origin: X ray induced in y+Y. cytology: T(l;Y;2)17C;Ys;40-41; involvement of
discoverer: Nicoletti. chromosome 2 inferred from genetic data; not cyto-
references: Nicoletti and Lindsley, I960, Genetics logically observable; therefore new order ambig-
45: 1705-22. uous.
1960, DIS 34: 95-97. origin: X ray induced in BSY.
genetics: Male viable but sterile. discoverer: Nicoletti.
T(l;Y;2)109 references: Nicoletti and Lindsley, 1960, Genetics
cytology: T(l;Y;2)3C;YL;40-41; involvement of chro- 45: 1705-22.
mosome 2 inferred from genetic data; not cytologi- 1960, DIS 34: 95-97.
cally observable; new order therefore ambiguous. genetics: Male viable but sterile.
origin: X ray induced in y/BsY sperm.
discoverer: Nicoletti. *T(J;Y;3)5
references: Nicoletti and Lindsley, 1960, Genetics cytology: T(1;Y)11D;YL + T(1;3)14F;72.
45: 1705-22 (fig.). new order: 1 - 1 1 D | Y L P - Y s ;
I960, DIS 34: 95-97. 20 - 14F|72 - 61;
genetics: Male viable but sterile. Y L D | n D _ I4p]72 - 100.
*T(l;Y;2)110 origin: X ray induced in y+Y.
cytology: T(1;2)19D;55F + T(Y;2)YS;45F. discoverer: Nicoletti.
new order: 1 - 19D|55F - 45F|Y SP - yL; references: Nicoletti and Lindsley, 1960, Genetics
20 - 19D|55F - 60; 45: 1705-22.
YSD|45F _ 21. 1960, DIS 34: 95-97.
origin: X ray induced in BSY. genetics: Male viable but sterile.
references: Nicoletti and Lindsley, 1960, Genetics T(1;Y;3)121
45: 1705-22. cytology: T(1;Y;3)6F;YS;86D.
1960, DIS 34: 95-97. new order: 1 — 6 F | Y S P - Y L ;
genetics: Male viable but sterile. 20 - 6F|86D - 100;
YSD|86D - 61.
*T(l;Y;2)130 origin: X ray induced in B^F.
cytology: T(l;Y;2)llF;YL;40-41; involvement of discoverer: Nicoletti.
chromosome 2 inferred from genetic data; not cyto- references: Nicoletti and Lindsley, 1960, Genetics
logically observable; new order therefore ambig- 45: 1705-22.
uous. 1960, DIS 34: 95-97.
origin: X ray induced in BSY. genetics: Male lethal.
references: Nicoletti and Lindsley, 1960, Genetics T(1;Y;3)127
45: 1705-22 (fig.). cytology: T(1;Y;3)19F;YS;85E.
I960, DIS 34: 95-97. new order. 1 - 19F|85E - 61;
genetics: Male viable but sterile. 20 - 19F|YSI>;
*T(1;Y;2)U6 YL _ YSPJ85E - 100.
cytology: T(1;Y)7D;YL + T(l;2)20A;57F. origin: X ray induced,
new order: 1 - 7 D | Y L P - Y s ; discoverer: Nicoletti.
20F - 20AJ57F - 60; references: Nicoletti and Lindsley, 1960, Genetics
Y L D |7D - 20A|57F - 21. 45: 1705-22.
origin: X ray induced in B^F. 1960, DIS 34: 95-97.
discoverer Nicoletti. genetics: Male viable but sterile.
references: Nicoletti and Lindsley, I960, Genetics *T(1;Y;3)134
45: 1705-22. cytology: T(1;Y)12E;YS + T(1;3)19B;62A.
1960, DIS 34: 95-97. new order: 1 — 1 2 E | Y S P — Y**;
genetics: Male viable but sterile. 20F - 19E|62A - 61;
*T(J;Y;2)1S3 YSD|i2E - 19E|62A - 100.
cytol ogy: T(l;Y;2)l 7A; YS;3SD. origin: X ray induced in BiSy.
new order: 1 - 17A|35D - 60; discoverer: Nicoletti.
20 - 17A|Y SD ; references: Nicoletti and Lindsley, 1960, Genetics
yL _ Y SP |35D - 2 1 . 45: 1705-22.
origin: X ray induced in BSY. 1960, DIS 34; 95-97.
discoverer: Nicoletti. genetics: Male viable but sterile.
*T(l;Y;3)m origin: X ray induced in BSY.

L discoverer: Nicoletti.
cytology: T(l;Y;3)7A;Y ;70C.
new order: 1 - 7A|70C - 100; references: Nicoletti and Lindsley, 1960, Genetics
20 - 7A|YLD; 45: 1705-22.
yS _ yLP|70C - 61. 1960, DIS 34: 95-97.
origin: X ray induced in BSY. genetics: Male lethal.
references: Nicoletti and Lindsley, 1960, Genetics T(1;Y;4)U6
45: 1705-22. cytology: T(1;Y)14D;YS + T(l;4)9C;101.
1960, DIS*34: 95-97. new order: 1 - 9C|l01;
genetics: Male viable but sterile. 20 - 1 4 D | Y S D ;
*T(1;Y;3)138 yL _ YSP|l4D - 9C|l02.
cytology: T(1;Y;3)11A;Y^;84B. origin: X ray induced in BSY.
new order: 1 - 11A|84B - 61; discoverer: Nicoletti.
20-llAJYLD ; references: Nicoletti and Lindsley, 1960, Genetics
yS _ y L P | 8 4 B - 100. 45: 1705-22.
origin: X ray induced in BSY, 1960, DIS 34: 95-97.
discoverer: Nicoletti. genetics: Male viable but sterile.
references: Nicoletti and Lindsley, 1960, Genetics
45: 1705-22. *T(I;2)7
I960, DIS 34: 95-97. origin: X ray induced.
genetics: Male viable but sterile. discoverer: Bonner, 1931.
T(1;Y;3)W references: Dobzhansky, 1935, Z. Induktive
cytology: T(1;Y)12A;Y^ + T(1;3)3F;69C. Abstammungs- Vererbungslehre 68: 134—62.
new order. 1 - 3F|69C - 100; genetics: X broken between rb and cv; chromosome 2
20-12A|yLD; to the right of sp. Male and heterozygous female
yS _ yLP|i2A - 3F|69C - 61. viable and fertile; homozygous female poorly viable
origin: X ray induced in y/BsY sperm. and sterile. XP2D recoverable as an aneuploid
discoverer: Nicoletti. segregant that is duplicated for the loci of y
references: Nicoletti and Lindsley, 1960, Genetics through rb but is not demonstrably deficient for 2R
45: 1705-22. markers; nothing written to indicate that it is de-
1960, DIS 34: 95-97. ficient for M(2)c.
genetics: Male viable with a free Y but sterile. Male *T(1;2)26
lethal without a free F. origin: X ray induced in R(l)2.
T(1;Y;3)144 discoverer: Pontecorvo, 1941.
cytology: T(1;Y;3)15E;Y^;74D. synonym: T(Xc2;2)26.
new order. 1 - 15E|74D - 100; references: 1942, DIS 16: 65.
20-15E|YLD; genetics: Section of X including car and bb inserted
y S _ y L P | 7 4 _ gj,. into base of 2L. Homozygous lethal.
origin: X ray induced in BSY. T(l;2)51b
discoverer Nicoletti. cytology: T(l;2)3Cl-2;3D6-7;20A;52F.
references: Nicoletti and Lindsley, 1960, Genetics new order: 1 - 3C1 |20A - 3D7|20A - 20F;
45: 1705-22. 21 - 52F|(3C2 - 3D6)|52F - 60.
1960, DIS 34: 95-97. origin: X ray induced w.ln(l)w™4 =In(l)3Cl-2;20A.
genetics: Male with free Y viable but sterile. Male discoverer Lefevre, 51b7.
without free Y lethal. synonym: T(l;2)w*Slb7\
T(1;Y;3)IS4 references: 1951, DIS 25: 71.
cytology: T(l;Y;3)10A;Ys;97A. 1952, DIS 26: 66.
new order. 1 - 10A|97A - 61; Ratty, 1954, Genetics 39: 513-28.
20 - 10A|ySD; genetics: Segregant Dp(l;2)51b = Dp(l;2)3Cl-
yL _ ySPJ97A _ 100, 2;3D6-7;52F survives; duplicated for loci of w,
origin: X ray induced in B^Y. rst, spl, fa, and dm. Duplication used to cover
discoverer Nicoletti. lethality of N in studies of pseudoallelism at the TV
references: Nicoletti and Lindsley, i960. Genetics locus (Welshons and Von Halle, 1962, Genetics 47:
45: 1705-22. 743-59).
1960, DIS 34: 95-97. *T(l;2)106
genetics: Male viable bat sterile. origin: X ray induced.
T(1;Y;3)U1 discoverer: Sturtevant, 1930.
cytology: TX1;Y;3)17A;YL;94. genetics: Break in X chromosome near centromere to
new order: 1 - 17AJ94 - 61; right of /; break in chromosome 2 near centromere,
20- probably in 2L. Male fertile; homozygous female
viable and fertile. Crossing over and disjunction
for both chromosomes X and 2 normal in genetics: Left break between ras and v; right break
T(l;2)106/+ female. T(l;2)106/+/Y female shows between g and pi. Male fertile. The segregant
nondisjunction of X's. Dp(l;2)A124 =Dp(l;2)10A;13Al-2;59, which is
*T(l;2)260-31 duplicated for v+ through g+, survives as fairly
cytology: T(1;2)9A;24;29. viable and fertile female, but male carrying
new order: 1 - 9A|(24 - 29)|9A - 20; Dp(l;2)A124 dies as embryo. The complementary
21 - 24J29 - 60. Dt(l)A124 -Df(l)10A;13Al-2 survives as a fertile
origin: X ray induced simultaneously with y260-31% X/X/Df triplpid female and as an X/X/Di diploid
discoverer: Fano, 1941. metafemale but not as an X/Df diploid.
references: Sutton, 1943, Genetics 28: 210—17. *7(\;2)B**v: Translocation(1;2) Bar
genetics: Male lethal; lethality attributable to the cytology: T(1;2)15F-1$A1;33B superimposed on
independent mutation to y260-31 since T(l;2)260- In(l)lB3-4;l 9F-20C1.
31) y260-31/D£(l)sc260-25 is lethal. new order: 1A1 - lB3|l9F - 16A1 |33B - 60;
*T(l;2)271b 20F - 20Cl|lB4 - 15FJ33B - 21.
cytology: T(l;2)3C3-7;40; inferred from Mackensen's origin: X ray induced in In(l)sc4.
fig. 15A, G, and H (1935). discoverer. Yu, 48g.
origin: X ray induced, genetics: Position effect at B. Male sterile.
discoverer: Patterson. T(l;2)Bbd: Translocation(l;2) Bar-baroid
synonym: Df(l)Del271b. cytology: T(l;2)16Al-2;48C2-3 + In(2R)41A;47A
references: 1932, Am. Naturalist 66: 193-206. (Bridges in Morgan, Bridges, and Schultz, 1936,
Mackensen, 1935, J. Heredity 26: 163-74 (fig.). Carnegie Inst. Wash. Year Book 35: 291).
genetics: Mutant for N. new order 1 - l6Al|48C2 - 47A|41A - 47A|4lA -
21;
*T(l;2)A50b: Translocation(1;2) from Austin 2 0 - 16A2J48C3 - 60.
cytology: T(1;2)2B;15F;41; inferred from fig. 17H of origin: X ray induced simultaneously with In(2R)41-
Mackensen (1935). 47.
new order: 1 - 2B|l5F - 20; discoverer: Dobzhansky, 31b5.
21 -41J15F - 2BJ41 - 60. references: 1932, Genetics 17: 369—92.
references: Mackensen, 1935, J. Heredity 26: 163— genetics: Recessive position effect for 15. Translo-
74 (fig.). cation and inversion probably not separable.
genetics: Left break in X between br and pn; right *T(\;2)B*>G: Translocation(l;2) Bar of Dubinin
break between r and t. Mutant for f. and Goldat
*T(l;2)A6Jb cytology: T(1;2)4;15F-16A;20;40-41; inferred from
cytology: T(1;2)15F; breakpoint in chromosome 2 at Dubinin and Goldat's figure,
unknown position in left arm, which also carries an new order: 1 - 4|l5F - 4|20;
inversion. Breakpoint in X inferred from 21 - 40|(16A - 20)|41 - 60.
Mackensen's fig. 17G (1935). origin: X ray induced.
references: Mackensen, 1935, J. Heredity 26: 163— discoverer: Dubinin and Goldat, 1936.
74 (fig.). references: 1936, Biol. Zh. (Moscow) 5: 881—84
genetics: Mutant for f. (fig-).
*TO;2)A106 genetics: Position effect for B. Male lethal.
cytology: T(l;2)6-7;12;17; rough estimates of break- T(l;2)BId: Translocaiion0;2) Blond
points in X from Mackensen's fig. 171 (1935); chro- cytology: T(1;2)1C3-4;6OB12-13 + In(2R)42A2-
mosome 2 broken in euchromatin of left arm. 3;58A4-B1.
new order: 1 — 6J17 — 20; new order: 1A - 1C3|6OB12 - 58Bl|42A3 -
21 - ?|12 - 17|7 - 12|? - 6 0 . 58A4|42A2 - 21;
references: Mackensen, 1935, J. Heredity 26: 1 6 3 - 20 - 1C4|6OB13 - 60F.
74 (fig.), origin: Spontaneous in In(2R)Cy.
genetics: Mutant for Bx. discoverer: Burkart, 1930.
*T(l;2)A124 references: 1931, Rev. Fac. Argon. Vet. Univ.
cytology: T(l;2)10A;13Al-2;59. Buenos Aires 7: 393-491.
new order: 1 - 10AJ13A2 — 20; Burkart and Stern, 1933, Z. Induktive Abstammungs-
21 - 59|(10A - 13A1)|59 - 60. Vererbungslehre 64: 310—25.
origin: X ray induced. Bridges, 1937, Cytologia (Tokyo), Fuji! Jub. Vol.
discoverer: Patterson, Stone, Bedichek, and Suche, 2: 745-55.
1934. Morgan, Bridges, and Schultz, 1938, Carnegie Inst.
references: Macfcensen, 1935, J. Heredity 26: 163— Wash. Year Book 37: 307.
74 (fig.). genetics: Associated with Bid. Both aneuploid seg-
Patterson, Stone, and Bedichek, 1935, Genetics regants survive. The XD2P element is duplicated
2Ch 259-79 (f%.). for y, ac, sc, Hw, svr, su(s), 1(1 )7e, su(b), and
1937, Genetics 22: 407-26. M(l)Bld and deficient for sp, bs, be, Pin, and
Pipkin, 1940, Texas Univ. Publ. 4032: 126-56. M(2}c; heterozygote extreme Plexate and slight
Minute with small dark body and slow development; new order: 1 - 8C|l8D - 8D|l8D - 20|40 - 60;
viability low; male sterile, female but slightly fer- 2Q|40 - 21.
tile. 2DXP is reciprocally duplicate-deficient; Tentative.
heterozygous female Blond and extreme Minute origin: X ray induced,
\M(l)Bld\ with short, broad, occasionally downward discoverer: Roberts, 1965.
curved wings; ecloses 3—4 days late; male lethal. genetics: Male lethal. Recombination reduced in X
chromos ome.
T(1;2)C6: Translocation(l;2) Crossover T(1;2)C179
suppressor cytology. T(1;2)9A;49A +ln(l)5C;20.
cytology: T(l;2)12E;40-41;60B; position of break- new order: 1 - 5C|20 - 9A|49A - 21;
point in chromosome 2 with respect to centromere 20|5C - 9A|49A - 60.
not determined, origin: X ray induced.
new order: 1 - 12EJ41 - 60B|40 - 21; discoverer: Roberts, 1965.
20 - 12E|60B - 60F. genetics: Male lethal. Recombination reduced in X
origin: X ray induced, chromosome.
genetics: Male lethal. Recombination reduced in 2R. T(1;2)C183
T(J;2)C20 cytology: T(l;2)12E;40-41 + In(2L)24C;30A; trans-
cytology: T(l;2)12E;30B. location breakpoint in chromosome 2 not determined
origin: X ray induced, with respect to centromere; new order therefore am-
discoverer: Roberts, 1964. biguous.
genetics: Male viable but sterile. Recombination re- new order: 1 — 12E|40 — 60;
duced in 2L. 20 - 12E|40 - 30A|24C - 30A|24C - 21.
T(1;2)C54 Tentative.
cytology: T(1;2)12E;32F. origin: X ray induced.
origin: X ray induced. discoverer: Roberts, 1965.
discoverer: Roberts, 1964. genetics: Male lethal. Recombination reduced in 2L.
genetics: Male lethal. Recombination reduced in 2L.
T(l;2)C60 T(1;2)C239
cytology: T(l;2)20;52B. cytology: T(1;2)7A-B;36C;39E.
origin: X ray induced. new order: 1 - 7A|36C - 39E|7B - 20;
discoverer. Roberts, 1964. 21 - 36C|39E - 60.
genetics: Male viable and fertile. Recombination re- origin: X ray induced,
duced in 2R. discoverer: Roberts, 1965.
genetics: Male lethal. Recombination reduced in X
T(J;2)C84 chromosome. The segregant Dp(2;l)C239 =
cytology: T(l;2)3F;17E-F;30A. Dp(2;l)7A-B;36C;39E survives.
new order. 1 - 3F|l7F - 20; T(1;2)C2S6
21 - 30A|3F - 17E|30A - 60. cytology: T(l;2)2A;40-41 + In(l)7E;17A;18B; posi-
origin: X ray induced. tion of breakpoint with respect to centromere in
discoverer: Roberts and D. Stewart, 1964. chromosome 2 not determined; new order therefore
genetics: Male viable but sterile. Recombination re- ambiguous. For example; if chromosome 2 is
duced ia X and 2L. broken in 2L:
T(1;2)C121 new order: 1 — 2 A | 4 O — 60;
cytology: T(l;2)20;35F;40. 20 - 18B|17A - 18B|7E - 17A|7E -
new order: 1 - 20|(35F - 40)J20; 2A|40 - 21.
21 -35FJ40 - 6 0 . origin: X ray induced.
origin: X ray induced, discoverer: Roberts, 1965.
discoverer: Roberts, 1965. genetics: Male lethal. Recombination reduced in X
genetics: Male viable and fertile. Recombination re- chromosome.
duced in 2L.
T(1;2)C171 T(J;2)C26J
cytology: T(ls2)12A;40-41. cytology: T(l;2)14C;40-41,' breakpoint in chromo-
origin: X ray induced. some 2 with respect to centromere not determined.
discoverer: Roberts, 1965. origin: X ray induced.
genetics: Male viable but sterile. Recombination re- discoverer: Roberts, 1965.
duced in X chromosome. genetics: Male viable but sterile. Recombination re-
T(1;2)Cm duced in X chromosome.
cytology: T(l;2)20;40-41 +ln(t)8C-D;l8D; transloca- T(1;2)C262
tion breakpoint in chromosome 2 with respect to cytology: T(1;2)11A;18A;4O-41; position of break-
centromere not determined; n*w order therefore am- point in chromosome 2 with respect to centromere
biguous. not determined.
new order: 1 — 11A|4O — 60;

20 - 18A|llA - 18A|40 - 21. cytology: T(l;2)7B2-4;19-20;41El-2 superimposed on
Tentative. R(1)1A3-4;19F-2OA1.
origin: X ray induced. new order: |lA4 - 7B2|20-20F - 20A11;
discoverer: Roberts, 1965. 21 - 4lEl|7B4 - 19|41E2 - 60.
genetics: Male lethal. Recombination reduced in X origin: X ray induced in R(l)2.
chromosome. discoverer: Hannah, 1947.
T(1;2)C3U genetics: Mutant for ct but not cm, sn, or oc. Male
cytology: T(l;2)5D;40-41 + T(1;2)9D;51D + lethal. T(l;2)ctl4*2/Dp(l;3)sn1:3al male rarely
T(l;2)20;56F; position of left breakpoint in chromo- survives; probably sterile.
some 2 with respect to centromere not determined. *T(l;2)ct268-i7
new order: 1 - 5D|40 - 51D|9D - SD|40 - 21; cytology: T(l;2)7B2-5;41E2-4 (Hoover).
20|S6F - 51D|9D - 20J56F - 60. origin: X ray induced.
Tentative because heterochromatic realignments discoverer: Demerec, 34h.
ambiguous. genetics: Mutant for ct but not scp or sn. Male
origin: X ray induced. lethal.
discoverer: Roberts, 1965. *T(1;2)ct268.24
genetics: Male viable but sterile. Recombination re- cytology: T(l;2)7B2-5;41F6-42Al.
duced in X and 2R. origin: X ray induced.
discoverer: Hoover, 35i.
T(1;2)C324 genetics: Mutant for ct but not scp or sn. Male
cytology: T(l;2)15F;20;30A. lethal.
new order: 1 - 15F|20 - 15F|3OA - 60; *T(J;2)ct268-26
20|30A - 2 1 . cytology: T(J;2)7B3-C1;36E.
discoverer: Roberts, 1965. discoverer: Hoover, 35j.
genetics: Mutant or deficient for f. Male lethal. Re- genetics: Mutant for ct but not scp or sn. Male
combination reduced in 2L. lethal.
T(1;2)C349 *T(1;2)ct268.32
cytology: T(1;2)6C;47D +In(l)2E;20. cytology: T(J;2)lE-F;3D-E;7B2-5;46 (Hoover),
new order: 1 - 2E|20 - 6C|47D - 21; new order: 1A - 1EJ3E - 7B2|46 - 21;
20|2E - 6CJ47D - 60. 20 - 7B5|3D - 1F|46 - 60.
origin: X ray induced, origin: X ray induced in y.
discoverer. Roberts, 1965. discoverer. Demerec, 38e.
genetics: Male lethal. Recombination reduced in X genetics: Mutant for ct but not fa, dm, scp, or sn.
chromosome. Male lethal.
T(1;2)C357 *T(l;2)ct268.33
cytology: T(l;2)20;56F. cytology: T(1;2)7B2-5;41E (Hoover).
discoverer: Roberts, 1965. discoverer: Demerec, 38e,
genetics: Male viable but sterile. Recombination re- genetics: Mutant for ct but not en. Male lethal.
duced in 2R. *T(l;2)ct268~4l
cytology: T(l;2)7B2-S;37C2-3 (Sutton).
U Translocation(l;2) cut origin: X ray induced.
cytology: T(1;2)7B; other breakpoints not recorded, discoverer Demerec, 391.
origin: X ray induced in R(l)2. genetics: Mutant for ct but not cm or sn. Male
discoverer: Hannah, 1947. lethal.
genetics: Mutant for ct but not y, ac, sc, cm, sn, or *T(J;2)DJ: Translocation(l;2) from deoxycytidine
oc. Male lethal. cytology: T(1;2)6F;26C.
origin: Induced by tritiated deoxycytidine.
cytology: T(1;2)7B2-3;8E2-3;25C superimposed on discoverer: Kaplan, 1965.
R(1)1A3-4;19F-2QA1. references: 1966, DIS 41: 59.
new order. |lA4 - 7B2|8E2 - 20-20F - 20Al|; genetics: Male lethal.
21 - 25C|(7B3 - 8E2)|25C - 60. 7(1,2)02
origin: X ray induced in R(l)2. cytology: T(1;2)8B;46B.
discoverer. Hannah, 1947. origin: Induced by tritiated deoxycytidine.
genetics: Mutant for ct but not cm or sn; male lethal. discoverer: Kaplan, 1965.
T(l;2)ct7cl/Dp(l;3)m13al male survives and is references: 1966, DIS 41: 59.
fertile. The segregant Dp(I;2)ct7cI » Dp(l;2)7B2- genetics: Male lethal.
3;8E2-3;25C survives; duplicated for an but not cm; *T(J;2)ef: Trtmslacation(h'2} elfin
male and female have darker, rooflike wings, en- cytology: T(1;2)14C8-D1;2R.
larged abdomens, and are sterile. origin: Induced by trie thy len erne la mine (CB. 1246).
discoverer: Fahmy, 1952. origin: X ray induced simultaneously with

references: 1959, DIS 33: 86. T(2;3)135 = T(2;3)37;85A, which has been lost.
genetics: Mutant for ef. Male sterile. discoverer: Lindsley, Edington, and Von Halle.
*T(1;2){2S7.15: Translocaiion(l;2) forked references: 1960, Genetics 45: 1649—70.
cytology: T(l;2)13E9-10;15E2-3;24F (Sutton). genetics: Variegated for a lethal.
new order: 1A - 13E9|l5E3 - 20; T(l;2)l-vJ50
21 - 24F|(13E10 - 15E2)|24F - 60. cytology: T(l;2)16-17;40.
discoverer: Demerec, 35a. discoverer: Lindsley, Edington, and Von Halle.
genetics: Mutant for f but not M(l)o or B. Male references: 1960, Genetics 45: 1649—70.
lethal. genetics: Variegated for a lethal; male sterile.
*T(J ;2)f2S7-22 T(l;2)l-v219
cytology: T(1;2)4D2-3;8F;15E4-F1;39E;41F-42A cytology: T(l;2)10A;40.
superimposed on Dp(l;l)15F9-16Al;16A7-Bl. origin: X ray induced.
new order: 1 - 4D2|(8F - 15E4)|41F - 39E|(4D3 - discoverer: Lindsley, Edington, and Von Halle.
8F)|39E - 21; references: 1960, Genetics 45: 1649-70.
2 0 - 16Al|l6A7 - 16A1|16A7 - genetics: Variegated for a lethal; male sterile.
15Fl|42A - 6 0 . *T(l;2)l-v223
origin: X ray induced in y BlBl. cytology: T(l;2)14F;41;50E.
discoverer: Demerec, 36c. new order: 1A - 14F|(41 - 50E)|l4F - 20;
genetics: Mutant for / but B unaffected. Male lethal. 21A - 41 |50E - 60.
T(1;2)K1: Translocation(l;2) of Krivshenko origin: X ray induced.
cytology: T(1;2)1A5-B3;2O;29A-B; deficient for 1B3- discoverer: Lindsley, Edington, and Von Halle.
20. references: 1960, Genetics 45: 1640—70.
new order: 1A1 - 1A5|29B - 60; genetics: Variegated for a lethal and defective ex-
20|29A - 2 1 . ternal male genitalia; male sterile.
discoverer. Krivshenko, 56cl2. T(l;2)lt: Translocation(1;2) light
references: 1956, DIS 30: 75. cytology: T(1;2)20C-D;40F.
genetics: Homozygous lethal. Fly hyperploid for the origin: X ray induced in chromosome carrying eq.
2LDXP element survives. discoverer: Schultz.
genetics: Variegated for It.
T(l;2)l-v25: Translocationfl ;2) lethal-variegated *T(l;2)lf»i6: Translocation(l;2) light-mottled
cytology: T(1;2)19-20;40-41; position of breakpoint cytology: T(1;2)11A;12F;22D;4OB-F.
in chromosome 2 with respect to centromere not de- new order: 1 - HA|l2F - 20;
termined. 21 - 22D|llA - 12F|40B - 22D|40F -
origin: X ray induced. 60.
discoverer: Lindsley, Edington, and Von Halle. origin: X ray induced.
genetics: Variegated for a lethal; male sterile. references: 1958, Genetics 43: 395—403.
*T(1;2)l-v47 genetics: Variegated for It.
cytology: T(1;2)8F-9B; heterochromatic material in- *T(1;2)ltm31
serted in X; genetic results suggest linkage be- cytology: T(l;2)8F;28D;40B-F.
tween X and 2. new order. 1 - 8F|28D - 40B|8F - 20;
origin: X ray induced. 21 - 28D|40F - 60.
discoverer Lindsley, Edington, and Von Halle. origin: X ray induced,
references: I960, Genetics 45: 1649-70. discoverer. Hessler, 1957.
genetics: Variegated for a lethal; g£-like phenotype. references: 1958, Genetics 43: 395-403.
T(J;2)l-v75 genetics: Variegated for It.
cytology: T(l;2)19-20;41. *T(l;2)lz: Translocation(l;2) lozenge
origin: X ray induced. cytology: T(1;2)8D12-E1;33A-B (Hannah).
discoverer. Lindsley, Edington, and Von Halle. origin: X ray induced.
references: 1960, Genetics 45: 1649—70. discoverer. Green and Green,
genetics: Variegated for a lethal; male sterile. references: 1956, Z. Induktive Abstammungs-
T(l;2)l'v129 Vererbungslehre 87: 708-21.
cytology: T(1;2)18B;41. genetics: Mutant for lz.
discoverer: Lindsley, Edington, and Von Halle. *T(J;2)N2"'9: Translocation(l;2) Notch
references: 1960, Genetics 45: 1649—70. cytology: T(1;2)3C;41. Cytology not examined;
genetics: Variegated for a semilethal; male sterile. breakpoints inferred from genes affected (Schultz).
T(t;2)l-vUS origin: X ray induced,
cytology: T(t;2)l8~19;41. discoverer: Demerec, 3315.
genetics: Variegates for rst, N, dm, and, at low tem- genetics: Variegates for w, rst, fa, dm, and ec but
peratures, w; also variegates for abnormal abdomen not pn, kz, or bi.
and M(2)S2 but not for stw, ap, or msf (Schultz). *T(l;2)N264-69
T(1;2)N264-io cytology: T(l;2)3C7-8;44C4-5 (Demerec).
discoverer: Demerec, 331. discoverer: Demerec, 38k.
genetics: Variegates for rst, N, and dm but not w. references: 1941, Proc. Intern. Congr. Genet., 7th.
Carries normal alleles of M(2)S2, ap, msf, and tk pp. 99-103.
(Schultz). X/Y male lethal; X/Y/Y survives. genetics: Carries a mutant allele of N and normal
*T(J;2)N264-23 alleles of w, rst, and dm.
cytology: T(1;2)3C8-9;41A (Demerec and Hoover). *T(l;2)N264-80
origin: X ray induced. cytology: T(l;2)3C6-7;36;40 +ln(l)ll;20 (Sutton).
discoverer: Demerec, 35h. new order: 1 - 3C6|(36 - 40)|3C7 - 11120 - ll|20;
references: 1941, Proc. Intern. Congr. Genet., 7th. 21 - 36|40 - 60.
pp. 99-102. origin: X ray induced,
genetics: Variegates for rst and fa but not w or dm. discoverer: Demerec, 39d.
*T(J;2)N264-24 references: 1941, Proc. Intern. Congr. Genet., 7th.
cytology: T(l;2)3C8-9;40F (Demerec). pp. 99-103.
origin: X ray induced. genetics: Contains mutant allele of N but normal
discoverer: Demerec, 35h. alleles of w, rst, dm, and ec.
references: 1941, Proc. Intern. Congr. Genet., 7th. *T(l;2)N264-82
pp. 99-103. cytology: T(1;2)3C3-4;41A + T(l;2)20A;57.
genetics: Variegates for w, rst, and fa. new order: 1 - 3C3|41A - 57|20A - 20F;
*T(l;2)N264.so 21 - 41A|3C4 - 20A|57 - 60.
cytology: T(l;2)3C7-9;20Cl-F;22A2-3 (Hoover), Tip of 2L also in chromocenter (Sutton).
new order: 1 - 3C7|20F; origin: X ray induced,
21 - 22A2|3C9 - 20Cl|22A3 - 60. discoverer Demerec, 39d.
origin: X ray induced. genetics: Variegates for w, rst, fa, and dm but not
discoverer: Demerec, 37k. pn, ec, or in.
references: 1941, Proc. Intern. Congr. Genet., 7th. *T(1;2)N264-1O2
pp. 99-103. cytology: T(l;2)3C6-7;50E;56C (Sutton).
genetics: Variegates for fa but not w, rst, or drn. new order: 1 - 3C6J(50E - 56C)J3C7 - 20;
21 - 50E|56C - 60.
*T(1;2)N264-S3 discoverer: Demerec, 391.
cytology: T(1;2)3C6-7;34C7-D1. genetics: Carries mutant allele of N and normal
origin: X ray induced. alleles of w, rst, and dm.
discoverer: Demerec, 38a. T(1;2)OR6: Translocation(l;2) from Oak Ridge
references: 1941, Proc. Intern. Congr. Genet., 7th. cytology: T(l;2)2A;60D.
pp. 99-103. origin: X ray induced in y.
genetics: Carries normal alleles of rst, fa, and dm. discoverer Waiters, 1959.
*T(l;2)N264.S9 genetics: Male viable and fertile. Homorygous fe-
cytology: T(l;2)3C8-9;40F (Hoover). male viable. XD2P element can replace one chro-
origin: X ray induced. mosome 2, producing a deficiency for the tip of 2R;
discoverer. Demerec, 38d. resulting progeny are Minute owing to inclusion of
references: 1941, Proc. Intern. Congr. Genet., 7th. M(2)c locus in the deficiency.
pp. 99-103. T(\;2}OR7
genetics: Variegates for w, rst, and spl but not pn, cytology: T(1;2)3A;41E.
kz, or dm. origin: X ray induced in y.
*T(l;2)N264-62 discoverer Waiters, 1959.
cytology: T(1;2)3C7-8;41A-B (Sutton). genetics: Male lethal. Male survives with BSw+Y
origin: X ray induced. but is sterile.
discoverer Demerec, 38e. 7(7/2)0*8
references: 1941, Proc. Intern. Congr. Genet., 7th. cytology: T(l;2)20;4O-41; position of breakpoints
pp. 99-103. with respect to centromeres not determined.
genetics: Variegates for w, rst, and fa but not dm. origin: X ray induced.
T(l;2)N264-66 discoverer Waiters, 1959.
cytology: T(1;2)3C6-7;41 -f T(1;2)7C9-D1;53F genetics: Male viable and fertile. Homozygous fe-
(Hoover), male viable.
new order: 1 - 3C6|41 - 53FJ7D1 - 20; TO;2}OR9
21 - 41J3C7 - 7C9|53F - 60. cytology: TX1;2)3A;J4F;41.
origin: X ray induced, new order. 1 - 3AJ14F - 20;
discoverer: Demerec, 38e. 21 _ 4 l | l 4 F - 3A|41 - 60,
origin: X ray induced in y. T(l;2)0R23

discoverer: Warters, 1959. cytology: T(l;2)6B;40.
genetics: Male lethal; lethality not covered by origin: X ray induced.
Bsw+Y; therefore probably associated with break discoverer: Warters, 1961.
at 14F. genetics: Male viable but sterile.
T(1;2)OR11 T(1;2)0R25
cytology: T(1;2)14F;41. cytology: T(1;2)1B;38E.
origin: X ray induced. origin: X ray induced in y.
discoverer: Warters, 1959. discoverer: Warters, 1961.
genetics:. Variegated for a lethal. X/Y male viable genetics: Male viable but sterile.
but sterile. T(l;2)0R26
T(J;2)ORU cytology:' T(1;2)15A;41.
cytology: T(1;2)18D;46B. origin: X ray induced.
origin: X ray induced. discoverer: Warters, 1961.
discoverer: Warters, 1959. genetics: Male viable but sterile; X/0 male has
genetics: Male quite inviable; rare survivor has un- melanotic, distended wings.
expended wings and crossed scute liars. T(l;2)0R27
T(1;2)OR15 cytology: T(1;2)16D;34B.
cytology: T(1;2)11B;6OE. origin: X ray induced.
discoverer: Warters, 1959. genetics: Male viable but sterile.
genetics: Male viable but sterile. T(l;2)0R28
T(1;2)OR17 cytology: T(1;2)3B;39E.
cytology: T(1;2)3C;37C. origin: X ray induced in y.
origin: X ray induced in y. discoverer: Warters, 1961.
discoverer: Warters, 1961. genetics: Male viable but sterile, with either a
genetics: Male viable but sterile, with either a normal Y or Bsw+Y.
normal Y or with Bsw+Y. T(1;2)0R29
cytology: T(l;2)8D;40.
T(1;2)ORW origin: X ray induced.
cytology: T(l;2)20B;30E. discoverer: Warters, 1961.
origin: X ray induced. genetics: Male viable but sterile.
discoverer: Warters, 1961. T(1;2)OR30
genetics: Male viable but sterile. 2LDXP element cytology: T(l;2)20;40-41; position of breakpoints
recoverable in viable hyperploids with outstretched with respect to centromeres not determined.
wings. origin: X ray induced,
T(1;2)OR19 discoverer: Warters, 1961.
cytology: T(l;2)20;51F +In(2R)42B;48E;57C. genetics: Male viable but sterile.
new order: 1 - 20|51F - 48E|57C - 51F|20; T(1;2)OR32
21 - 42B|48E - 42B|57C - 60E. cytology: T(l;2)20;54A.
discoverer. Warters, 1961. discoverer: Warters, 1961.
genetics: Male viable and weakly fertile. Homozy- genetics: Male viable but sterile. Male hyperploid
gous female viable. Dp(2;l)OR19 ~ for 2RDXP element survives.
Dp(2;l)20;48E;51F;57C survives in both male and T(1;2)OR33
female. cytology: T(1;2)17;47A.
T(1;2)OR20 origin: X ray induced.
cytology: T(1;2)16C;43B. discoverer: Warters, 1961.
origin: X ray induced, genetics: Male viable but sterile.
discoverer: Waiters, 1961. T(1;2)OR36
genetics: Male lethal; lethal originated after trans- cytology: T(l;2)20;40-41; position of breakpoints
location. with respect to centromeres not determined.
cytology: T(1;2)19E;32D. discoverer Warters, 1961.
genetics: Male lethal. cytology: T(l;2)10A;S0C.
T(J;2)OR22 origin: X ray induced,
cytology: T(1;2)8D;22B. discoverer: Warters, 1961.
discoverer: Warters, 1961. T(J;2)OR38
genetics: Male lethal. cytology. T(l;2)20B;S0A.
origin: X ray induced, discoverer: Warters, 1961.

discoverer: Warters, 1961. genetics: Male lethal; lethal originated after trans-
genetics: Male lethal. Male hyperploid for the location.
2RDXP element probably survives. T(J;2)OR52
T(1;2)OR39 cytology: T(1;2)4F;41A.
cytology: T(1;2)1D;46E. origin: X ray induced.
origin: X ray induced in y. discoverer: Warters, 1961.
genetics: Male viable but sterile. T(1;2)ORS4
T(1;2)OR41 cytology: T(1;2)19E;32E.
cytology: T(1;2)12D;25E. origin: X ray induced.
discoverer: Warters, 1961. genetics: Male viable with slightly curled wings;
genetics: Male lethal. sterile.
T(1;2)OR42 T(1;2)ORSS
cytology: T(1;2)12F;58F. cytology: T(l;2)20A;44D.
genetics: Male lethal. genetics: Male viable but sterile.
T(1;2)OR43
cytology: T(l;2)15E;40D. T(1;2)OR56
origin: X ray induced, cytology: T(1;2)18F;47D.
discoverer: Warters, 1961. origin: X ray induced,
genetics: Male viable but sterile, with small rough discoverer: Warters, 1961.
eyes; perhaps mutant for un. genetics: Male lethal.
T(1;2)OR44 T(1;2)OR58
cytology: T(1;2)16F;28F. cytology: T(1;2)11B;4O-41 + T(l;2)19C;30B; position
origin: X ray induced. of right breakpoint in chromosome 2 with respect to
discoverer: Warters, 1961. centromere not determined.
genetics: Male viable but sterile. new order: 1 - 11B|4O - 30B|l9C - 11B|41 - 60;
T(1;2)OR45 20 - 19C|3OB - 21.
cytology: T(1;2)7D;40-41; position of breakpoint in origin: X ray induced.
chromosome 2 with respect to centromere not deter- discoverer: Warters, 1961.
mined. genetics: Male lethal. Male hyperploid for 2LDXP
origin: X ray induced. element survives and has outstretched wings.
genetics: Male viable but sterile. cytology: T(1;2)19E:57B.
T(J;2)OR47 origin: X ray induced,
cytology: T(1;2)19E;53B. discoverer Warters, 1961.
origin: X ray induced, genetics: Male lethal. Male hyperploid for the
discoverer: Warters, 1961. 2RDXP element survives.
T(1;2)OR48 T(1;2)OR60
cytology: T(1;2)17A;31F. cytology: T(l;2)10A;52D.
discoverer. Warters, 1961. discoverer Warters, 1961,
genetics: Male viable but sterile. genetics: Male viable but sterile.
T(1;2)OR49 T(1;2)OR61
cytology. T(l;2)20;40-41; position of breakpoints cytology: T(1;2)18D;31F,
with respect to centromeres not determined. origin: X ray induced,
origin: X ray induced, discoverer Warters, 1961.
genetics: Male viable but sterile. T(1;2)OR62
T(1;2)OR5Q cytology: T(l;2)8F;17F;40-41; position of breakpoint
cytology: T(1;2)19C;33F. in chromosome 2 with respect to centromere not de-
origin: X ray induced, termined,
discoverer Warters, 1961. new order: 1 — 8 F | 4 O — 60;
genetics: Male viable and fertile. Homozygous fe- 20 - 17FJ8F - 17F]40 - 21.
male weakly viable. Tentative.
TO;2)OR5J origin: X ray induced,
cytology: T(1;2)8D;41F. discoverer; Warters, 1961.
origin: X ray induced. genetics: Male lethal.

cytology: T(1;2)11A;53F. genetics: Male lethal.
origin: X ray induced, T(1;2)OR78
discoverer: Waiters, 1961. cytology: T(l;2)19E;30B.
T(J;2)OR65 discoverer: Warters, 1961.
cytology: T(l;2)7A;40. genetics: Male viable but sterile. Male hyperploid
origin: X ray induced. for 2LDXP element survives and has outstretched
discoverer: Warters, 1961. wings.
genetics: Male lethal. T(1;2)OR82
T(1;2)OR66 cytology: T(l;2)13B;30B.
cytology: T(1;2)8B;23C. origin: X ray induced.
genetics: Male viable and fertile. T(1;2)OR83
T(1;2)OR67 cytology: T(1;2)12A;22B.
cytology: T(1;2)12B;41 + T(l;2)20A;45B. origin: X ray induced.
new order: 1 - 12B|41 - 45B|20A; discoverer Warters, 1961.
21 - 41 |l2B - 20A|45B - 60. genetics: Male lethal.
Tentative. T(1;2)OR84
origin: X ray induced. cytology: T(1;2)3C;38E.
discoverer: Warters, 1961. origin: X ray induced in y.
genetics: Variegated for a lethal. Male fertile. discoverer: Warters, 1961.
T(1;2)OR68 genetics: Male viable but sterile, with either a
cytology: T(1;2)16E;41 + Tp(2)25E;33A;40. normal Y or Bsw+Y.
new order: 1 - 16E|41 - 40|(25E - 33A)|40 - T(1;2)OR85
33A|25E - 21; cytology: T(l;2)10;38.
2 0 - 16E|41 - 60. origin: X ray induced.
Tentative. discoverer: Warters, 1961.
genetics: Male viable but sterile. cytology: T(1;2)11A;32B.
cytology: T(1;2)3C;41C. genetics: Male viable but sterile.
origin: X ray induced in y. *T(l;2)ret: Translocation(l;2) reticulated
discoverer: Warters, 1961. cytology: T(l;2)20A5-B2;2R.
genetics: Variegated for a lethal; male sterile with origin: Induced by L-p-NN-di-(2chloroethyl)amino-
either a normal Y or Bsw+Y. phenylalanine (CB. 3025).
TO;2)OR72 discoverer: Fahmy, 1953.
cytology: T(l;2)l 9E;29F + In(2LR)24F;54B. references: 1958, DIS 32: 73.
new order. 1 - 19E|29F - 54B|24F - 21; genetics: Associated with ret; male sterile.
20 - 19E|29F - 24F|54B - 60. T(T;2)sc19: Transhcation(l;2) scute
origin: X ray induced. cytology: T(l;2)lBl-2;lB4-7;25-26; breaks in X esti-
discoverer. Warters, 1961. mated from fig. 1 of Muller and Prokofyeva (1934,
genetics: Male viable but sterile. Male hyperploid Dokl. Akad. Nauk. n.s. 4: 74-83), but the left
for the 2RD2LMXP element may survive. break, which genetically is to the left of y, is in-
T(1;2)OR73 consistent with the cytological location of y in re-
cytology: T(1;2)19E;57E. gion 1A5-8. Break in chromosome 2 estimated from
origin: X ray induced. position of y+ 1—2 units to the right of dp.
discoverer Warters, 1961. new order 1A - lBl|lB7 - 20;
genetics: Male viable bat sterile. Male hyperploid 21 - 25|(1B2 - 1B4)|26 - 60.
for the 2RDXP element survives. origin: X ray induced,
T(1;2)OR74 di scoverer League.
cytology: T(t;2)l9E;56C. references: Muller, 1935, Genetica 17: 237-52.
origin: X ray induced, genetics: Mutant for sc. A small subterminal piece
discoverer: Warters, 1961. of X is inserted into 2L 1 or 2 units to the right of
genetics: Male viable but sterile. Male hyperploid dp. The two halves of the translocation are re-
for 2RDXP element survives. coverable independently as Df(l)sc19 = Df(l)lBl-
T(1;2)OR75 2;lB4-7 and Dp(l;2)mcl* = Dp(l;2)lBl-2;lB4-
cytology: T(1;2)12E;32B. 7;25-26. Dt(l)scl9 is deficient few y, ac, ac, and
origin: X ray induced. l(l)mc but not l(l)Jl, cm, or M(l)Bld; it is male
lethal but survives in the heterozygous female. T(1;2)SPW

Dp(l;2)sc19 carries, in addition to sc19, normal cytology: T(l;2)10;50.
alleles of y, ac, and l(l)sc; it is viable homozy- origin: Gamma ray induced.
gous and does not affect crossing over in 2L. discoverer: Lindsley and Musatti, 1961.
*T(l;2)sc"s genetics: Male viable but sterile.
cytology: T(1;2)1A6-B1;25F; inferred from Goldat's T(1;2)SP16
fig. 3. cytology: T(l;2)20;40-41; position of breakpoints
origin: X ray induced derivative of sc6. with respect to centromeres not determined.
discoverer: Goldat. origin: Gamma ray induced.
references: 1936, Biol. Zh. (Moscow) 5: 803—12. discoverer: Lindsley and Musatti, 1961.
genetics: Mutant for sc. genetics: Male viable but sterile.
*T(l;2)sc260.J7 T(1;2)SP18
cytology: T(1;2)1B2-3;31C. cytology: T(1;2)1A;56A.
origin: X ray induced. origin: Gamma ray induced in y w.
discoverer: Sutton, 39d. discoverer Lindsley and Mussatti, 1961.
references: 1943, Genetics 28: 210—17. genetics: Male viable but sterile.
genetics: Mutant for sc but not y, ac, or svr. T(1;2)SP19
*T(l;2)sc260-26 cytology: T(l;2)20;40-41; position of breakpoints
cytology: T(l;2)lB4-5;41F2-3;58B2-3 + In(2LR)27D2- with respect to centromeres not determined.
3;41A. origin: Gamma ray induced.
new order: 1A - 1B4|41F3 - 58B2|lB5 - 20; discoverer: Lindsley and Musatti, 1961.
21 - 27D2|41A - 27D3|41A - genetics: Male viable but sterile.
41F2|58B3 - 60. T(l-2)SP20
origin: X ray induced. cytology: T(l;2)20;40-41; position of breakpoints
discoverer: Sutton, 391. with respect to centromeres not determined.
references: 1943, Genetics 28: 210—17. origin: Gamma ray induced.
genetics: Mutant for sc but not y, ac, or svr. discoverer: Lindsley and Musatti, 1961.
*T(l;2)sc260.27 genetics: Male viable but sterile.
cytology: T(1;2)15E;33-34;57B-C + Dp(l;f)lA8- T(1;2)SP31
B1;19F. cytology: T(l;2)20;56B.
new order: 1A1 - lA8|l9F - 20; origin: Gamma ray induced.
21 - 33|15E - 19F|1B1 - 15E|57B - discoverer: Lindsley and Musatti, 1961.
34|S7C - 60. genetics: Male viable but sterile. Male hyperploid
origin: X ray induced, for the 2RDXP element survives.
discoverer Satton, 391.
references: 1943, Genetics 28: 210—17. *T(J;2)SP33
genetics: Mutant for sc but not y, ac, or svr. Male cytology: T(l;2)14;41.
sterile. Dp(l;i)sc260-2 7 =Dp(l;f)lA8-Bl;19F seg- origin: Gamma ray induced.
regates free from translocation; carries normal discoverer: Lindsley and Musatti, 1961.
alleles of y and ac. genetics: Male viable but sterile.
T(1;2)SP36
T(l;2)scS2: Translocation(1 ;2) scute of cytology: T(l;2)20;40-41; position of breakpoints
Sinitskaya with respect to centromeres not determined.
cytology: T(1;2)1B4-7;6OC-E (inferred from genetic origin: Gamma ray induced,
results). discoverer: Lindsley and Musatti, 1961.
discoverer Sinitskaya, 1934. genetics: Male viable but sterile.
genetics: Mutant for sc. X chromosome broken to the T(1;2)SP42
right of l(l)sc in same place as right breakpoint of cytology: T(l;2)20;40-41; position of breakpoints
T(l;2)8c19 and 2R broken between sp and M(2)c with respect to centromeres not determined.
(Muller). Aneuploid segregants XD2P and 2RDXP origin: Gamma ray induced.
should survive. discoverer: Lindsley and Musatti, 1961.
T(1;2)SP1: Translocation(1;2) from $db Paulo genetics: Male viable but sterile.
cytology: T(1;2)8B;41. T(1;2)SP43
origin: Gamma ray induced. cytology: T(l;2)16A;60C.
discoverer: Lindsley and Musatti, 1961. origin: Gamma ray induced.
genetics: Variegated for a lethal; male sterile. discoverer: Lindsley and Musatti, 1961.
TO;2)SP4 genetics: Male viable but sterile.
cytology: T(l;2)20;40-41; position of breakpoints T(1;2)SP48
with respect to centromeres not determined. cytology: T(1;2)15F;35A.
origin: Gamma ray induced. origin: Gamma ray induced.
discoverer Lindsley and Musatti, 1961. discoverer: Lindsley and Musatti, 1961.
genetics: Male viable but sterile. genetics: Male viable but sterile.
T(1;2)SP49 discoverer: Lindsley and Musatti, 1961.

cytology: T(l;2)12;40-41; position of breakpoint in genetics: Male viable but sterile.
chromosome 2 with respect to centromere not deter- T(1;2)SP71
mined. cytology: T(l;2)20;40-41; positions of breakpoints
origin: Gamma ray induced. with respect to centromeres not determined.
genetics: Male viable but sterile. discoverer: Lindsley and Musatti, 1961.
T(l;2)SP50 genetics: Male viable but sterile.
cytology: T(l;2)20;29-30. *T(1;2)SP75
origin: Gamma ray induced. cytology: T(1;2)8C;35D.
discoverer: Lindsley and Musatti, 1961. origin: Gamma ray induced in y w.
genetics: Male viable but sterile. Male hyperploid discoverer: Lindsley and Musatti, 1961.
for 2LDXP survives. genetics: Male viable but sterile.
T(l;2)SPSl T(1;2)SP77
cytology: T(l;2)20;40-41; position of breakpoints cytology: T(1;2)9A;41.
with respect to centromeres not determined. origin: Gamma ray induced.
origin: Gamma ray induced. discoverer: Lindsley and Musatti, 1961.
discoverer: Lindsley and Musatti, 1961. genetics: Male viable but sterile.
genetics: Male viable but sterile. T(1;2)SP81
T(1;2)SP52 cytology: T(l;2)20;24F-25A.
cytology: T(1;2)12E;57F. origin: Gamma ray induced.
discoverer: Lindsley and Musatti, 1961. genetics: Male viable but sterile. Male hyperploid
genetics: Male viable but sterile. for 2LDXP element survives.
T(1;2)SP5S T(1;2)SP84
cytology: T(1;2)1A;41 + T(l;2)4B;30B + cytology: T(1;2)4C;42C.
In(l)12D;14B. origin: Gamma ray induced in y w.
new order: 1A|41 - 30B|4B - 1A|41 - 60; discoverer: Lindsley and Musatti, 1961.
20 - 14B|l2D - 14BJ12D - 4B|30B - 21. genetics: Male viable but sterile.
origin: Gamma ray induced in y w. T(1;2)SP87
discoverer. Lindsley and Musatti, 1961. cytology: T(l;2)9A4-Bl;58A3-4.
genetics: Male lethal. origin: Gamma ray induced.
T(1;2)SP58 discoverer Lindsley and Musatti, 1961.
cytology: T(l;2)10A;34A. genetics: Male viable but sterile.
discoverer: Lindsley and Musatti, 1961. T(1;2)SP88
genetics: Male viable but sterile. cytology: T(l;2)20;32F-33A.
T(1;2)SP60 origin: Gamma ray induced.
cytology: T(1;2)17E;35A. discoverer Lindsley and Musatti, 1961.
origin: Gamma ray induced. genetics: Male viable but sterile. Male hyperploid
discoverer: Lindsley and Musatti, 1961. for the 2LDXP element survives.
genetics: Male viable but sterile. *T(1;2)SP89
T(1;2)SP61 cytology: T(1;2)4E;35A.
cytology: T(1;2)18F;47D, origin: Gamma ray induced in y w.
origin: Gamma ray induced, discoverer Lindsley and Musatti, 1961.
discoverer: Lindsley and Musatti, 1961. genetics: Male viable but sterile. May be mutant for
genetics: Male viable but sterile,
T(1;2)SP64 T(1;2)SP93
cytology: T(1;2)3C;28C. cytology: T(1;2)18C-D;22A-B.
origin: Gamma ray induced toy w. origin: Gamma ray induced,
discoverer Lindsley and Musatti, 1961. discoverer Lindsley and Musatti, 1961.
genetics: Male viable but sterile, with either a genetics: Male viable but sterile. Male hyperploid
normal Y or Bsw+Y. for the 2LDXP element survives.
T(1;2)$P67 T(1;2)SP94
cytology: T(l;2)20;40-41; positions of breakpoints cytology: T(1;2)14B-C;23F.
with respect to centromeres not determined, origin: Gamma ray induced,
origin: Gamma ray induced. discoverer. Lindsley and Musatti, 1961.
discoverer Lindsley and Musatti, 1961. genetics: Male viable but sterile.
genetics: Male viable but sterile. T(1;2)SP96
T(1;2)SP69 cytology: T(l;2)20;40-41; positions of breakpoints
cytology: T(1;2)7C;41. with respect to centromeres not determined.
origin: Gamma ray induced. origin: Gamma ray induced.
discoverer Lindsley and Musatti, 1961. references: Lefevre, 1953, DIS 27: 57.
genetics: Male viable but sterile. genetics: Variegated for w. The segregant
*T(1;2)SP97 Dp(l;2)W"53° = Dp(l;2)3B2-Cl;3C9-Dl;40-41 sur-
cytology: T(1;2)9E-F;35A-B. vives and is duplicated for the loci of w, rst, and
origin: Gamma ray induced in. y w. TV. Df(l)wm^3a - Df(l)3B2-Cl;3C9-Dl survives as
discoverer: Lindsley and Musatti, 1961. Notch female; deficient for w, rst, and N.
genetics: Male viable but sterile. *TO;2)w">S3*
W;2)SP102 cytology: T(l;2)3C3-4;20A2-3;58F8-59Al.
cytology: T(1;2)16A;41. new order: 1 - 3C3|58F8 - 21;
origin: Gamma ray induced. 20F - 20A3|3C4 - 20A2|59Al - 60.
discoverer: Lindsley and Musatti, 1961. origin: Neutron induced,
geneticsf Male viable but sterile. discoverer: Mickey, 53ell.
*T(1;2)SP106 synonym: T(X'2)Inx *nd 3.
cytology. T(l;2)6B;40. references: 1963, DIS 38: 29.
origin: Gamma ray induced in y w. genetics: Variegated for w.
discoverer: Lindsley and Musatti, 1961. *T(l;2)w>»2S8-34
genetics: Male viable but sterile. cytology: T(1;2)3C3-5;41A (Demerec and Hoover).
T(1;2)SP11O origin: X ray induced.
cytology: T(1;2)13A;57E. discoverer: Demerec, 38b.
origin: Gamma ray induced. genetics: Variegated for w but not rst, fa, or dm.
discoverer Lindsley and Musatti, 1961. Male viable.
genetics: Male viable but sterile. *T(l;2)w«> 258-3 6
T(1;2)SP111 cytology: T(l;2)3C6-7;4C2-3;41A-B;41F5-6 (Demerec
cytology: T(l;2)20;40-41; position of breakpoints and Hoover).
with respect to centromeres not determined. new order. 1 - 3C6J(41B - 41F5)|4C3 - 20;
origin: Gamma ray induced. 21 - 41A|(3C5 - 4C2)|41F6 - 60.
discoverer: Lindsley and Musatti, 1961. Insertions said to be in inverted order but not spec-
genetics: Male viable but sterile. ified with respect to centromere or numerical order.
*T(J;2)Sy: Translocation(l;2) Stubby origin: X ray induced.
origin: Spontaneous. discoverer: Demerec, 38b.
discoverer: Ives, 34J31. references: Sutton, 1940, Genetics 25: 534—40
genetics: Associated with Sy. Male sterile. Prob- (fig-)-
ably reciprocal translocation with breaks near the genetics: Variegated for w and rat but not pn, fa, or
base of X and 2L. dm. Male viable. Cytology predicts that each ele-
*T(l;2)v267'4: Translocation(l;2) vermilion ment of the translocation should survive as aneu-
cytology: T(l;2)llA7-8;36 (Sutton). ploid but not so recorded.
origin: X ray induced. *T(7;2)w»258-37
discoverer: Hoover, 35i. cytology: T(I;2)3C3-4;40-41A (Sutton),
genetics: Mutant for v (breakpoint not at v locus). origin: X ray induced.
Semilethal. ras, dwx, sbr, m, dy, and fw not af- discoverer Demerec, 33j.
fected. genetics: Variegated for w but not kz, rat, fa, or dm.
T(l;2)w+sib?: see T(l;2)51b *T(l;2)w°2S8-39
*T(l;2)w 13G2: Translocation(l;2) white cytology: T(l;2)3C3-5;40E-F (Demerec and Hoover),
cytology: T(1;2)3C3-5;56F; also inversion in 2R. origin: X ray induced,
origin: X ray induced. discoverer: Demerec, 38e.
discoverer: Gans. genetics: Variegated for w but not pa, r&t, fa, or dm.
genetics: Variegated for w. Male viable.
*T(l;2)wmS2b12: Translocation(l;2) white-mottled
cytology: T(l;2)lE5-Fl;3C3-4;20B;40-41. cytology: T(1;2)3C3S;41 (Demerec and Hoover),
new order: 1A - 1E5|2OB - 3C4J20B - 20F; origin: X ray induced.
21 - 4O|(1F1 - 3C3)|41 - 60. discoverer Demerec, 38e.
origin: X ray induced inln(l)rst3 -In(l)3C3-4;20B. genetics: Variegated for w and rst but not pn, kz, fa,
discoverer: Ratty, 52bl2. or dm.
references: Lefevre, 1953, DIS 27: 57. *T(7;2>w"»OI; Translocation(l;2) white-mottled of
genetics: Variegated for w. Dp(l;2)w^^2bl2 « Dubinin
Dp(l;2)lES-Fl;3C3-4;40-41 survives. cytology: T(l;2)3B;19-20;21F.
new order: 1 — 3BJ21F — 60;
cytology: T(l;2)3B2-Cl;3C9-Dl;40-41. 20|3B - 19J21F - 21A.
new order: 1 - 3B2J3D1 - 20; origin: X ray induced,
21 - 4O|(3C1 - 3C9)|41 - 60. discoverer: Dubinin.
origin: X ray induced, references: Sachorov, 1936, Biol. Zh. (Moscow) 5:
discoverer: P. Farnsworth, 53a4. 293-302.
*T(l;2)wvD4: Translocation(1 ;2) white-variegated origin: X ray induced.

of Demerec discoverer: Roberts, 1965.
cytology: T(1;2)3D6-E1;4OF (Schultz). genetics: Male sterile. Recombination reduced in
origin: X ray induced. 2L.
discoverer: Demerec, 33k2. *TO;2;3)ct268-40: Translocation(l;2;3) cut
genetics: Variegated for A?, rst, w, and dm. X/Y cytology: T(l;2;3)7D2-3;10A5-6;21B-C;28-29;40-
male survives only rarely as rst with mottled eye 41;75B-C;87D;88C;92; new order not determined.
color; X/Y/Y male more viable, slightly rst, and origin: X ray induced.
sterile. Variegation for It in X/X/Y female. discoverer: Demerec, 39k.
T(J;2)w-ec: Translocation(l;2) white-echinus references: Sutton, 1940, Genetics 25: 534-40
cytology: T(1;2)3C1-2;3E7-8;37D. (fig-)-
new order: 1 - 3Cl|3E8 - 20; genetics: Mutant at ct but not scp, cm, sn, v, sbr,
21 - 37DJ(3C2 - 3E7)|37D - 60. dy> £> ty> na, ph ^d, or me. Male lethal.
origin: X ray induced. T(l;2;3)Din: Translocation(l;2;3) Dinty
references: Lefevre and Wilkins, 1966, Genetics 53: cytology: T(1;3)3C;63A + T(2;3)39D;73A (Lindsley).
175-87. new order: 1A - 3C|63A - 73A|39D - 60;
genetics: T(l;2)w-ec male is phenotypically white 20 - 3C|63A - 61;
and echinus; TV not affected. Does not complement 21 - 39D|73A _ 100.
with w"P. Gives rise to Df(l)w-ec. origin: X ray induced.
*T(J;2)y260-13: Translocation(l;2) yellow discoverer: Braver, 55a.
cytology. T(1;2)1A4-5;36D. references: 1955, DIS 29: 70.
origin: X ray induced. Pollock, 1963, DIS 38: 50.
discoverer: Sutton, 1939. genetics: Associated with Din. Male viable and
references: 1943, Genetics 28: 210-17. fertile. The two translocations should be easily
genetics: Mutant for y. separable, and Din is, in all probability, associ-
*T(1 ;2)yvh Translocation(l;2) yellow-variegated ated with only one.
cytology: T(1;2)1A;39. *T(l;2;3)l-v2U: Translocation(l;2;3) lethal-
origin: X ray induced. variegated
discoverer. Schultz, 33all. origin: X ray induced.
genetics: Variegated for y. discoverer: Lindsley, Edington, and Von Halle.
origin: X ray induced. genetics: Variegated for a lethal; male sterile.
discoverer: Imazumi. T(l;2;3)l-v459
references: 1961, DIS35: 87-88. cytology: T(l;2;3)3D-F;XR;50;80-81.
1962, DIS 36: 80. new order: 1A - 3D|50 - 21;
1962, Cytologia 27: 212-28 (fig.). |3F - 20F-;
genetics: Distal one-third of 2L appended to X chro- 20F-80 - 61;
mosome as short arm. Also T(2;3) with 2R broken 6 0 - 50-81 - 100.
between en and vg and 3L broken between se and Tentative.
st. Male lethal in embryo. Postulated that centromere of chromosome 3 split
W;2;3)220 or double with one half capped by 2RD and the
cytology: T(1;2;3)14A;50A;75C. other by XRD. Xp in the form of a ring.
new order: 1 - 14A|50A - 21; origin: X ray induced.
20 - 14A|75C - 61; discoverer: Lindsley, Edington, and Von Halle.
60 - 50A|75C - 100. references: 1960, Genetics 45: 1649-70.
origin: X ray induced. genetics: Variegated for a lethal; male fertile.
discoverer: Lindsley, Edington, and Von Halle, *T(1;2;3)N264.74: Translocation(h2;3) Notch
references: 1960, Genetics 45: 1649—70. cytology: T(l;2;3)3C10-ll;20D-E;40C-D;92E6-8;
genetics: Male viable but sterile. 20D-E break claimed to be to the left of the nucle-
T(l;2;3)C232: Translocation(l;2;3) Crossover olus organizer (Sutton).
suppressor new order: 1 - 3C10|40D - 60;
cytology: T(2;3)35D;71E; additional presence of 20F - 20E|40C - 21;
T(l;2)20;40-41 or T(l;3)20;80-81 inferred from 61 - 92E6|20D - 3C11 |92E8 - 100.
genetic data, origin: X ray induced.
discoverer: Roberts, 1965. references: Sutton, 1940, Genetics 25: 534—40
genetics: Male viable and fertile; homozygous fe- (fig.)-
male lethal. Recombination reduced in 2L. genetics: Variegates for w, rst, and N but not kz,
W;2;3)C312 pn, or dm.
cytology: T(2;3)32C;87E; additional presence of *T(l;2;3)N264-87
T(t;2)20;40-41 or T(l;3)20;80-81 inferred from cytology: T(l;2;3)3C7-9;10A2-Bl;45F-46A;59F'
genetic data. 60A;97C-D;100E-F (Sutton).
new order: 1 - 3C7|97D - 100E|59F - 46A|lOBl - new order: 1 - 14C|87B - 61;

20; 20 - 14C|27D - 21;
21 - 45F|3C9 - 10A2|60A - 60F; 60 - 27D|87B - 100.
61 - 97C|lOOF. origin: X ray induced.
discoverer: Demerec, 39j. genetics: Male lethal.
references: Sutton, 1940, Genetics 25: 534—40. T(1;2;3)0R24
genetics: Carries a mutant allele of/V but normal cytology: T(1;2)14B;39D + T(l;3)2C;80C +
alleles of w, rst, and dm. T(1;3)19;87A,
new order: 1 - 2C|80C - 87A|l9 - 14BJ39D - 21;
T(1;2;3)OR9: Translocation(l;2;3) from Oak Ridge 20 - 19|87A - 100;
cytology: T(l;2;3)19-20;49F;81F. 60 - 39D|l4B - 2C|80C - 61.
new order: 1 — 19|81F — 61; origin: X ray induced in y.
20|49F - 60; discoverer: Warters, 1961.
21 _ 49F|8lF - 100. genetics: Male lethal.
origin: X ray induced, T(J;2;3)OR25
discoverer: Warters, 1961. cytology: T(l;2;3)19E;29B;80-81; position of break-
genetics: Male lethal. point in chromosome 3 with respect to centromere
T(1;2;3)OR10 not determined; therefore new order ambiguous.
cytology: T(l;2;3)18A;41;73F. new order: 1 - 19EJ80 - 100;
new order: 1 - 18A|73F - 100; 20 - 19E|29B - 21;
20 - 18A|41 - 60; 60 - 29B|80 - 61.
21 - 4l|73F - 61. Tentative.
genetics: Male viable but sterile. genetics: Male viable but sterile. Hyperploid male,
T(1;2;3)OR12 presumably carrying 2LDXP, survives.
cytology: T(1;2)3A;41 + T(1;3)7E;78F. T(l;2;3)0R26
new order: 1 - 3A|41 - 21; cytology: T(1;2)2D;56F + T(1;3)3F;96B,
20 - 7E|78F - 61; new order: 1 - 2D|56F - 21;
60 - 4l|3A - 7E|78F - 100. 20 - 3F|96B - 100;
origin: X ray induced in y. 60 - 56F|2D - 3FJ96B - 61.
discoverer: Warters, 1961. origin: X ray induced in y.
genetics: Male viable but sterile, with either a discoverer: Warters, 1961.
normal Y or Bsw+Y. genetics: Male viable and fertile.
T(l;2;3)0R31
T(7;2;3)ORU cytology: T(l;3)20;92A + T(2;3)38D;87E.
cytology: T(1;2;3)5E;21D;62C. new order. 1 - 20J92A - 87E|38D - 60;
new order: 1 — 5E|21D — 60; 20J92A - 100;
20 - 5E|62C - 61; 21 - 38DJ87E - 61.
21A - 21D|62C - 100. origin: X ray induced,
discoverer: Warters, 1961. genetics: Male viable but sterile. Male hyperploid
genetics: Male viable and fertile. Homozygous for 3RDXP element survives.
female viable. T(1;2;3)0R34
T(1;2;3)ORU cytology: T(1;3)18F;84B + T(2;3)28B;75F +
cytology: T(1;2)1A;57D + T(l;3)20;72E. T(2;3)44C;63A.
new order: 1A|57D — 21; new order: 1 - 18FJ84B - 75F|28B - 21;
20|72E - 61; 20 - 18F|84B - 100;
60 - 57D|lA - 20|72E - 100. 60 - 44C|63A - 75FJ28B - 44C|63A -
origin: X ray induced in y. 61.
discoverer: Warters, 1961. Tentative.
genetics: Male viable but sterile. origin: X ray induced.
T(1;2;3)OR17 discoverer: Warters, 1961.
cytology: T(1;2;3)20;40-41;61F; neither breakpoints genetics: Male lethal.
in X and 2 with respect to centromere nor new *T(l;2;3)sc260.l8: Translocation(l;2;3) scute
order determined. cytology: T(1;2)1A6-B1;4W-E + T(1;3)7A2-B1;®QC.
origin: X ray induced. new orden 1A1 - 1A6|41D - 21;
discoverer: Warters, 1961. 20 - 7B1J80C - 61;
genetics: Male viable but sterile. 60 - 41B|lBl - 7A2J8QC - 100.
T(1;2;3)OR23 origin: X ray induced.
cytology: T(l;2,3)14C;27D;87B. discoverer. Suttoa, 39d.
references: 1943, Genetics 28: 210-17. new order: 1 - 10E|40 - 60D|64D - 61;
genetics: Mutant for sc but not y, ac, or svr. Male 21 - 11A|6OD - 60F;
sterile. 21 - 40|64D - 100.
*T(l;2;3)sc260.29 origin: Gamma ray induced.
cytology: T(1;2;3)1A6-B1;22A-B;34A-B;75C-E. discoverer: Lindsley and Musatti, 1961.
new order: 1A1 - 1A6|34A - 22B|34B - 60; genetics: Male viable but sterile.
2 0 - 1B1|75C - 61; T(l;2;3)$P40
21 -22AJ75E - 100. cytology: T(l;2;3)4-5;50A;80 + T(2;3)40;86.
origin: X ray induced. new order: 1 _ 4 J80 — 86|40 - 21;
discoverer: Sutton, 40a. 20 - 5|50A - 60;
references: 1943, Genetics 28: 210-17. 61 - 80|50A - 40|86 - 100.
genetics: Mutant for sc but not y, ac, or svr. Tentative.
origin: Gamma ray induced in y w.
*T(l;2;3)scPi: Translocation(l;2;3) scute o/ discoverer: Lindsley and Musatti, 1961.
Panshin genetics: Variegated for a lethal. Male sterile.
discoverer: Panshin, 1934. W;2;3)SP57
genetics: Mutant for sc. cytology: T(l;2;3)20;40-41;75A; breakpoint in chro-
mosome 2 inferred from genetic data.
T(1;2;3)SP3: Translocation(l;2;3) from Sdb Paulo new order: 1 — 20|75A — 100;
cytology: T(l;2;3)20;23A-B;96B. 20|40 - 21;
new order: 1 — 20|96B - 61; 60 -40|75A - 61.
20|23A - 21; Tentative.
60 - 23BJ96B - 100. origin: Gamma ray induced.
discoverer: Lindsley and Musatti, 1961. genetics: Male viable but sterile.
genetics: Male lethal. Male hyperploid for 2LDXP T(1;2;3)SP65
element survives. cytology: T(1;2;3)18A;39E;76A.
T(1;2;3)SP5 new order: 1 - 18A|76A - 100;
cytology: Six-break rearrangement with two breaks 20 - 18A|39E - 21;
in 2R, one in 3L, and two in 3R. 60 - 39E|76A - 61.
new order: XD|2RP - 2L; origin: Gamma ray induced.
| discoverer: Lindsley and Musatti, 1961.
2RDJ3LP - 3RP|3RD. genetics: Male viable but sterile.
discoverer: Lindsley and Musatti, 1961. *T0;2;3)w>»2SB-44: Translocation(l;2;3) white-
genetics: Male sterile. mottled
T(1;2;3)SP6 cytology: T(1;2;3)3C3-4;4D2-E1;56E1-F1;8OD
cytology: T(l;2;3)20;40-41;80-81; neither position of (Sutton).
breakpoints with respect to centromeres nor new new order: 1 - 3C3|80D - 100;
order determined. 20 -4E1J80D - 61;
origin: Gamma ray induced. 21 - 56El|(3C4 - 4D2)|56F1 - 60.
discoverer Lindsley and Musatti, 1961. origin: X ray induced.
genetics: Male viable but sterile. discoverer: Demerec, 38k.
T(1;2;3)SP8 genetics: Variegated for w but not pn, rst, or la.
cytology: T(l;2;3)5;17F;44B;90A. T(l;2;3)wm258-44 may be separated into
new order. 1 - s|l7F - 20; T(l;3)vi^3S8-44 „ T(l;3)3C3-4;4D2-El;8QD, which
21 - 44B|5 - 17F|90A - 100; is deficient for 3C4 through 4D2 (i.e.,
61 - 90A|44B - 60. Df(l)w^258-44 = Dt(l)3C3-4;4D2-El), and
origin: Gamma ray induced, Dp(t;2)wo>2S8-44 = Dp(l;2)3C3-4;4D2-El;56El-Fl,
discoverer: Lindsley and Musatti, 1961. which is duplicated for the same region. The defi-
genetics: Male viable but sterile. ciency includes the loci of fa, dm, M(1)3E, ec,
*T(1;2;3)$P25 M(1)4BC, W, peb, and rb but not rat or rg. 3C3-4
cytology: T(l;2;3)19;54;86. breakpoint inconsistent with genetic data on rst.
new order: 1 — 19J54 - 21; * should be viable.
2 0 - 19|86 - 100;
6 0 - 5 4 | & 6 - 61. *T(l;2;3)wy274.2: Translocation(l;2;3) wavy
origin: Gamma ray induced. cytology: T(l;2)8F-9A;20A-B;26B-D + T(1;3)11D-
discoverer Lindsley and Musatti, 1961. E;65C-D (Sutton).
genetics: Male viable but sterile. Male hyperploid new order: 1 — 8FJ26D — 60;
for 3RDXP apparently survives. 20F - 20B|9A _ 11D165C - 61;
T(1;2;3)SP29 21 - 26BJ20A - 11E|65D - 100.
cytology: T(1;2;3)10E-11A;40;60D;64D. origin: X ray induced.
discoverer: Demerec, 34a. origin: X ray induced.

genetics: Mutant for wy but not iw, dy, g, or s. Male discoverer: Warters, 1961.
lethal. genetics: Male viable but sterile. Produces a hyper-
ploid female that may carry the 4DXP element.
T(l;2;3;4)l-v454: Translocation(J;2;3;4) lethal- *TC7;2;4)w"6wV; Translocation(l;2;4) white-
variegated mottled brown-Variegated
cytology: T(1;2;3)12B;22-23;81 + T(2;4)44F;101F. cytology: T(l;2)12F3-4;59C4-5 + T(1;4)3C3-
new order 1 — 12B|81 — 61; 4;101E4-5.
20 - 12B|22 - 21; new order: 1 - 3C3|l01E4 - 101A;
60 - 44F|l01F - 101 A; 20 - 12F4|59C5 - 60;
100 - 81 (23 - 44F|l01F - 102. 21 - 59C4|l2F3 - 3C4|l0lE5 - 102F.
origin: X ray induced. origin: Neutron induced.
discoverer: Lindsley, Edington, and Von Halle. discoverer: Mickey, 53fl5.
genetics: Associated with I(l)vf54# Male sterile. genetics: Variegated for w. Also claimed to varie-
gate for bw, which is unusual since the T(l;2) is
*T(1;2;4)A12: Trarrslocation(l;2;4) from Austin completely euchromatic.
cytology: T(l;2;4)lB-C;7A;7B;13Bl-5; 101-102; *T(1;2;4)wyD2: Translocation(l;2;4) white-
breakpoints in chromosomes 2 and 4 not determined variegated of Demerec
(Mackensen, 1935, Texas Univ. Publ. 4032: fron- cytology: T(l;2;4)3C4-5;18F;38;101A-C (Schultz).
tispiece). new order: 1 - 3C4|l01C - 102F;
new order: 1A - 1 B | 1 3 B 5 - 20; 20 - 18F|3C5 - 18F|38 - 21;
21 - ?|(7A - 7B)|? - 60; 60 - 38|l01A.
10l|((lC - 7A)|(7B - 13Bl))|lO2. Tentative.
origin: X ray induced. origin: X ray induced,
discoverer: Patterson, Stone, Bedichek, and Suche. discoverer: Demerec, 33k27.
references: Stone, 1934, Genetica 16: 506—19. genetics: Variegates for w but not rst in male and
Mackensen, 1935, J. Heredity 26: 163-74. for w and occasionally for rst in female. Absence
Patterson, Stone, and Bedichek, 1935, Genetics 20: of effect on ci a criterion for postulating break in
259-79. 4L. Fly hyperploid for the XD4P element survives.
1937, Genetics 22: 407-26.
genetics: A section from between sc and br on the *T(J;3)3
left to between g and sd on the right is inserted origin: X ray induced.
into chromosome 4. The ct locus but not cm, sn, or discoverer: Bonner, 1931.
oc is deleted from the insertion; i.e., Df(l)A12 = references: Dobzhansky, 1935, Z. Induktive
Df(l)7A;7B, and inserted into chromosome 2; i.e., Abstammungs- Vererbungslehre 68: 143—46.
Dp(l;2)A12 = Dp(l;2)7A;7B. Female hyperploid for genetics: X chromosome broken between rb and rg;
the XDXP element; i.e., Dp(l;t)A12 =Dp(l;f)lB- 3R broken to the right of ca. Male and homozygous
C;13Bl-5, survives and is claimed to be fertile. and heterozygous females viable and fertile. Cross-
Female hyperploid for XM; i.e., Dp(l;4)A12 - ing over in heterozygous female nearly absent in
Dp(l;4)lB-C;7A;7B;13Bl-S;101-102, occasionally left end of X; rises to about normal around ct; may
survives and is sterile. be increased at right end. Crossing over in chromo-
*T(1;2;4)N264.85: Translocation(l;2;4) Notch some 3 in translocation heterozygote normal be-
cytology: T(l;2;4)3B4-Cl;6A2-Bl;60A4-5;101F-102A tween e 8 and TO and reduced to two-thirds normal
[Sutton, 1940, Genetics 25: 534-40 (fig.)], between ro and ca. Male carrying the XD3P element
new order: 1 - 3B4|60A4 - 21; in place of a normal 3 nearly lethal; female has
2 0 - 6Bl|60A5 - 60F; narrow wings, occipital bristles, and branched pos-
101A - 1O1F|(3C1 - 6A2)|lO2A - 102F. terior veins. Crossing over between normal X chro-
origin: X ray induced, mosomes about one-third of normal at left end in
discoverer. Demerec, 39d. duplication-bearing female, but nearly normal to
genetics: Variegates for w, rst, fa, dim, rg, ex, cv, right of ct.
nix, and va but not pn, ec, bi, peb, or rb. Carries *T(1;3)S4a
normal alleie of ci (Stern). Dp(l;4)N264'85 - origin: X ray induced,
Dp(l;4)3B4-Cl;6A2-Bl;101F-102A viable in both discoverer: Lefevre, 54a4.
sexes but sterile in male. Complementary synonym: T(l;3)w+54*4,
D%ipja64~ss inviable. references: 1955, DIS 29; 73.
T(l;2;4)0R24: Translocation(l;2;4) from Oak genetics: A section of X chromosome including w+
Ridge but not Bpl inserted into chromosome 3.
cytology: T(1;2)3C;38A + T(1;4)11A;1O2C. *W;3)$4c
new order: 1 - 3C|38A - 60; origin: X ray induced.
2 0 - 11AJ102C - 102F; discoverer: L*fevre, 54c 10.
21 - 38AJ3C - 11A|1O2C - 101A. synonym; T
370 GEKETIC VARIATIONS OF DROSOPHILA MELANOGASTER
references: 1955, DIS 29: 73. references: Ogaki, 1960, DIS 34: 97.
genetics: Section of X chromosome including w+ 1960, Japan J. Genet. 35: 282.
inserted into chromosome 3. genetics: Position effect at B. Male sterile.
*T(1;3)65 T(l;3)Bs3i: Translocation(l;3) Bar-Super inserted
cytology: T(1;3)16-17;79D. in chromosome 3
origin: X ray induced. cytology: T(l;3)15F9-16Al;16A7-Bl;19-20;Y;66B13-
discoverer: Lindsley, Edington, and Von Halle. Cl (Muller; Lindsley); translocation between
references: I960, Genetics 45: 1649—70. Dp(l;l)B =Dp(l;l)15F9-16Al;16A7-Bl and chromo-
genetics: Male viable but sterile. some 3. X break can be shown genetically to sepa-
*TO;3)102 rate f+ from B and is assumed here to separate the
prig in: X ray induced. two halves of the Bar duplication.
discoverer: Sturtevant, 1930. new order: 1 — 16A1 |20-Ys;
references: Dobzhansky, 1932, Biol. Zentr. 52: 495. 61 - 66B13|(16A1 - 19)|66C1 - 100.
genetics: Breakpoint in X chromosome between bb Tentative.
and centromere; break in 3L between TU and se. origin: Neutron induced in X-Ys, sc w B.
Crossing over in 3L greatly reduced. Male and discoverer: Norby.
homozygous female fertile. Male and female hyper- references: Muller and Norby, 1949, DIS 23: 61.
ploid for the 3LDXP element survive and are fer- genetics: Associated with BS3i. Male viable. Homo-
tile; duplicated for locus of ru but not se, ft, car, zygous female lethal. Chromosome 3 containing in-
or bb. serted X material survives as duplication in pres-
*T(l;3)107 ence of normal X chromosomes; male sterile;
discoverer: Sturtevant, 1930. female fertile. Duplication has extreme B pheno-
genetics: Probably a segment from chromosome 3 is type.
.intercalated into X chromosome since segregants *T(l;3)Bb: Translocation(l;3) Bubble
are a Minute -bearing 3, presumably Df(3)107, and a cytology: T(1;3)13E;84F (Morgan, Bridges, and
Minute-suppressing X, presumably Dp(3;l)107. Schultz, 1937, Carnegie Inst. Wash. Year Book 36:
Male viable but homozygous female lethal. 301).
*T(1;3)U3-3 origin: X ray induced.
origin: X ray induced. discoverer: R. L. King, 32d.
discoverer: Neuhaus. genetics: Associated with Bb. Male sterile.
synonym: T(l;3)DeU43-3. T(1;3)C48: Translocation(l;3) Crossover
references: 1941, DIS 15: 16. suppressor
genetics: Two breaks in X chromosome, one be- cytology: In(l)10E-F;l8C-D; additional presence of
tween sc and br and another near the centromere. T(l;3)20;80-8l inferred from genetic data.
A break in chromosome 3 is between st and the origin: X ray induced.
centromere of 3. The y+ and s c + loci are then discoverer: Roberts, 1964.
attached to proximal end of 3L, and the distal end genetics: Male lethal. Recombination reduced in X
of 3L is attached to centromere of X. Bulk of the chromosome.
X chromosome is acentric and lost. T(1;3)C151
*T(1;3)26Q-21 cytology: T(l;3)9D;80-81; position of breakpoint in
cytology: T(l;3)6C;70E-F. chromosome 3 with respect to centromere not deter-
origin: X ray induced simultaneously with mined.
discoverer: Sutton, 1939. discoverer: Roberts, 1965.
references: 1943, Genetics 28: 210—17. genetics: Male viable but sterile. Recombination
reduced in X chromosome.
*T(l;3)Ah Translocathn(h3} from Austin
origin: X ray induced, cytology: T(l;3)20;90E + Df(3R)88B-C;94A.
discoverer Muller, 1926. new order 1 - 20|(90E - 88C|94A - 90E)|20;
references: Painter and Muller, 1929, J. Heredity 61 - 8 8 B | 9 4 A - 100.
20: 287-98. origin: X ray induced,
genetics: Breakpoints in X chromosome between dm discoverer. Roberts, 1965.
and ec and between car and bb. Midsection of X genetics: Male fertile. Recombination reduced in
translocated to 3R. Fly hyperploid for Dp(l;i)Al 3R.
survives; duplicated for loci of y through dm as
well as bb (Schultz). cytology. T(l;3)14B;80-81; position of breakpoint in
see T chromosome 3 with respect to centromere not deter-
mined.
T(1;3)BS8I; Transtocation(l;3 Bar origin: X ray induced.
cytology: T(1;3)16A;88F. discoverer Roberts, 1965.
origin: X ray induced. genetics: Male lethal. Recombination reduced in X
discoverer: E. B. Lewis, 5814. chromosome.
T(1;3)C195 new order: [lA4 - 7B2|7D6 - 20-20F - 20A11;

cytology: T(1;3)1W;71A-B. 61 - 85|(7B3 - 7D2)|85 - 100.
origin: X ray induced. origin: X ray induced in R(l)2.
discoverer: Roberts, 1965. discoverer: Hannah, 1947.
genetics: Male viable but sterile. Recombination genetics: Mutant for ct but not cm or sn. Male
reduced in X chromosome. lethal. The derived Dp(l;3)ct*2cl = Dp(l;3)7B2-
T(l;3)C250 3;7D2-6;85 survives as female and as sterile male;
cytology: In(l)9F;15D-E; additional presence of duplicated for sn.
T(l;3)20;80-81; inferred from genetic data. *T(l;3)ct 268-5
origin: X ray induced., cytol ogy: T(l;3)7B2-3;90C4-Dl.
genetics: Male viable and fertile; homozygous fe- discoverer: Demerec, 33k.
male viable. Recombination reduced in X chromo- genetics: Mutant for ct but not scp or en.
some. *T(1;3)ct268.2i
T(1;3)C291 cytology: T(1;3)7B3-4;7B4-5;96F.
cytology: T(l;3)16C;20;87F;98E. new order: 1A - 7B3|96F - 61;
new order: 1 _ 16C|98E - 87F|(16C - 20)|87F - 61; 20 - 7B5|96F - 100;
20|98B - 100. deficient for 7B4.
discoverer: Roberts, 1965. discoverer: Hoover, 35i.
genetics: Male viable and fertile; homozygous fe- genetics: Mutant for ct but not scp or sn. Male
male viable. Recombination reduced in 3R. lethal.
T(l;3)C300 *T(l;3)ct268-n
cytology: T(1;3)12C;61F;66E;68D. cytol ogy: T(1;3)3D2-3;7B2-5;84D4-5;86B4-C1;88F
new order: 1 — 12C|68D — 100; (Hoover),
61 - 61F|66E - 61F|68D - 66E|l2C - new order: 61 - 84D4|(3D3 - 7B2)|88F - 100;
20. remainder not described,
discoverer: Roberts, 1965. discoverer: Demerec, 38d.
genetics: Male dies in third larval instar. Recombi- genetics: Mutant for ct and dm but not scp, sn, or fa.
nation reduced in 3L. Male lethal.
T(1;3)C315 *T(l;3)ct268-36
cytology: T(l;3)20;70F. cytology: T(1;3)7B2-C1;66F (Sutton).
discoverer: Roberts, 1965. discoverer: Demerec, 39j.
genetics: Male fertile; homozygous female lethal. genetics: Mutant for ct. Male lethal.
Recombination reduced in 3L. *T(1;3)ct268-37
T(1;3)C329 cytology: T(l;3)5D2-3;7B2-3;80C-F.
cytology: T(l;3)3F;80-81; position of breakpoint in new order: 1 - 5D2|7B3 - 20;
chromosome 3 with respect to centromere not deter- 61 - 80C|7B2 - 5D3|80F - 100.
mined. origin: X ray induced,
discoverer: Roberts, 1965. references: Sutton, 1940, Genetics 25: 534—40
genetics: Male viable but sterile. Recombination (fig-).
reduced in X chromosome. Demerec, 1940, Genetics 25: 618—27.
genetics: Mutant for ct; variegated for tux and vs;
T(7;3Jcf n °: Translocation(1;3) cut shf, cm, and sn not affected. The segregant
cytology: T(1;3)IB;7B2-3;8E-F;84B superimposed on Dp(l;3)ct268-37 = Dp(i;3)5D2-3;7B2-3;80C-F
R(1)1A3-4;19F-2OA1. viable and fertile in both male and female. Its
new order. |lA4 - 1 B | 8 E - 7B3|8F - 2O-2OF - complement, Dt(l)ct268-37 ^Df(l)5D2-3;7B2-3,
20Al|; survives as a Minute female; deficient for M(l)30,
61 - 84B|(1B - 7B2)|84B - 100. TUX, vs, shf, and cm but not r&, ex, cv, or an-
origin: X ray induced in R(l)2. nuitant for ct.
discoverer Hannah, 1947.
genetics: Mutant for ct but not y, me, sc, cm, mi, or *T(l;3)cul°0.69: Trafislocation(l;3) cwr/ec/
oc. Male lethal. Female carrying Dp(l;3)ctllm = cytology: T(l;3)6Bl-Ct;88A4-Bl.
Dp(l;3)lB;7B2-3;84B survives and has small eyes origin: X ray induced,
and arclike wings with deltalike venation; dupli- d i scoverer: Alexander.
cated for cm. references: Ward and Alexander, 1957, Genetics 42:
42-54.
cytology: T(l;3)7B2-3;7D2-6;85 superimposed on genetics: Mutant for cu.
R( 1 )1A 3-4;19F-2QA 1. T(l,3)Del 143-3: see T(l;3)143-3
T(1;3)D3 references: 1960, Genetics 45: 1649—70.

cytology: T(1;3)4F;62A. genetics: Variegated fora lethal. Male sterile.
origin: Induced by tritiated deoxycytidine. T(l;3)l-vl63
discoverer: Kaplan, 1965. cytology: T(l;3)17A-B;80-81; position of chromo-
references: 1966, DIS 41: 59. some 3 breakpoint with respect to centromere not
genetics: Male lethal. determined.
*T(1;3)f257-29: Translocation(l;3) forked origin: X ray induced.
cytology: T(1;3)15F5-16A1;64. discoverer: Lindsley, Edington, and Von Halle.
origin: X ray induced, references: 1960, Genetics 45: 1649—70.
discoverer: Bishop, 401. genetics: Variegated for a lethal. Male sterile.
genetics: Mutant for / but not M(l)o, B, or os. Male *T(l;3)l-v252
viable but sterile. origin: X ray induced.
*T(l;3)fd: Translocation(l;3) furled discoverer: Lindsley, Edington, and Von Halle.
cytology: T(1;3)7A;86E superimposed on references: 1960, Genetics 45: 1649—70.
In(3R)89C;96A (Darby), genetics: Variegated for a lethal. Male sterile.
new order: 1 - 7A|86E - 61; T(l;3)l-v361
20 - 7A|86E - 89C|96A - 89C|96A - cytology: T(l;3)19-20;80-81; position of breakpoints
100. with respect to centromeres not determined.
origin: Induced by P3 2 in In(3R)P. origin: X ray induced.
discoverer: Bateman, 1949. discoverer: Lindsley, Edington, and Von Halle.
references: 1950, DIS 24: 54. references: 1960, Genetics 45: 1649—70.
1951, DIS 25: 77. genetics: Variegated for a lethal. Male sterile.
genetics: Associated with Id, T(l;3)t-v453
*T(1;3)H: Translocation(l;3) Hairless cytology: T(l;3)12D;80-81; position of breakpoint in
discoverer Efroimson. chromosome 3 with respect to centromere not deter-
references: Kamshilov, 1933, Biol. Zh. (Moscow) 2: mined.
161-83. origin: X ray induced.
genetics; Break in X chromosome to the left of w; discoverer: Lindsley, Edington, and Von Halle.
3R broken near H. references: 1960, Genetics 45: 1649—70.
genetics: Variegated for a lethal. Male sterile.
T(1;3)K2: Translocation(l;3) of Krivshenko
cytology: T(l;3)20A-B;20D-F;80-8l superimposed on cytology: T(1;3)3C;81.
ln(l)lB2-3;20B-Dl. Inferred from genetic data origin: X ray induced.
since salivary chromosomes do not reveal an aber- discoverer: Lindsley, Edington, and Von Halle.
ration. In ganglia 1 metaphase, chromosome J is a references: 1960, Genetics 45: 1649—70.
rod-shaped and a J-shaped element. genetics: Variegated for w and a lethal. Male
new order: 1A - 1B2|2OBJ8O - 100; sterile.
20F|80 - 6 1 . T(1;3)l-v463
Tentative. cytology: T(l;3)19-20;81-82; position of breakpoints
origin: X ray induced in In(l)sc8. with respect to centromeres not determined,
discoverer: Krivshenko, 55g3. origin: X ray induced.
references: 1956, DIS 30: 76. discoverer: Lindsley, Edington, and Von Halle,
genetics: Irradiated In(l)sc8 broken in distal region references: 1960, Genetics 45: 1649—70.
between y + and 6fe+ and also near the centromere. genetics: Variegated for a lethal. Male sterile.
Chromosome 3 broken near the centromere, whether *T(1;3)lz268.29: Translocation(l;3) lozenge
to left or right of the centromere is not known. Tip cytology: T(1;3)8D8-9;81F.
of X chromosome with y + and ac + is attached to the origin: X ray induced.
chromosome 3 centromere and one arm of this chro- discoverer: Hoover, 38d.
mosome is attached to the X centromere. Bulk of genetics: Mutant for Iz and independently for ct but
the X chromosome is thus acentric and lost. Homo- not sn, t, dvr, tip, or ras. T(l;3)lz268-29/lz f e m a le
zygote viable and moderately fertile. This chromo- fertile. Male lethal.
some may be considered as Dp(l;3)K2. *T(1;3)N34b: Translocation(h3) Notch
*T(1;3H-184: Translocation(h3) lethal origin: X ray Induced.
cytology: T(1;3)18A;81. discoverer Oliver, 34b3.
discoverer: Lindsley, Edington, and Von Halle. genetics: Carries mutant allele of TV and normal
references: 1960, Genetics 45: 1649-70. alleles of w and ec.
gtMrici: Associated with 1(1)184. other information: Reported as suspected of being a
T(l;3)I-v3: Translocatiort(l;3) lethal-variegated T(l;3); basis of suspicion not given.
cytology: T(1;3)4A;81. T(l;3)NS0kii
origin: X ray induced. cytology: T(1;3)1E3-4;3C6-7;3C8-9;89A; 3C7-8
discoverer: Lixklsley, Edington, and Von Halle. missing (Lefevre).
new order: 1A1 - 1E3|3C9 - 20; genetics: Variegates for w, rst, N, and its pseudo-
61 - 89A|(1E4 - 3C6)|89A - 100. alleles (Cohen, 1962, Genetics 47: 647—59); seems
origin: X ray induced, to carry a mutant allele of dm. The segregant,
discoverer: Lefevre, 50kll. Df(iyM264-58 =Df(l)3B2-3;3D6-7 survives in>heter-
references: 1951, DIS 25: 71. ozygous female and is deficient for w, rst, fa, and
1952, DIS 26: 66. dm but not pn or ec. Dp(l;3)N264-S8 = Dp(l;3)3B2-
Ratty, 1954, Genetics 39: 513-28. 3;3D6-7;80D-F survives as both male and female.
genetics: Mutant for N. The segregant *T(1;3)N 264-64
Dp(l;3)N50k11 =Dp(l;3)lE3-4;3C6-7;89A is viable cytology: T(l;3)3E5-6;80C-F (Hoover).
and carries normal alleles of pn, w, and rst. origin: X ray induced.
T(J;3)N264.6 discoverer: Demerec, 38e.
cytology: T(l;3)3C9-Dl;62A;73E;80C (Schultz). references: 1941, Proc. Intern. Congr. Genet., 7th.
new order: 1 - 3C9|80C - 73E|3D1 - 20; pp. 99-103.
61 - 62A|73E - 62A|80C - 100. genetics: Variegates for w, rst, fa, and dm but not
origin: X ray induced. pn, kz, or ec.
discoverer: Demerec, 33k20. *T(1;3)N 264-65
genetics: Variegates for w and N; position effect on cytology: T(l;3)2B10-16;3D4-5;81F;96C4-5 (Hoover).
pb and Dfd. X/Y male lethal; X/Y/Y viable and new order: 1 - 2B10J(81F - 96C4)|3D5 20;
sterile. Dp(3;ipi264'6 - Dp(3;l)3C9-Dl;73E;80C 61 - 81F|(2B16 - 3D4)|96C5 100.
viable. DitfLyX2*4-6 = Df(3L)73E;80C survives origin: X ray induced.
and is Minute, possibly deficient for M(3)S34. discoverer: Demerec, 38e.
*T(l;3)N264-29 genetics: Variegates for w, rst, fa, and dm but not
cytology: T(l;3)3D4-5;80 (Hoover). kz.
origin: X ray induced. *T(T;3)N264-70
discoverer: Demerec, 36d. cytology: T(l;3)3C4-5;80D-F + T(1;3)6F2-
references: 1941, Proc. Intern. Congr. Genet., 7th. 7Al;100B2-3 (Sutton).
pp. 99-103. new order: 1 - 3C4|80F - 100B2|6F2 - 3C5|80D -
genetics: Variegates for rst, fa, and dm but not w or 61;
ec. X/Y male lethal; X/Y/Y occasionally sur- 20 - 7Al|lO0B3 - 100F.
vives. origin: X ray induced.
*T(l;3)N264-34 discoverer: Demerec, 38k.
cytology: T(l;3)3C3-5;70C2-3 (Hoover). references: Sutton, 1940, Genetics 25: 534—40.
origin: X ray induced. genetics: Variegates for w, rst, fa, and dm but not
discoverer: Demerec, 37a, kz, pn, ec, cm, scp, or shf.
references: 1941, Proc. Intern. Congr. Genet., 7th. *T(l;3)N264-83
pp. 99-103. cytology: T(l;3)3C6-7;12F2-4;79E2-3 +In(3R)81;88
genetics: Contains mutant allele of TV but normal (Sutton).
alleles of w, rst, and dm. new order: 1 - 3C6|l2F2 - 3C7|79E3 - 81 [SB -
8l]88 - 100;
*TO;3)N264.ss 20 - 12F3|79E2 - 6l.
cytology: T(l;3)3D4-5;80F9-81Fl; chromosome 3 origin: X ray induced.
claimed broken in 3R. discoverer: Demerec, 39d.
origin: X ray induced, references: 1941, Proc. Intern. Congr. Genet., 7th.
discoverer: Demerec, 38b. pp. 99-103.
references: 1941, Proc. Intern. Congr. Genet., 7th. genetics: Carries mutant allele of N but normal
pp. 99-103. alleles of w, rst, and dm.
genetics: Variegates for w, rst, fa, and dm but not
pn, kz, or ec. cytology: T(l;3)3B4-Cl;4B4-S;80 [Suttoo, 1940,
*T(1;3)N264-S6 Genetics 25: 534—40 (fig.); Gersh, 1959, Genetics
cytology: T(l;3)3D4-5;80 (Sutton). 44: 163-72].
origin: X ray induced. new order: 1 - 3B4|4B5 - 20;
discoverer: Demerec, 38c. 61 - 80|4B4 - 3C1J80 - 100.
genetics: Variegates for w and probably N. origin: X ray induced.
cytology: T(l;3)3B2-3;3D6-7;80D-F (Sutton). references: 1940, Genetics 25: 618—27.
new order. 1 - 3B2|3D7 - 20; genetics: Variegates for w, rat, fa, dm, and me bat
61 - SCOLDS - 3B3J80F - 100. not pn or hi. The segregant, D§(1)N3*4"1O@ «
origin: X ray induced, Df(l)3B4'Ct;4B4-S, deficient for IV &nd M{1)3E,
discoverer: Demerec, 3&d. survives In heterozygous female. Dp(l;3)N264-10Q
synonym: TXltfysv**364-**. Dp(l;3)3B4-Cl;4B4~5;80 originally survived in
references: 1940, Genetics 25: 618—27. female but not mate; more recently, male carrying
Sutton, 1940, Genetics 25: 534-40 (fig.). duplication foond to awvive (Gersh, 1959).
*T(J;3)N 264-104 discoverer: Warters, 1961.

cytology. T(1;3)3C7-9;87D1-E1 + In(l)lB4-5;18-19 genetics: Male lethal. Male hyperploid for 3LDXP
(Sutton). element survives.
new order: 1A1 - lB4|l8 - 3C9|87D1 - 61; T(1;3)OR6
20 - 19|lB5 - 3C7J87E1 - 100. cytology: T(1;3)4D;87F.
discoverer: Demerec, 39j. discoverer: Warters, 1961.
genetics: Mutant for svr and N but not ac, sc, sta, genetics: Male viable but sterile.
tw, w, rst, p, ss, k, or e. T(1;3)OR7
*T(l;3)N264.m cytology: T(1;3)14D;91E.
cytology: T(1;3)3C7-9;81F;86B6-C1 (Sutton). origin: X ray induced.
new order: 1 - 3C7|81F - 86B6|81F - 61; discoverer: Warters, 1961.
2O-3C9|86C1 - 100. genetics: Male viable but sterile.
origin: X ray induced. W;3)OR9
discoverer: Demerec, 40j. cytology: T(1;3)6D;66B.
genetics: Carries mutant allele of N and normal origin: X ray induced.
alleles of kz, w, and dm. discoverer: Warters, 1961.
T(1;3)O4: Trons/ocaf/on(7;3) of Oliver genetics: Variegated for a lethal; male sterile. Male
origin: X ray induced. has small rough eyes; perhaps mutant for rux.
discoverer: Oliver, 29k24. T(l;3)0 R11
references: Dobzhansky and Schultz, 1934, J. Genet. cytology: T(1;3)18F;84B.
Oliver, 1937, DIS 7: 19. discoverer: Warters, 1961.
genetics: X chromosome broken between m and g genetics: Male viable but sterile.
and between f and car. Center section of X then T(h3)OR12
inserted into 3L. The segregant Df(l)O4 is in- cytology: T(1;3)2B6-13;84A + T(1;3)18D;98F-99A
viable when added to a normal male genotype, (Becker),
poorly viable when added to a normal female geno- new order: 1 - 2B6|84A - 98F|l8D - 2B13|84A _
type, and survives well when added to an intersex 61;
(2X:3A) genotype, where it confers a low degree of 20 - 18D|99A - 100.
fertility. The reciprocal segregant, Dp(l;3yO4, is Tentative.
lethal in the male, survives well in the female, and origin: X ray induced in y.
poorly in intersexes. discoverer: Warters, 1961.
T(l;3}05 genetics: Male viable but sterile. Male hyperploid
cytology: T(1;3)4F2-3;62B-C;88A-C;92C-D (Lewis, for 3RPXD element survives.
1951, DIS 25: 108-9). T(1;3)OR13
new order, 1 - 4F2|88C - 92C|4F3 - 20; cytology: T(1;3)15A;84E.
61 - 62B|88A - 62BJ92D - 100. origin: X ray induced.
discoverer: Oliver, 29130. genetics: Male viable but sterile.
references: 1937, Am. Naturalist 71: 560—66. T(l;3)0RU
1938, Genetics 23: 162. cytology: T(l;3)17A;80B.
genetics: Male viable and fertile. Homozygous fe- origin: X ray induced.
male viable but sterile. The segregant, Dp(3;l)O5 = discoverer: Warters, 1961.
Dp(3;l)4F2-3;88A-C;92C-D is viable and fertile in genetics: Variegated for a lethal; male sterile.
male and female. It is duplicated for loci of red, T(1;3)OR1S
jvl, cv-c, so(Hw), sbd, ss, bar, at, gl, k, and Dl but cytology: T(1;3)18D;88A.
not cu, ry, kar, or e (Lindsley and Grell, 1958, DIS origin: X ray induced,
32: 136; E. B. Lewis). Produces roughish eyes, discoverer: Warters, 1961.
spread, nicked wings, coarse bristles, and a darkly genetics: Male viable but sterile. Subsequently ac-
pigmented abdomen. quired a male lethal.
T(1;3)O6 T(1;3}OR17
origin: X ray induced. cytology; T(t;3)19E;67C.
discoverer: Oliver, 34d24. origin: X ray induced,
genetics: Mutant for ec. Break in 3L between nt and discoverer: Warters, 1961.
h. Break in X not determined. Male and homozy- genetics: Male viable and fertile. Homozygous fe-
gous female viable and fertile. male viable. Male hyperploid for 3LDXP survives.
T(1;3)OR1: Translocation(l;3) from Oak Ridge
cytology: T(l;3)5A;20;66B;79E. cytology: T(1;3)19B;8OA.
new order: 1 - 5Ai79E - 66B]SA - 20J79E - 100; origin: X ray induced,
20}66B - 6 1 . discoverer: Warters, 1961.

cytology: T(1;3)12E;75F. discoverer: Warters, 1961.
origin: X ray induced. genetics: Male virtually lethal.
genetics: Male viable but sterile. cytology: T(1;3)3A;65A.
T(J;3)OR2J origin: X ray induced in y,
cytology: T(1;3)19E;61F. discoverer: Warters, 1961.
origin: X ray induced. genetics: Male viable but sterile, with either a
discoverer: Warters, 1961. normal Y or Bsw+Y.
genetics: Male viable but sterile; X/0 male dies. T(1;3)OR35
Male hyperploid for 3LDXP element survives. cytology: T(1;3)19E;75C.
T(1;3)OR22 origin: X ray induced,
cytology: T(1;3)6C;98C. discoverer Warters, 1961.
genetics: Male viable and fertile. Homozygous fe- cytology: T(1;3)7D;62A;87E.
male viable. new order 1 - 7D|62A - 87E|62A - 61;
T(1;3)OR23 20 - 7D|87E - 100,
cytology: T(l;3)20;80-81; positions of breakpoints origin: X ray induced,
with respect to centromeres not determined, discoverer: Warters, 1961.
discoverer Warters, 1961. T(1;3)OR37
genetics: Male viable but sterile. cytology: T(1;3)3C;97F.
T(];3)OR24 origin: X ray induced in y.
cytology: T(l;3)12F;80B. discoverer Warters, 1961.
origin: X ray induced, genetics: Male viable and fertile; homozygous fe-
discoverer: Warters, 1961. male viable.
genetics: Male viable but sterile. T(h3)OR38
T(1;3)OR25 cytology: T(1;3)18D;61D.
cytology: T(l;3)20B;99B. origin: X ray induced.
discoverer: Warters, 1961. genetics: Male viable but sterile. Male hyperploid
genetics: Male viable but sterile. Male hyperploid for 3LDXP survives.
for 3RDXP element survives.
T(1;3)OR28 T(1;3)OR39
cytology: T(1;3)11A;8OC cytology: T(1;3)6B'F;75C.
genetics: Male lethal. genetics: Male viable but sterile.
T(1;3)0R29 T(1;3)OR40
cytology: T(1;3)16F;84B. cytology: T(1;3)6F;62D + r(l;3)16B;20;84F.
origin: X ray induced, new order 1 - 6F|62D - 84F|20 - 16BJ84F - 100;
discoverer Warters, 1961. 20|l6B - 6FJ62D - 61.
genetics: Male viable but sterile. origin: X ray induced,
T(T;3)OR3O discoverer Warters, 1961.
cytology: T(1;3)19E;65D. genetics: Male viable but sterile.
discoverer: Warters, 1961. cytology: T(1;3)9F;98E.
genetics: Male viable but sterile. Male hyperploid origin: X ray induced,
for 3LDXP element survives. discoverer Warters, 1961.
T(1;3)OR31 genetics: Male viable but sterile.
cytology: T(l;3)10A;68D. T(1;3)OR43
origin: X ray induced, cytology: T(l;3)20A;97D.
discoverer: Warters, 1961. origin: X ray induced.
genetics: Male viable but sterile. discoverer Warters, 1961.
T(1;3)OR32 genetics: Male viable but sterile. Male hyperploid
cytology. T(1;3)16A;71B. for 3RDXP element survives.
discoverer: Waiters, 1961. cytology: T(1;3)17A;61D.
genetics: Male viable but sterile. origin: X ray induced.
T(1;3)OR33 discoverer Warters, 1961.
cytology* 7X1;3)13E;62F. genetics: Male viable but sterile.

cytology: T(l;3)12C;80A. discoverer Warters, 1961.
discoverer: Waiters, 1961. T(J;3)OR63
genetics: Variegated for a lethal; male sterile. cytology: T(l;3)20;80-81; positions of breakpoints
T(1$)OR47 with respect to centromeres not determined.
cytology: T(l;3)20;93D. origin: X ray induced.
discoverer. Waiters, 1961. genetics: Male sterile.
genetics: Male viable and fertile; homozygous fe- T(1;3)OR66
male viable. Male hyperploid for 3RDXP element cytology: T(1;3)3F;71E,
survives. origin: X ray induced in y.
T(1;3)OR49 discoverer Warters, 1961.
cytology: T(1;3)11A;66D. genetics: Male lethal; lethality not covered by
origin: X ray induced, Bsw+Y.
discoverer Warters, 1961. T(1;3)OR67
genetics: Male viable but sterile. cytology: T(1;3)4C;73C.
cytology: T(1;3)12D;97A. discoverer: Warters, 1961.
discoverer. Warters, 1961. T(1;3)OR69
genetics: Male lethal. cytology: T(l;3)20;80-81; positions of breakpoints
T(J;3)OR52 with respect to centromeres not determined.
cytology: T(l;3)19E;70C;83F. origin: X ray induced.
new order. 1 - 19EJ83F - 7OCJ83F - 100; discoverer: Warters, 1961.
20 - 19E|70C - 61. genetics: Male viable but sterile.
Tentative. T(1;3)OR71
origin: X ray induced. cytology: T(l;3)20;71D.
discoverer: Warters, 1961. origin: X ray induced,
genetics: Male viable but sterile. discoverer: Warters, 1961.
T(U3)ORS4 genetics: Male viable but sterile.
cytology: T(1;3)12F;83A. T(1;3)OR72
origin: X ray induced. cytology: T(l;3)20;80-8l; positions of breakpoints
discoverer: Warters, 1961. with respect to centromeres not determined.
*T(1;3)OR55 discoverer: Warters, 1961.
cytology: T(1;3)11C;67C. genetics: Variegated for a lethal. Male sterile.
discoverer: Waiters, 1961. T(l;3)osb<i": Translocation(l;3) outstretched
genetics: Male viable but sterile. small eye-bending wings
*T(l;3)ORS7 cytology: T(l;3)16E;80C (Nicoletti).
cytology: T(1;3)3E;5B;61C. origin: X ray induced.
new order: 1 - 3E|5B - 3E|61C - 100; discoverer: Halfer, 1960.
20 - 5BJ61C - 61A. genetics: Associated with osbdw. Male sterile.
origin: X ray induced in y. T(J;3)P104: Translocation(l;3) from Pasadena
discoverer. Warters, 1961. cytology: T(l;3)l9-20;87F-88A.
genetics: Male lethal. Lethality not covered by origin: X ray induced.
Bsw+Y; therefore probably associated with break discoverer. E. B. Lewis.
in SB. 7(l;3)ras^: TranslocationfJ;3) raspberry-
T0',3)0RS9 variegated
cytology: T(l;3)20;80-81; positions of breakpoints cytology: T(1;3)9E;13C;81F (E. B. Lewis).
with respect to centromeres not determined. new order. 1 - 9EJ13C - 20;
origin: X ray induced, 61 - 81F|(9E - 13C)|81F - 100.
discoverer: Warters, 1961. origin: Fast neutron induced.
genetics: Male lethal. discoverer: E. B. Lewis, 1953.
T(1;3)OR60 references: Brokaw, 1954, DIS 28: 73.
cytology: T(1;3)4B;88A. genetics: Shows recessive variegation for ras and a
origin: X ray induced. rough eye and dominant variegation for a wing
discoverer Warters, 1961. effect resembling Bg/+. No variegation for m, vb,
genetics: Male lethal. sot, si, or un. Is probably an enhancer of J3; a few
T(1;3)OR62 msv/rmmv female® somewhat resemble £?/+.
cytology: 7X1;3)1OF;88C. Dp(l;3)r®$sv » Dp(l;3)9E;13C;81F male dies but fe-
male survives; duplicated for ras. v, m, dy, and g ac but is not demonstrably deficient for 3L factors
but not un or r. Df(l)rasv is lethal in both sexes. since it is homozygous viable. y+ localizes about
*T(l;3)rst: Translocation(J;3) roughest four units to the left of ru.
discoverer Ball, 32b25. *T(l;3)scK: TranslocationO;3) scute of
genetics: Associated with rst. Breakpoints in X Krivshenko
chromosome near w and bb; position of breakpoint discoverer: Krivshenko.
in chromosome 3 unknown. references: Agol, 1936, DIS 5: 7.
T(l;3)ry35: Translocation(l;3) rosy genetics: Mutant for sc. Three-break rearrangement
cytology: T(l;3)20;87C-E;91B-C (Lindsley). with XD translocated to 3LP; 3LD translocated to
new order: 1 - 20|(87E - 91B)|20; 3Rp, and 3RD translocated to Xp.
61 - 87C|91C - 100. T(1;3)scK3
origin: X ray induced in cu kar chromosome. cytology: T(l;3)lB2-3;61Al-2.
discoverer: Schalet. origin: X ray induced.
references: 1964, DIS 39: 62-64. discoverer: Krivshenko, 53j29.
Schalet, Kernaghan, and Chovnick, 1964, Genetics references: 1959, DIS 33: 95-96.
50: 1261-68. genetics: Mutant for sc. Male fertile. Two halves of
genetics: Deficient, mutant, or variegated for ry and the translocation recoverable separately. XD3P
pic. The segregant, Dp(3;l)ry35 = Dp(3;l)20;87C- element is viable homozygous, although males are
E;91B-C is viable and fertile in male and female; somewhat infrequent. 3DXP is inviable in male
duplicated for loci of Sb and Ubx. and homozygous female but survives in heterozy-
T(1;3)sc260-J5: TranslocationO ;3) scute gous female.
cytology: T(1;3)1B4-5;71C-D. T(l;3)sn13al: TranslocationO;3) singed
origin: X ray induced, cytology: T(l;3y6C;7C9-l0;79D2-El; chromosome 3.
discoverer: Demerec, 381. X material inserted into chromosome 3.
references: Sutton, 1943, Genetics 28: 210—17. new order: |lA4 - 6C|7C10 - 20-20F - 20Al|;
genetics: Mutant for sc but not y or ac. Male sterile. 61 - 79D2|(6C - 7C9)|79E1 - 100.
k
T(l;3)sc260-20 origin: X ray induced in R(l)2.
cytology: T(l;3)lA8-Bl;61Al-2. discoverer: Hannah, 1947.
origin: X ray induced. references: Valencia, 1966, DIS 41: 58.
discoverer: Sutton, 39e. genetics: Mutant for &n. The segregant
references: 1943, Genetics 28: 210—17. Dp(l;3)sn13al = Dp(l;3)6C;7C9-Dl survives and is
genetics: Mutant for sc but not y, ac, or svr. Male duplicated for cm and ct.
and homozygous female viable and fertile. The two
halves of the translocation are recoverable inde- cytology: T(l;3)3Cl-2; 7C9-10;72A-B superimposed
pendently. The 3LDXP element should be deficient on In(l)lB3-4;2QB-DlI*iB2-3;20B-DlR +Ia(l)4D7-
for y and ac but carry sc 2 * 0 " 2 0 ; it presumably is El;llF2-4.
male lethal but survives in heterozygous female. new order: 1A - 1B3|2OB - 11F4|4E1 - 7C9|3C1 -
The XD3P element carries normal alleles of y and 1B3|2OD1 -2 OF;
ac but not sc260~20 or svr+; should also carry 61 - 72A|(3C2 - 4 D 7 | l l F 2 -
normal alleles of ve and ru. 7C10)|72B - 100.
T(l;3)scJ4: TranslocationO ;3) scute of Jacobs- origin: X ray induced in ln(l)scsi1Lsc8R+dl-49.
duller discoverer: Muller, Valencia, and Valencia, 1946-53.
cytology: T(1;3)1B;3A3-C2;61A (inferred from ge- references: Valencia, 1966, DIS 41: 58.
netic tests); 1B-3A3 lost. genetics: Associated with an1 *Bb5. w n o t affected.
new order. 1A1 - 1B|61A - 100;
2 0 - 3C2|61A. *TO;3}SP2: TranslocationO;3) from $db Paulo
origin: X ray induced. cytology: T(l;3)20;90E.
discoverer Jacobs-Muller. origin: Gamma ray induced in y w.
references: Muller, 1932, Proc. Intern. Congr. discoverer: Lindsley and Musatti, 1961.
Genet., 6th. Vol. 1; 225. genetics: Male viable but sterile.
1934, DIS 2: 60. T(1;3)SP11
genetics: The section of the X chromosome from IB cytology: T(l;3)20;7SB.
through 3A was presumably inserted elsewhere in origin: Gamma ray induced in y w.
the genome; it subsequently separated from the rest discoverer Lindsley and Musatti, 1961.
of the configuration and was lost. Base of the X, genetics: Male viable but sterile.
presumably capped by the indemonstrable terminus T(1;3)SP13
of 3L, i.e., 3DXp, is deficient for the tip of X cytology: T(l;3)20;80-81; positions of breakpoints
through z and may be stocked in combination with with respect to centromeres not determined,
a duplication for the tip of X, much mm the XD4P origin: Gamma ray induced.
element from T(l;4)w®»5 or Dp(l;f)x9. The XD3P discoverer: Lindsley and Musatti, 1961.
segregant carries normal alleles of 1(1 )J 1, y, and genetics: Male viable but sterile.
T(1;3)SPU origin: Gamma ray induced.

cytology: T(l;3)20;80-81; positions of breakpoints discoverer: Lindsley and Musatti, 1961.
with respect to centromeres not determined. genetics: Male viable but sterile.
origin: Gamma ray induced. T(1;3)SP54
discoverer: Lindsley and Musatti, 1961. cytology: T(l;3)20;67B.
genetics: Male viable but sterile. origin: Gamma ray induced.
*T(1;3)SP15 discoverer: Lindsley and Musatti, 1961.
cytology: T(l;3)20;67. genetics: Male viable but sterile. Male hyperploid
origin: Gamma ray induced in y w. for 3LDXP element survives.
genetics: Male viable but sterile. Male hyperploid cytology: T(l;3)20;83C.
for 3LDXP element survives. origin: Gamma ray induced.
T(1;3)SP21 discoverer: Lindsley and Musatti, 1961.
cytology: T(1;3)1B;83F. genetics: Male viable but sterile.
origin: Gamma ray induced in y w. T(1;3)SP62
discoverer: Lindsley and Musatti, 1961. cytology: T(l;3)20;89A.
genetics: Male viable but sterile. origin: Gamma ray induced.
T(1;3}SP22 discoverer: Lindsley and Musatti, 1961.
cytology: T(l;3)20;80-81; positions of breakpoints genetics: Male viable but sterile.
with respect to centromeres not determined. T(1;3)SP63
origin: Gamma ray induced. cytology: T(l;3)20;65.
discoverer: Lindsley and Mussati, 1961. origin: Gamma ray induced.
T(1;3)SP26 genetics: Male viable but sterile. Male hyperploid
cytology: T(l;3)20;80-81; positions of breakpoints for the 3LDXP element survives.
with respect to centromeres not determined. T(1;3)SP68
origin: Gamma ray induced. cytology: T(1;3)11A;8O-81.
cytology: T(1;3)8A;84A.
origin: Gamma ray induced. T(l;3)SP70
discoverer Lindsley and Musatti, 1961. cytology: T(l;3)20;80-81; positions of breakpoints
genetics: Male viable but sterile. with respect to centromeres not determined.
cytology: T(1;3)8F;64E. discoverer: Lindsley and Musatti, 1961.
origin: Gamma ray induced. genetics: Male viable but sterile.
discoverer: Lindsley and Musatti, 1961. *T(J;3)SP73
genetics; Male viable but sterile. cytology: T(l;3)20;89E.
T(1;3)SP38 origin: Gamma ray induced in y w.
cytology: T(l;3)10;84. discoverer: Lindsley and Musatti, 1961.
origin: Gamma ray induced. genetics: Mutant for Ubx. Male viable but sterile.
genetics: Male viable but sterile. cytology: T(1;3)13D;64A.
T(1;3)SP41 origin: Gamma ray induced,
cytology: T(1;3)3E;67C-D. discoverer: Lindsley and Musatti, 1961.
origin: Gamma ray induced in y w. genetics: Male viable but sterile.
discoverer: Lindsley and Musatti, 1961. *T(h3)SP90
genetics: Male viable but sterile. cytol ogy: T(l;3)18C;l 00A.
T(1;3)SP44 origin: Gamma ray induced in y w.
cytology: T(l;3)20;80-81; positions of breakpoints discoverer: Lindsley and Musatti, 1961.
with respect to centromeres not determined, genetics: Male viable but sterile. Male hyperploid
origin: Gamma ray induced. for 3RDXP element survives.
genetics: Male viable but sterile. cytology: T(1;3)5B-C;81.
cytology: T(l;3)ll;98. discoverer: Lindsley and Musatti, 1961.
origin: Gamma ray induced. genetics: Male viable but sterile.
discoverer: Lindsley and Musatti, 1961. T(1;3)SP8S
genetics: Male viable but sterile. cytology: T(l;3)16B;80-81; position of chromosome
T0;3)SPS3 3 breakpoint with respect to centromere not deter-
cytology: T(l;3)12;92. mined.
CHROMOSOME ABERRATIONS- TRANSLOCATIONS 379
origin: Gamma ray induced. 1926, Papers Mich. Acad. Sci. 7: 273-78.
discoverer: Lindsley and Musatti, 1961. 1929, Z. Induktive Abstammungs- Vererbungslehre
genetics: Male viable but sterile. 51: 397-411.
T(l;3)SP90 genetics: Inseparable from v. Male viable but
cytology: T(1;3)18D;68A. sterile. Primary nondisjunction occurs with a fre-
origin: Gamma ray induced. quency of about 2 percent in heterozygous females,
discoverer: Lindsley and Musatti, 1961. secondary nondisjunction is 23 percent. Crossing
genetics: Male viable but sterile. over is reduced near v but approaches normal on
T(1;3)SP99 both ends of the X.
cytology: T(1;3)12E;64E. *T(1;3)Vel: Translocation(J;3) Velvet
origin: Gamma ray induced. origin: X ray induced in In(l)sc8.
discoverer: Lindsley and Musatti, 1961. discoverer: Patterson,
genetics: Male viable but sterile. references: 1934, DIS 2: 10.
T(1;3)SP112 genetics: Associated with Vel. Homozygous viable
cytology: T(1;3)11B;8SD. and fertile.
origin: Gamma ray induced. T(l;3)w+S4a4: s e e T(l;3)54a
discoverer: Lindsley and Musatti, 1961. T(l;3)w+S4cxo: S ee T(l;3)54c
T(J;3)SP122 *T(l;3)w»h Translocation(l;3) white-mottled
cytology: T(1;3)11E;92E. origin: X ray induced,
origin: Gamma ray induced. discoverer: Muller, 1927.
discoverer: Lindsley and Musatti, 1961. references: 1930, J. Genet. 22: 299-334.
genetics: Male viable but sterile. genetics: Variegated for w and N. X/Y lethal,
T(1;3)ssv: Translocation(l;3) spineless- X/Y/Y viable and sterile,
variegated other information: First recorded case of variegated
cytology: T(1;3)20;89B;100F; breakpoint in X chro- position effect.
mosome inferred from genetic results; not visible *T(l;3)w>»2
cytologically. origin: X ray induced.
new order: 1 - 20|(89B - 100F)|20; discoverer: Patterson.
61 - 89BJ100F. references: Muller, 1930, J. Genet. 22: 299—334.
Tentative. genetics: Variegated for w. Male sterile.
discoverer: E. B. Lewis. cytology: T(l;3)3A10-Bl;3E2-3;80.
genetics: Variegated for ss and mutant for ar is ta- new order: 1 - 3A1QJ3E3 - 20;
pe dia. Male viable but sterile. 61 - 80|(3Bl - 3E2)|80 - 100.
T(l;3)sta: Translocation(l;3) stubarfsta origin: X ray induced.
cytology: T(1;3)1D3-E1;2A;89B21-C4 (E. B. Lewis), discoverer: Lefevre, 49a7.
new order: 1A - 1D3|2A - 20; synonym: T(l;3)wmSP: Translocation(l;3) white-
61 - 89B2l|(lEl - 2A)J89C4 - 100. mottled Spotted,
discoverer: Oliver, 32122. 1951, DIS 25: 71.
references: 1935, DIS 4: 15. Ratty, 1954, Genetics 39: 513-28.
genetics: Mutant for sta and as*. Male viable and genetics: Variegated for w, rst, and spl. The two
fertile; homozygous female lethal. The segregant elements of the trans location can be separated;
Dp(l;3)sta = Dp(l;3)lD3-El;2A;89B21-C4 is viable. Dt(l)w**4»* = Df(lpA10-Bl;3E2-3 survives in het-
The complementary Df(I)ata = Df(l)lD3-El;2A is erozygous female and is N; Dp(l;3)wm49a *=
viable in heterozygous female either as Di(l)sta/+; Dp(l;3)3AlQ-Bl;3E2-3;80 survives in both male
+/+ or Df(l)sta/Df(l)sta; Dp(l;3)sta/+ but the and female and carries the loci of w, rst, N, and
second type is sterile. (from the cytology) presumably dm.
T(l;3)Thl: Translocation(l;3) from Thy mi dine
cytology. T(1;3)12C;65B. cytology: T(1;3)3C3*5;81 (Demerec and Hoover).
origin: From male treated with H3-thymidine as origin: X ray induced.
larva. discoverer: Demerec, 371.
discoverer: Kaplan, genetics: Variegated for w but not mt, fa, or din.
genetics: Male lethal. Male viable.
T(l;3)v: Translocation(l ;3) vermilion
cytology: T(l;3)10;93B (Lewis, 1951, DIS 25: 108- cytology: T(l;3)3C3-4;4D2-El;80D; deficient for
9). 3C4-4D2.
origin: X ray induced in a chromosome carrying v. origin: Aneuploid segregant from
discoverer. E. G. Anderson, 1924.
references: 1925, Papers Mich. Acad. ScL 5: 3 5 5 -
66. cytology: T(l ;3)3B2-Cl; 19F2-20A1 ;2®E;63C7-8.
new order: 1 - 3B2|63C8 - 100; new order: 1 - 3C3|3P;

20F|l9F2 - 3C1J20A1 - 2OE|63C7 - 61. 20 - 3C7|1O1F - 102F;
origin: X ray induced. 3 D |l01F - 101A.
discoverer: Sutton, 40e. origin: X ray induced.
genetics: Variegated for w and rst, but not pn or discoverer: Mackensen.
spl. Male lethal. references: 1935, J. Heredity 26: 163-74 (fig.).
T(l;3)wm264-sa: S ee T genetics: Variegated for w.
T(l;4)2: see T(1;4)A1
cytology: T(l;3)3C2-3;100C3-4. T(l;4)t: see T(l;4)Bs
origin: X ray induced. T(1;4)3A: see T(1;3;4)A
discoverer: Patterson. T(l;4)4: see T(1;4)B*
synonym: T(l;3)Aw'r>609e,
references: Griffen and Stone, 1938, Genetics 23: *T(l;4)231b
genetics: Variegated for w. Seems likely that the discoverer: Patterson.
rearrangement is more complicated, since a euchro- references: Patterson and Painter, 1931, Science
tnatic-euchromatic translocation would not be ex- 73: 530-31.
pected to produce variegation. Patterson, 1932, Am. Naturalist 66: 193—206.
T(l;3)w™sP: see T(l;3)w^4 9a 1932, Genetics 17: 38-59.
T(l;3)wyco: Translocationfl;3) white-variegated genetics: Variegated for N and w. Left end of X
cobbled from sc to ec attached to chromosome 4.
cytology: T(l;3)2B17-Cl;3C4-5;77D3-5;81 (Schultz). *T(1;4)A7: Translocation(l;4) from Austin
new order. 1A - 2B17J3C5 - 20; cytology: T(l;4)9B;20;101-102.
61 - 77D3|2C1 - 3C4|81 - 77D5J81 - new order: 1 — 9 B | 2 0 ;
100. 10l|9B -20J102.
discoverer: Clausen. origin: X ray induced.
genetics: Variegated for w and rst and apparently discoverer. Muller, 1928.
mutant for in, but eg, Did, Dfdr, pb, and p not af- synonym: CRB; W13; T(l;4)wl3; T(l;4)l.
fected, in effect probably associated with 77D3-5 references: Muller and Stone, 1930, Anat. Record
break. Each element of the translocation survives 47: 393-94.
as an aneuploid. Df(l)wvc° = Df(l)2B17-Cl;3C4-5 Muller and Painter, 1932, Z. Induktive
is deficient for recessives from kz through the Abstammungs- Vererbungslehre 62: 316—65.
dwarf character of rst2 (i.e., rst but not vr)- Painter, 1934, Genetics 19: 448-69.
Df(l)w™°/T(l;3)wvco ^ Wf extreme rst, and genetics: X chromosome broken between Iz and ras
highly infertile. Dp(l;3)wvc0 = Dp(l;3)2B17- and between bb and the centromere. The segregant,
C1;3C4~5;77D3-5;81 covers w and tst. Dp(l;4)Al = Dp(l;4)9B;20;101-102, is lethal when
*T(l;3)y260-n: Translocation(l;3) yellow added to a normal male genotype, causing failure
cytology: T(l;3)lB2-3;8SFl-5. of separation of the germ layers (Poulson, 1940,
origin: X ray induced. J. Exptl. Zool. 83; 271-325). Segregant added to
discoverer: Sutton, 39a. a normal female or triploid intersex genotype pro-
references: 1943, Genetics 28: 210—17. duces females with low fertility (Pipkin, 1940,
genetics: Mutant for y but not BC, SC, or svr. Male Texas Univ. Publ. 4032: 126—56). The reciprocal,
viable but sterile. Df(l)Al =DfCl)9B;20, causes death associated
with incomplete blastoderm formation when added
cytology: T(l;3)6C;70E-F + In(l)lA6-7;SD8-El. to male genotype (Poulson, 1940); it results in
new order: 1A1 - 1A6|5D8 - 1A7|5E1 - 6C|70F - poorly fertile females when added to normal female
100; or triploid intersex genotypes (Pipkin, 1940).
20 - 6CJ70E - 61. *TO;4)A2
origin: X ray induced, cytology: Chromosome 4 appended to X as second
discoverer: Sutton, 1939. arm.
genetics: Mutant for y but not sc. Male lethal. discoverer: Patterson, Stone, Bedichek, and Suche,
1933.
*T(1£;4)A: Translocation(l;3;4) from Austin references: Painter and Stone, 1935, Genetics 20:
origin: X ray induced. 327-41 (fig.).
discoverer: Patters on, Stone, Bedichek, and Suche. genetics: Translocation involves short arm of one
synonym: T(l;4)3A. and base of long arm of the other. Unlike most X~4
references: Painter and Stone, 1935, Genetics 20: pseudofusions, crossing over between t and the
327-41. centromere is virtually eliminated.
*T(l;3;4)A96b *T(1;4)A3
cytology: T(1;3;4)3C3-7;IQIF; break in chromosome cytology: About 10 percent of metaphase length of X
3 not determined (Mackensoo, 1935). chromosome transferred to chromosome 4.
origin: X ray induced. references: Stone, 1934, Genetica 16: 506—20.

discoverer: Patterson, 301. Painter and Stone, 1935, Genetics 20: 327—41
synonym: T(l;4)wm^. (fig.)-
references: Patterson and Painter, 1931, Science genetics: Same as T(1;4)A5.
73: 530-31. *T(1;4)A8
Patterson, 1932, Genetics 17: 38—59. cytology: T(l;4)llA6-7 (1940, Texas Univ. Publ.
Muller and Painter, 1932, Z. Induktive 4032, frontispiece); breakpoint in chromosome 4
Abstammungs- Vererbungslehre 62: 316—65. unknown.
genetics: Variegated for w and N. X broken between origin: X ray induced.
w and ec. discoverer: Patterson, Stone, Bedichek, and Suche,
1933.
*T(1;4)A4 references: 1934, Am. Naturalist 68: 359—69.
cytology: T(1;4)13F6-14A1;1O2F Inferred from fig. Stone, 1934, Genetica 16: 506-20.
17D, E, and F of Mackensen (1935), also frontis- Patterson, Stone, and Bedichek, 1935, Genetics
piece of Texas Univ. Publ. 4032. 20: 259-79 (fig.).
discoverer: Patterson, Stone, Bedichek, and Suche, Pipkin, 1940, Texas Univ. Publ. 4032: 126-56.
1933. genetics: X chromosome broken between tw and wy
references: 1934, Am. Naturalist 68: 359-69. and chromosome 4 to the left of bt. Homozygous
Stone, 1934, Genetica 16: 506-20. viable and fertile. Both the X&4P and the 4DXP
Mackensen, 1935, J. Heredity 26: 163-74 (fig.). elements added to a normal diploid female geno-
Patterson, Stone, and Bedichek, 1935, Genetics type produce weakly fertile hyperploid females and
20: 259-79 (fig.). when added to a triploid intersex genotype produce
1937, Genetics 22: 407-26. sterile hypoploid triploid females.
Pipkin, 1940, Texas Univ. Publ. 4032: 126-56.
genetics: X chromosome broken between sd and f. *T(1;4)A9
Homozygous viable and fertile. Both the 4DXP and cytology: T(l;4)5Al-4 (1940, Texas Univ. Publ.
the reciprocal XD4P elements survive when added 4032, frontispiece).
to diploid female or intersex genotypes. X/X/4DXP origin: X ray induced.
females, but not the other genotypes, are fertile. discoverer Patterson, Stone, Bedichek, and Suche;
1933.
T(1;4)A4: see T(l;4)Bs references: 1934, Am. Naturalist 68: 359—69.
*T(1;4)A5 Stone, 1934, Genetica 16: 506-20.
cytology: Chromosome 4 appended to X as second Patterson, Stone, and Bedichek, 1935, Genetics
arm. 20: 259-79.
discoverer: Patterson, Stone, Bedichek, and Suche, Pipkin, 1940, Texas Univ. Publ. 4032: 126-56.
1933. genetics: X chromosome broken between rg and cv
references: Stone, 1934, Genetica 16: 506—20. and 4 broken to the left of bt. Homozygous viable
Painter and Stone, 1935, Genetics 20: 327-41 and fertile. The XD4P element added to a normal
(fig-)- diploid female genotype produces partially fertile
genetics: Translocation involves short arm of one hyperploid females; it survives when added to a
chromosome and base of long arm of the other. triploid intersex genotype. Its complement, 4DXP,
Crossing over at base of X normal. is virtually lethal when added to a diploid female
but produces a partially fertile hypotriploid when
T(1;4)A5: see T(l;4)wms added to triploid intersex genotype.
*T(1;4)A6 *T(1;4)AJ0
cytology: Chromosome 4 appended to X as second cytology: T(I;4)1A5-6;1O2A2~4.
arm. origin: X ray induced.
origin: X ray induced. discoverer Patterson, Stone, Bedichek, and Suche,
discoverer Patterson, Stone, Bedichek, and Suche, 1933.
1933. references: 1934, Am. Naturalist 68; 359—69.
references: Stone, 1934, Genetica 16: 506—20. Stone, 1934, Genetica 16: 506—20.
Painter and Stone, 1935, Genetics 20: 327—41 Stone and Griffen, 1940, Texas Univ. Publ. 4032:
(fig-)- 208-17 (fig.),
genetics: Same as T(1;4)A5. genetics: Homozygous viable and fertile.
*T(1;4)A7 *T(1;4)A71
cytology: Chromosome 4 appended to X as second cytology: Chromosome 4 appended to X as second
arm. arm.
origin: X ray induced. origin; X ray induced,
discoverer: Patterson, Stone, Bedichek, and Suche, discoverer Patterson,, Stone, Bedichek, &ad Suche,
1933. 1933.
references: Stone, 1934, Genetica 16: 506—20. new order: 1 - 16A7|l02F - 100;
Painter and Stone, 1935, Genetics 20: 327—41 20 - 16A1J102F.
(fig-). origin: X ray induced in Dp(l;l)B = Dp(l;l)15F9-
Brown, 1940, Texas Univ. Publ. 4032: 65-72. 16A1.16A7-16B1.
genetics: Same as T(1;4)A5. discoverer. Stone, 1931.
T(1;4)A13 synonym: T(l;4)l; T(l;4)4; T(1;4)A4.
cytology: T(l;4)18C5-Dl (1940, Texas Univ. Publ. references: 1934, Genetica 16: 506-20.
4032: frontispiece). genetics: Position effect at B more extreme than in
origin: X ray induced, treated chromosome. Male and homozygous female
discoverer: Patterson, Stone, Bedichek, and Suche, viable and fertile. The 4DXP segregant carries no
1933. known markers from chromosome 4 and Bs through
references: 1934, Am. Naturalist 68: 359-69. bb+ from X. Female hyperploid for this element
Stone, 1934, Genetica 16: 506-20. viable and fertile. Hyperploid male poorly viable
Patterson, Stone, and Bedichek, 1935, Genetics and sterile.
20: 259-79. other information: Used by Stern in cytological dem-
1937, Genetics 22: 407-26. onstration of crossing over (1931, Biol. Zentr. 51:
Pipkin, 1940, Texas Univ. Publ. 4032: 126-56. 547_87). 4DXp from T(1;4)BS used by Lindsley
genetics: X chromosome broken between (u and car and Sandier (1963, In Methodology in Basic
and in chromosome 4 to the right of ey. Homozy- Genetics, W. J. Burdette, ed. Holden-Day, Inc.
gous viable and fertile. No ci position effect. The pp. 390—403) in construction of compound-gener-
4DXP element sterile in male and fertile in female. ating Bs duplications. Reciprocal products of
Complementary XD4P produces fertile hypotriploid meiosis in male not always recovered with equal
females when added to triploid intersex phenotype; frequency (Novitski and Sandier, 1957, Proc. Natl.
it produces a virtually lethal superfemale when Acad. Sci. U.S. 43: 318-24; Zimmering, 1960,
added to normal diploid female genotype. Genetics 45: 1253—68; Zimmering and Barbour,
1961, Genetics 46: 1253—60; Zimmering and
*T(1;4)A14 Perlman, 1962, Can. J. Genet. Cytol. 4: 333-36).
cytology: Chromosome 4 appended to X as second
arm. 1: Translocation(1;4) cut
origin: X ray induced, cytology: T(1;4)7B2-3;20;101A-D superimposed on
discoverer: Patterson, Stone, Bedichek, and Suche, R(1)1A3-4;19F-2OA1.
1933. new order: |lA4 - 7B2|20-20F - 20A11;
references: Stone, 1934, Genetica 16: 506—20. 101A|7B3 - 20J101D - 102.
Painter and Stone, 1935, Genetics 20: 327-41 Tentative.
(fig-)- origin: X ray induced in R(l)2.
Brown, 1940, Texas Univ. Publ. 4032: 65-72. discoverer: Hannah, 1947.
genetics: Translocation involves short arm of one genetics: Mutant for ct but not y, ac, sc, cm, sn, or
chromosome and base of long arm of the other. oc. Male lethal.
Stone (1934), but not Brown (1940), reports strong T(7;4)/-v77: Translocation(T;4) lethal-variegated
reduction in crossing over at base of X chromo- cytology: T(1;4)15;1Q1.
some. origin: X ray induced.
T(h4)A17 discoverer Lindsley, Edington, and Von Halle,
cytology: T(1',4)7F5-8B1 (1940, Texas Univ. Publ. references: 1960, Genetics 45: 1649-70.
4032: frontispiece). genetics: Variegated for a lethal. Male fertile.
origin: X ray induced. T(l;4)m*: see T(l;4)wm5
discoverer: Mickey. *T(J;4)M-pro: Translocathn(l;4) M/nufe-proc/ocer
references: Patterson, Stone, and Bedichek, 1937, discoverer: Bridges, 33d26.
Genetics 22: 407-26. synonym: M-pro; Minute-producer.
Pipkin, 1940, Texas Univ. Publ. 4032: 126—56. genetics; Minutes produced are haplo-4's. The
genetics: X chromosome broken between t and lz, trans location causes nondis junction of chromosome
although the reported cytological breakpoint is to 4 centromeres (L. V. Morgan, 1940, DIS 13: 51).
the left of this interval. The XD4P element sur- TO;4)N264.U: Translocation(l;4) Notch
vives when added to either a normal diploid female cytology: T(l;4)3C6-7;101F (Sutton).
or a triploid intersex genotype; in the latter at origin: X ray induced,
least, the product is a fertile female. The comple- discoverer: Demerec, 34a.
mentary 4DXP is virtually lethal when added to a synonym: T(1;4)N«8.
diploid female genotype but produces partially fer- references: 1941, Proc. Intern. Congr. Genet., 7th.
tile females when added to a triploid intersex pp, 99-103.
genotype. Jodd, 1955, DIS 29: 126-27.
T(1;4)BS; Trmslocation(l ;4) Bar of Stone genetics: Carries mutant allele of N. Variegates tor
cytology: T(l;4)15F9-16Al;l6A7-Bl;102F (Griffen, w and rmt but not pn, kz, or <±n. The d + allele
1941, Genetics 26: 154-55). shows weakened dominance (Stem).
*T(l;4)N264-20 M(l)Bld and was used extensively by Sturtevant

cytology: T(l;4)3C4-5;3C7-8;101F; 3C5-7 missing (1934, 1936) in his studies on preferential segrega-
(Sutton). tion. The 4DXP element survives in the heterozy-
new order 1 — 3C4|l01F; gous female but is an extreme Minute and rarely
20 - 3C8|l01F - 102. fertile.
origin: X ray induced. T(1;4)scH: Translocation(l ;4) scute of Hack eft
discoverer: Demerec, 34g. cytology: T(1;4)1B4-C3;1O1-1O2; inferred from
genetics: Deficient for N; variegates for w and rst genetic results.
but not pn, kz, or dm. ci+ shows weakened domi- origin: Gamma ray induced.
nance (Stern). discoverer: Hackett, 46a.
*T(l;4)N264-86 references: Muller and Valencia, 1947, DIS 21: 70.
cytology: T(l;4)3C6-7;3C7-8;3E5-6;101F. genetics: Two halves of the translocation may be
new order: 1 - 3C7|3E6 - 20; recovered separately. 4DXP is deficient for y and
101A - 101F|3C7 - 3E5|l01F - 102; sc but not M(l)Bld and carries ey+. XD4P covers
band 3C7 present twice and considered Df(l)sc19 and therefore carries a normal allele of
to have been from each of two chromatids in the l(l)sc. T(l;4)scIi/ci is ci+.
sperm (Demerec and Sutton, 1940, Proc. Natl. T(l;4)w**: see T(1;4)A1
Acad. Sci. U.S. 26: 532-36). *T(l;4)w2S8-43
origin: X ray induced. cytology: T(l;4)3C3-5;102F4-5 (Demerec).
discoverer: Demerec, 39i. origin: X ray induced.
references: 1940, Genetics 25: 618-27. discoverer. Demerec, 38k.
Sutton, 1940, Genetics 25: 534-40 (fig.). genetics: Mutant for w but not kz, pn, rst, or (a.
genetics: Carries two AT loci, one mutant and one Male lethal.
variegated. Also carries a mutant allele of rst : see T(1;4)A3
(rst264~s6) and variegates for dm but not w or ec.
Carries normal allele of ci + (Stern). T(1;4)w">*: T(l;4) white-mottled
Dp(l;4)N264-s6 = Dp(l;4)3C6-7;3E5-6;101F-102 cytology: T(l;4)3C3-4;101Fl-2 (Griffen and Stone,
viable and fertile in both sexes; D((1^264-86 = 1938, Genetics 23: 149).
Dt(l)3C7-8;3ES-6 viable in heterozygote. origin: X ray induced.
*T(1;4)N 264-113 discoverer: Muller, 1929.
cytology: T(l;4)3C10-Dl;101; section 102 missing. synonym: T(l;4)m5: Translocation(l;4) mottled-5;
new order: 1 — 3ClQ|l01; T(1;4)A5.
20 - 3Dl|?. references: 1930, J. Genet. 22: 299-334.
Proximal portion of X chromosome considered to be Bolen, 1931, Am. Naturalist 65: 417-22.
terminally deficient although it occasionally genetics: Variegates for w and ci. [Dubinin,
appears to be capped by a small nucleolus-like Sokolov, and Tiniakov, 1935, Biol. Zh. (Moscow)
structure [Sutton, 1940, Genetics 25: 628—35 4: 716—20]. X chromosome broken between 1(1)3C3
(fig.)]. Not clear that a reciprocal translocation and rst, and chromosome 4 broken to the left of ey.
between X and short arm of 4 was adequately ruled XD4P added to a normal male genome produces
out. males with 20 percent normal viability that are
origin: X ray induced. weakly fertile; added to a diploid female genome,
discoverer. Demerec, 40c. it produces fertile hyperploid genome; but added to
genetics: Variegates for w and N but not ac, sc, dm, a triploid inters ex genome is virtually lethal.
or ec. Carries normal allele of ci (Stern). 4DXP is inviable when added to a male genome, is
T(1;4)N«*: see T(1;4)N264-12 virtually lethal when added to a female genome; it
*70-r4)sc102: Translocation(l;4) scute produces rather fertile hypotriploid females when
cytology: T(1;4)1D;1O1E-F [+ T(1;4)1A;1O2F ac- added to a triploid inters ex genome (Pipkin, 1940,
cording to Schultz]. Texas Univ. Publ. 4032: 126-56). Griffen and
new order. lA|l02F - 10lF|lD - 20; Stone (1940, Texas Univ. Publ. 4032: 190-200)
101A - 101F|lD - 1A|1O2F; produced and studied a number of X-ray-induced
Tentative. derivatives of T
origin: X ray induced in ac3.
discoverer: Sturtevant, 1930. cytology: T(l;4y3C3-4;i01A-D,
references: 1934, Proc. Natl. Acad. Sci., U.S. 20: origin: X ray induced.
515-18. discoverer: Panshin.
1936, Genetics 21: 444-66. references: Pmnshin and Khvostov®, 1938, Biol. Zh.
genetics: Mutant for sc. Virtually male lethal. X (Moscow) 7: 359-80.
chromosome broken between M(l)Bld and &*y chro- Pan*liin, 1938, Nature 142: 837.
mosome 4 broken proximal to ci. According to 1941, DIS 15: 33-34.
Schultz, both X and 4 also have breaks distal to genetics: Variegated for w but not ci. First rear-
all known loci and their termini are interchanged. mftgemeat to involve, mad therefor© to demonstrate,
XD4P carries X-chxocnosotoe loci from y through existence of 4L. Panshin and Khvostova [l938;
Panshin, 1938, Biol. Zh. (Moscow) 7: 837-65] pro- new order: 1A - 3Cl|l01E - 101A;
duced and studied a number of X-ray-induced deriv- 20 - 3C2|?.
atives of T(l;4)wm11. origin: X ray induced.
cytology: T(l;4)3Cl-2;3C4-7;20A;101. references: Sutton, 1940, Genetics 25: 628—35.
new order: 1 - 3Cl|20A - 3C7|20A - 20F; genetics: Variegated for w but not pn, rst, or spl.
10l|(3C2 -3C4)|l01 - 102. Male viable. Translocation-bearing fly carries two
origin: X ray induced in In(l)w™4 = In(l)3Cl-2;20A. normal fourth chromosomes.
discoverer: Lefevre, 51c2O. *lC\;4)vtmAt Translocation(l;4) white-mottled from
references: 1951, DIS 25: 71. Austin
1952, DIS 26: 66. cytology: T(1;4)3C2-3;101A2-3.
Ratty, 1954, Genetics 39: 513-28. origin: X ray induced.
genetics: Variegated for w and rst. Male lethal. discoverer: Stone.
Dp(l;4)wmSlc =Dp(l;4)3C2-3;3C4-7;101 viable and references: Griffen and Stone, 1939, Genetics 24:
fertile; carries loci of w and rst but not spl. 73.
*T(l;4)w»>52bU 1940, Texas Univ. Publ. 4032: 201-7 (fig.)-
cytology: T(l;4)2A2-3;3C3-4;20B;101. genetics: Variegated for w. Male viable and fertile.
new order: 1 - 2A2|20B - 3C4|20B - 2OF; Second demonstration of the existence of a left arm
10l|(2A3 -3C3)|l01 - 102. on chromosome 4.
origin: X ray induced inln(l)rst3 =In(l)3C3-4;20B. T(l;4)wmD3: Translocation(T;4) white-mottled of
discoverer: Ratty, 52bl3. Dubinin
references: Lefevre, 1953, DIS 27: 57. cytology: T(l;4)3C;101.
genetics: Variegated for w. discoverer: Dubinin.
T(l;4)w»258-18 references: Sacharov, 1936, Biol. Zh. (Moscow) 5:
cy to I ogy: T(l ;4)3 C4-5; 101. 293-302.
origin: X ray induced. genetics: Variegated for w.
discoverer: Demerec, 33k. *T(1;4)w">DV4: Translocation(l;4) white-mottled
references: Demerec and Slizynska, 1937, Genetics of Dubinin and Volotov
22: 641-49. cytology: T(l;4)3C3-7;3D;101A-D; 3C-3D missing;
genetics: Variegated for w and rst but not pn, fa, inferred from genetic data and from fig. 5, 6, and 7
dm, or ec. Also variegated for ci (Stern). Male and of Sacharov (1936), which indicate that the break
homozygous female viable and fertile. X chromo- in chromosome 4 is in the left arm.
some broken between rst and vt (Gersh, 1965, new order: 1 - 3C3|l01D - 102F;
Genetics 51: 477-80). The X®4P element sur- 20 - 3D|l01A.
vives as a duplication. discoverer: Dubinin and Volotov.
references: Sacharov, 1936, Biol. Zh. (Moscow) 5:
293-302 (fig.).
cytology: T(l;4)3B5-6;101F (Demerec and Hoover). genetics: Deficient for N; variegated for w. Male
origin: X ray induced. lethal. Since the XD4P element of T(l;4)waiDV4
discoverer: Demerec, 1934. survives as a duplication and carries wmDV4, the
synonym: T(l;4)wvD3: Tranalocation(l;4) white- left break in X chromosome is between w and N.
variegated of Demerec. T(l;4)w>"J: Translocation(};4) white-mottled of
genetics: Variegates for w, fa, spl, N, dm, and Jonsson
M(1)3E but not ec or W. Also variegates for ci cytology: T(l;4)3C2-3;20;102C.
(Gersh). Males usually lethal; survivors probably new order 1 - 3C2|20 - 3C3|lO2C - 101 A;
X/Y/Y. Cell lethal in X/0 tissue in gynandro- 2OjlO2C - 102F.
morphs (Judd, 1953, DIS 27: 95). origin: X ray induced,
*T( 7;4)w» 258-31 discoverer Jonsson, 61i28.
cytology: T(l;4)3C3-5;102F4-17 (Demerec and references: Lefevre, 1963, DIS 37: 49.
Hoover). Lefevre and Wilkins, 1966, Genetics 53: 175-87.
origin: X ray induced. genetics: Variegated for w. The 4DXP element of
discoverer: Demerec, 371. the translocation has become separated from the
genetics: Variegated for w but not rst. Male viable. XD4P element and lost. The XD4P element is vi-
able as an X/Y male but lethal as an X/0 male,
cytology: T(1;4)3C1-2;1O1E-F; distal part of chro- probably owing to deficiency for 06. Additional
mosome 4 lost. Sutton thought it a terminal defi- evidence for appreciable deficiency for proximal X
ciency of X. Evidence that chromosome 4 is heterochromatin is virtually random disjunction of
involved seems equivocal, especially since, ac- X and y chromosomes. XD4P carries ci + but not
cording to events postulated, the original mottled ey+. The variegation of white is unorthodox be-
fly should have been haplo-4. Alternative interpre- cause heterochroraatin has been moved to the white
tation is translocation between X and Y in X/Y locus rather than white moved into proximal heter-
sperm. ochromatin.
*T(l;4)w"Med: Translocation(l;4) white-mottled of genetics: Break between pr and tk. rl, M(2)S2, stw,
Medvedev ap, msf, tk, and ltd not affected.
discoverer: Medvedev, 1934. *T(Y;2)A3: Translocation(Y;2) from Austin
genetics: Variegated for w and probably rst. Arose origin: X ray induced.
in wa and therefore has light eye color. discoverer: Stone,
genetics: Variegated for bw.
; Transloccttion(l;4) zeste *T(Y;2)B
cytology: T(l;4)3Cl-2;102F2-4; genetic data more in cytology: T(Y;2)4OF-41A1, placed in 2R by
accord with breakpoint in 3C2-3 than 3C1-2. Whittinghill (1937, DIS 8: 82).
origin: X ray induced in a chromosome carrying z. origin: X ray induced.
discoverer: Gans. discoverer: Dobzhansky, 1929.
references: 1953, Bull. Biol. France Belg. Suppl. references: 1930, Biol. Zentr. 50: 671—85.
38: 1-90 (fig.). 1932, Z. Induktive Abstammungs- Vererbungslehre
Gersh, 1963, DIS 37: 80. 60: 235-86.
genetics: Suppresses z. The X&4*3 element is genetics: Lethal in combination with M(2)S210 and
poorly viable when added to male genome but vi- shows an extreme rl phenotype with rl. stw, ap,
able and fertile in female; duplicated for w but msf, tk, and ltd not affected.
does not cover lethality of Df(l)wm4Lrst3R = *T(Y;2)bw + Y: Translocation(Y;2) brown-wild
Df(l)3Cl-2;3C3-4 (Gersh, 1963). typeY
cytology: T(Y;2)Y^;58F1-59A2;6OE3-F1 (Gersh,
*T(l;A)pn-ec: Translocationfl; Autosome) prune- 1956, DIS 30: 115; Nicoletti).
echinus new order: Y L D |(59A2 - 6OE3)|yLP _ Y s ;
cytology: T(1;A)2D1-2;3F7-4A1;4O-41 or 50-52; posi- 21 - 58Fl|60Fl - 60F5.
tion of autosomal breakpoint not determined. origin: X ray induced.
new order: 1 - 2D1 |4A1 - 20; discoverer: Dempster.
21 _ 4O|(2D2 - 3F7)|41 - 60; references: Brosseau, Nicoletti, Grell, and
(for example). Lindsley, 1961, Genetics 46: 339—46.
origin: X ray induced. genetics: Dp(2;Y)bw+ carries loci from bw through
discoverer: Robins, 62g26. ba but not hv or M(2)c; it is used as a marked Y
references: Lefevre, 1963, DIS 37: 50. and referred to as bw+Y.
genetics: w not affected. Male lethal. Female het-
erozygous for the segregant Df(l)pn-ec = Df(l)2Dl- *T(Y;2)bwR27: Translocation(Y;2) brown-
2;3F7~4A1 survives though poorly viable and fer- Rearranged
tile. Male heterozygous for complementary cytology: T(Y;2)59D11-E1.
Dp(l;A)pn-ec = Dp(l;A)2Dl-2;3F7-4Al;40-41 or origin: X-ray-induced derivative of bw.
80-81 viable but sterile. discoverer: Slatis.
T(X-2)In* end 3: S ee T(l;2)wtn53e references: 1955, Genetics 40: 5—23.
: see T(l;2)26 genetics: Associated with bw**27.
*T(Y;2)bw**7
*T(Y;2)21E cytology: T(Y;2)59D5-6.
cytology: T(Y;2)21D4-E1. origin: X ray induced.
discoverer: Schultz. discoverer: Slatis.
references: Lewis, 1945, Genetics 30: 137—66. references: 1955, Genetics 40: 5—23.
genetics: Not mutant for S or ast. Chromosome 2 genetics: Associated with bwR57.
broken between ds and S. Both 2LDYP and YD2P T(Y;2)C
recoverable in aneuploid progeny. cytology. T(Y;2)4OF-41A1; placed in 2R by
T(Y;2)54a Whittinghill (1937, DIS 8: 82-84).
cytology: T(Y;2)YL;59C4-6. origin: X ray induced.
discoverer: Mickey, 54a. discoverer: Dobzhansky, 1929.
references: 1959, Texas Univ. Publ. 5914: 99-105- references: 1930, Biol. Zentr. 50: 671-85.
1963, DIS 38: 30. 1932, Z. Induktive Abstammungs- Vererbungslehre
genetics: Variegated for bw. Male fertile. Male 60: 235-86.
hyperploid for YF2RD survives, is not variegated, genetics: Does not affect d, M(2)S2, stw, ap, msf,
and is sterile. tk, or ltd.
T(Y;2)A T(Y;2)D: see T(Y;2;3)D
cytology: T(Y;2)4OF-41A1; placed in 2R by T(Y;2)dp *' •*; Tronslocation(Y;2) dumpy
Whittinghill (1937, DIS 8: 82-84). origin: X ray induced.
origin: X ray induced, discoverer: Thompson, 61d.
discoverer: Dobzhansky, 1929. genetics: Mutant for dp.
references: 1930, Biol. Zentr, 50: 671-85. *T(Y;2)o|pw2: Tr<mslocathn(Y;2) dumpy-warped
1932, Z. Induktive AbstaBunung*- Vererbungslehre origin: X ray induced.
60: 235-86. discoverer: Schalet, 55k.
references: Carlson and Schalet, 1956, DIS 30: 71. origin: X ray induced.
Carlson, 1958, DIS 32: 117-18. discoverer: Slatis.
genetics: Variegated for dp. references: 1955, Genetics 40: 8.
T(Y;2)E genetics: Induced simultaneously with (but indepen-
cytology: T(Y;2)36D2-3 (Whittinghill, 1937, DIS 8: dently of) bw24, an isoallele of bw. Associated
82-84). with a rough-eye phenotype. Male hyperploid for
origin: X ray induced. Dp(2;Y)R24 is viable but sterile.
discoverer: Dobzhansky, 1929. T(Y;2)w+Y
references: 1930, Biol. Zentr. 50: 671-85. Described as w+Y in subsection on Y derivatives.
1932, Z. Induktive Abstammungs- Vererbungslehre
60: 235-86. *T(Y;2;3)D
genetics: Male fertile, but Df(l)sc4Lsc8R/T(Y;2)E cytology: T(Y;2;3)29F-3OA1 + T(2;3)34C;78F +
male is sterile. Df(2R)41A;41C + Df(3L)61E2-Fl;62A4-6. May also
T(Y;2)F: see T(Y;2;3)F carry small inverted segment in region 41
T(Y;2)G (Whittinghill, 1937, DIS 8: 82-84).
cytology: T(Y;2)36B5-C1;4OF; metaphase chromo- new order: YD|30Al - 34C|78F - 100;
somes appear normal (Morgan, Bridges, and Y p |29F - 2 1 ;
Schultz, 1935, Carnegie Inst. Wash. Year Book 34: 60 - 41C|41A - 34C|78F - 62A6|61E2 -
287). 61A.
new order. YD|(36C1 - 40F)|yP; origin: X ray induced.
21 - 36B5|40F - 60. discoverer: Dobzhansky, 1929.
origin: X ray induced. synonym: T(Y;2)D.
discoverer: Dobzhansky, 1929. references: 1930, Biol. Zentr. 50: 671-85.
references: 1930, Biol. Zentr. 50: 671-85. 1932, Z. Induktive Abstammungs- Vererbungslehre
Rhoades, 1931, Genetics 16: 490-504. 60: 235-86.
genetics: Dp(2;Y)G has normal phenotype and is genetics: Deficient for M(2)S2 and stw, but not rl,
fertile when hyperploid in either sex; duplicated for ap, msf, tk, or ltd in chromosome 2 and for ru, aa,
the loci of M(2)m, M(2)H, hk, pr, Bl, It, and the and ve but not su(ve) or R in chromosome 3. The
lethal of bwV326 but not rd. 2LDYP element survives in hyperploids.
*T(Y;2)H T(Y;2;3)F
cytology: T(Y;2)37Bl-2;4QB2-3; also an inversion in origin: X ray induced.
2R from near centromere to left of px (Morgan, discoverer: Dobzhansky, 1929.
Bridges, and Schultz, 1935, Carnegie Inst. Wash. synonym: T(Y;2)F.
Year Book 34: 287). references: 1930, Biol. Zentr. 50: 671—85.
new order. YD|(37B2 - 40B2)|YP; 1932, Z. Induktive Abstammungs- Vererbungslehre
21 - 37B1J40B3 - | - | - 60. 235-86.
origin: X ray induced. genetics: Break in 2R to right of sp.
discoverer: Dobzhansky, 1929. *T(Y;2;3)I
references: 1930, Biol. Zentr. 50: 671—85. cytology: T(Y;2)47A2-3 + T(Y;3)91E2-4 +
Schultz and Bridges, 1932, Am. Naturalist 66: In(3LR)6 9C2-3;84E2-3 + In(3LR)74A -Bl; 99 C
323-34. (Whittinghill, 1938, DIS 8: 82-84).
genetics: Male fertile. Homozygote viable but male new order. YDJ47A2 - 21;
sterile. D£(2L)H survives and is deficient for YD|91E4 - 99CJ74B1 - 84E2|69C2 - 61;
M(2yfi, hk, and pr but not M(2)m or It; somewhat 60 - 47A3|YP|91E2 - 84E3|69C3 -
sterile. Dp(2;Y)H appears normal; duplicated for 74A|99C - 100.
the loci for which Di(2L)H is deficient. origin: X ray induced.
T(Y;2)J discoverer: Dobzhansky, 1929.
cytology: T(Y;2)40F-41Al;57Fl-2 (Whittinghill, references: 1930, Biol. Zentr. 50: 671-85.
1937, DIS 8: 82-84). 1932, Z. Induktive Abstammungs- Vererbungslehre
new order. YD|40F - 21; 60: 235-86.
YP|57F1 - 41A1J57F2 - 60.
origin: X ray induced. *T(Y;3)42i
discoverer: Dobzhansky, 1929. cytology: Break in middle of one arm of chromosome
references: 1930, Biol. Zentr. 50: 671—85. 3.
1932, Z. Induktive Abstammungs- Vererbungslehre origin: X ray induced.
60: 235-86. discoverer Poulson.
genetics: Does not affect rl, M(2)S2, ntw, ap, mat, references: 1943, DIS 17: 51.
tk, or ltd. *T(Y;3)HS8b: Tronslocathn(Y;3) Hairless
*T(Y;2)R24 origin: Gemma ray induced.
cytology: T(Y;2)4SA;51E. discoverer: Ives, 58b25.
new order: YD|(45A - 51E)|yP; references: 1959, DIS 33: 95.
21 - 45AI51E - 60. genetics: Mutant for H.
*T(Y;3)I origin: Gamma ray induced.

cytology: T(Y;3)YS;63C;72E. discoverer: C. Hinton, 63b.
new order: YL - YSPJ63C - 72E|63C - 61; references: 1964, DIS 39: 61.
Y SD |72E - 100. 1965, Genetics 51: 971-82.
origin: X ray induced, genetics: Homozygous lethal.
discoverer: Muller. 1(2:3)63-5
references: Painter and Muller, 1929, J. Heredity 20: cytology: T(2;3)40C;89E-F + In(3L)69-70;79-80.
287-98. new order: 21 - 40C|89E - 80|70 - 79J69 - 61;
Muller, 1930, J. Genet. 22: 299-334. 60 - 40C[89F - 100.
Mohr and Mossige, 1940, Hereditas 26: 202—8 origin: Gamma ray induced.
(fig-). discoverer: C. Hinton, 63b.
genetics: Right break in 3L between ffi and s£. The references: 1965, Genetics 51: 971—82.
3LDYP element recoverable in hyperploid and du- T(2;3)63-6
plicated for loci from ni through th. cytology: T(2;3)59E-F;89E-F.
*T(Y;3)K4: Translocation(Y;3) of Krivshenko origin: Gamma ray induced.
cytology: T(Y;3)Ys;8l; inferred from metaphase discoverer: C. Hinton, 63b.
cytology. Ganglion metaphases show break in Ys references: 1964, DIS 39: 61.
distal to 66 and break in 3R near centromere. 1965, Genetics 51: 971-82.
origin: X ray induced. genetics: Homozygous viable and fertile. Short
discoverer: Krivshenko, 59b7. bristles; wings obliquely creased, ovate, and often
references: 1959, DIS 33: 96. asymmetrical.
genetics: Homozygous viable but apparently sterile. 1(2,3)63-7
T(Y;3)P8Q: Translocation(Y;3) from Pasadena cytology: T(2;3)41C;92D-E.
cytology: T(Y;3)88C-F. origin: Gamma ray induced,
discoverer: E. B. Lewis. discoverer: C. Hinton, 63b.
T(Y;3)P102 references: 1964, DIS 39: 61.
cytology: T(Y;3)87B2-3. 1965, Genetics 51: 971—82.
discoverer: E. B. Lewis. genetics: Homozygous lethal.
*T(Y;3)srl00.23: Translocation(Y;3) stripe T(2;3)63-8
cytology: T(Y;3)90E2-3. cytology: T(2;3)36E;86B.
origin: X ray induced. origin: Gamma ray induced.
discoverer: Alexander. discoverer. C. Hinton, 63b.
42-54. 1965, Genetics 51: 971-82.
genetics: Mutant for sr. genetics: Homozygous lethal.
*T(Y;3)sf*00.i26: Translocation(Y;3) scarlet T(2;3)63-9
cytology: T(Y;3)73A2-3. cytology: T(2;3)34A-B;75C.
origin: X ray induced. origin: Gamma ray induced,
discoverer: Alexander. discoverer. C. Hinton, 63b.
42-54. 1965, Genetics 51: 971—82.
genetics: Mutant for st. genetics: Homozygous lethal.
T(Y;4) * 1(2:3)63-10
Described as 4Y in subsection on Y derivatives. cytology: T(2;3)33'34;76D-E.
T(2;3)63-1 origin: Gamma ray induced.
cytology: T(2;3)49D-E;79B-C. discoverer: C. Hinton, 63b.
origin: Gamma ray induced. references: 1964, DIS 39: 61.
discoverer. C. Hinton, 63b. 1965, Genetics 51: 971-82.
references: 1964, DIS 39: 61. genetics: Homozygous viable but sterile. Abdominal
1965, Genetics 51: 971-82. tergite pigmentation more intensive and extensive
genetics: Homozygous viable and fertile. Eyes than wild type.
slightly rough. 1(2:3)63-13
T(2;3)63-2 cytology: T(2;3y24-2S;94D-E.
cytology: T(2;3)27B-C;75C. origin: Gamma ray induced.
origin: Gamma ray induced, discoverer: C. Hinton, 63b.
discoverer: C. Hinton, 63b. references; 1964, DIS 39: 61.
references: 1964, DIS 39: 61. 1965, Genetics 51: 971-82.
1965, Genetics 51: 971-82. genetics: Homozygoos viable and fertile. Eye color
genetics: Hcmozygous lethal. variegated.
TX2;3)63-3 1(2:3)63-14
cytology: T(2;3)40-41;8Q-81; inferred from genetic cytology: T(2;3)38A-B;69A-B,
results since salivary chromosomes appear normal. origin: Gamma ray induced.
discoverer: C. Hinton, 63b. origin: Gamma ray induced.

references: 1964, OIS 39: 61. discoverer: C. Hinton, 63b.
1965, Genetics 51: 971-82. references: 1964, DIS 39: 61.
genetics: Homozygous lethal. 1965, Genetics 51: 971-82.
7(2,3)63-15 genetics: Homozygous lethal.
cytology: T(2;3)41D;64A. T(2;3)64-31
origin: Gamma ray induced. cytology: T(2;3)36D-E;96B-C + In(2R)41E-F;55F.
discoverer: C. Hinton, 63b. new order: 21 - 36D|96B - 61;
references'. 1964, DIS 39: 61. 60 - 55F|41F - 55F|41E - 36E|96C -
1965, Genetics 51: 971-82. 100.
genetics: Homozygous lethal. Eye color variegated origin: X ray induced.
in heterozygote. discoverer: C. Hinton, 1964.
T(2;3)63-16 references: 1965, Genetics 51: 971-82.
cytology: T(2;3)41C~D;93A-B. 1(2,3)64-32
origin: Gamma ray induced. cytology: T(2;3)35D-E;70C-D.
discoverer: C. Hinton, 63b. origin: X ray induced.
references: 1964, DIS 39: 61. discoverer: C. Hinton, 1964.
1965, Genetics 51: 971-82. references: 1965, Genetics 51: 971-82.
genetics: Homozygous viable and fertile. Eyes 7(2,3)64-33
slightly rough. cytology: T(2;3)40-41;80-81; inferred from genetic
T(2;3)63-17 data since salivary chromosomes appear normal.
cytology: T(2;3)40C;96A-B. origin: X ray induced.
origin: Gamma ray induced. discoverer: C. Hinton, 1964.
discoverer: C. Hinton, 63b. references: 1965, Genetics 51: 971—82.
references: 1964, DIS 39: 61. T(2;3)64-34
1965, Genetics 51: 971-82. cytology: T(2;3)25D;86C.
genetics: Homozygous lethal. Eye color in hetero- origin: X ray induced.
zygote variegated over SM5 but normal over bw and discoverer: C. Hinton, 1964.
+. Possibly contains light-mottled. references: 1965, Genetics 51: 971-82.
7(2;3)63-18
cytology: T(2;3)39B-C;80C. T(2;3)64-35
origin: Gamma ray induced. cytology: T(2;3)40B;92C.
discoverer: C. Hinton, 63b. origin: X ray induced.
references: 1964, DIS 39: 61. discoverer C. Hinton, 1964.
1965, Genetics 51: 971-82. references: 1965, Genetics 51: 971-82.
genetics: Homozygous viable and fertile. Ninety 7(2.3)64-36
percent have troughlike wing posture. cytology: T(2;3)40D;85E.
7(2.3)63-19 origin: X ray induced.
cytology: T(2;3)24D-E;80C. discoverer: C. Hinton, 1964.
origin: Gamma ray induced. references: 1965, Genetics 51: 971—82.
discoverer: C. Hinton, 63b. 7(2,3)64-37
references: 1964, DIS 39: 61. cytology: T(2;3)60E;82F.
1965, Genetics 51: 971-82. origin: X ray induced.
genetics: Homozygous lethal. discoverer: C. Hinton, 1964.
7(2;3)63-21 references: 1965, Genetics 51: 971-82.
cytology: T(2;3)32E;89C-E + ht(3LR)65B;84B. *7(2;3)100r20
new order: 21 - 32E|89C - 84B|65B - 84B|65B - cytology: T(2;3)35B2;3;40;80 superimposed on
61; Dp(l;3)3B4-Cl;4B4-5;80.
60-32E|89E - 100. new order 21 - 35B2|4B4 - 3Cl|80 - 100;
origin: Gamma ray induced. 60 - 40J35B3 - 40|80 - 61.
discoverer: C. Hinton, 63b. origin: X ray induced in Dp(l;3)N264-100,
references: 1965, Genetics 51: 971-82. discoverer: Gersh, 1959.
7(2;3)63-22 references: 1959, Genetics 44: 163-72.
cytology: T(2;3)40B;84D. genetics: Selected as a partial reversion from white-
origin: Gamma ray induced, mottled.
discoverer: C. Hinton, 63b. 7(2;3)101
references: 1964, DIS 39: 61. cytology: T(2;3)44B;83E-F (Lewis, 1956, DIS 30:
1965, Genetics 51: 971-82. 130).
genetics: Homozygous lethal. discoverer: Sturtevant.
7(2,-3)63-23 genetics: Homozygous viable; male fertile but fe-
cytology: T(2;3)40~41;80-81; inferred from genetic male sterile. Crossing over about normal in chro-
data since salivary chromosomes appear normal. mosome 2 of heterozygous female.
*T(2;3)103 references: Patterson, Stone, Bedichek, and Suche,

discoverer: Sturtevant. 1934, Am. Naturalist 68: 359-69.
genetics: Homozygous lethal. Reciprocal transloca- Pipkin, 1940, Texas Univ. Publ. 4032: 73-125.
tion with breaks in 2L and 3L. Crossing over in genetics: Homozygous viable and fertile. Chromo-
heterozygous female low in 2L, normal in 2R. somes 2 and 3 broken at chromocenter. 2L attached
T(2;3)108 to 3R and 3L to 2R.
cytology: T(2;3)37-40;42A2-3;52D-F;58A4-B1;80;81; *T(2;3)A26
inferred from a combination of cytological (52D-F origin: X ray induced.
by Lewis, 1951, DIS 25: 108-9) and genetic obser- discoverer: Muller.
vations. references: Painter and Muller, 1929, J. Heredity
new order: 21 - 37|(80 - 81)|52D - 42A3|58B1 - 20: 287-98.
60; Muller, 1930, J. Genet. 22: 299-334.
61 - 801(40 - 42A2|58A4 - 52F)|81 - genetics: Break in 3R between sr and e.
100.
origin: Arose in In(2R)Cy = Jn(2R)42A2-3;58A4-Bl. T(2;3)AntpYt>: Translocation(2;3) Antennapedta
discoverer: Sturtevant. ofYu
genetics: Mutant for Rev. Homozygous semilethal. cytology: T(2;3)22B;83E-F + T(2;3)38B;98A.
The segregant that receives a normal chromosome 2 new order: 21 - 22B|83F - 98A|38E - 22B|83E -
and the translocated element that might be desig- 61;
nated 3LD2P3RD survives and is fertile. It is du- 60 - 38E|98A - 100.
plicated for the loci of pr, It, rl, tk, and according origin: X ray induced,
to E. B. Lewis, for M(2)S7, sm, and hy; not defi- discoverer: Yu, 1948.
cient for chromosome 3 genes. references: 1949, Ph.D. Thesis, Calif. Inst.
Technol.
T(2;3)109 Lewis, 1956, DIS 30: 76.
cytology: T(2;3)22F-23B;55F-56A;80 (Lewis, 1951, genetics: Mutant for Antp; associated with 83E-F
DIS 25: 108-9). breakpoint. Homozygous lethal.
new order: 21 - 22F|55F - 23B|80 - 61;
60 - 56A|80 - 100. T(2;3)apX°: Translocafion(2;3) apterous-Xasta
discoverer: Sturtevant. cytology: T(2;3)41F;89E8-F1 superimposed on
genetics: Homozygous viable and wild type. Origi- In(2R)42A2-3;58A4-Bl + In(3R)89C2-3;96A18-l 9
nated in In(3R)P but separable from it. (Bridges in Morgan, Bridges, and Schultz, 1936,
*T(2;3)110 Carnegie Inst. Wash. Year Book 35: 294, with cor-
origin: X ray induced. rection by Lewis, 1951, DIS 25: 108-9).
discoverer: Sturtevant. new order: 21 - 41F|89E8 - 89C3|96A19 - 100;
genetics: Homozygous lethal. Wings short, ex- 60 - 58Bl|42A3 - 58A4|42A2 -
tended, coiled downward in spiral. L4 and marginal 41F|89F1 - 96A18|89C2 - 61.
veins thickened, L4 sometimes not reaching margin; origin: X ray induced in In(2R)Cy; In(3R)P.
posterior wing cell reduced. Posterior crossvein discoverer: Serebrovsky, 28a.
absent; L5 reduced and irregularly plexate. Break synonym: T(2;3)Xa: Translocation(2;3) Xasta.
in 2R near vg and one in 3R, which carries references: Serebrovsky and Dubinin, 1930, J.
In(3R)P. New order is 2L + 3L and 2R + 3R. Heredity 21: 259-65.
Sturtevant, 1934, DIS 2: 19.
*T(2;3)13S genetics: Dominant mutant for ap. Homozygote vir-
cytology: T(2;3)37;85A. tually lethal.
origin: X ray induced simultaneously with T(l;2)l- other information: The first X-ray-induced mutation
vl35. recovered in the USSR. Useful as a balancer of 2R
discoverer: Lindsley, Edington, and Von Halle, and 3R.
references: I960, Genetics 45: 1663. *T(2;3)as1 r*1; Translocation(2;3) asteroid-reverted
T(2;3)A cytology. T(2;3)21E2-3;68C2-3;8SD8-9.
cytology: T(2;3)39B-C;83B (Lewis, 1951, DIS 25: new order. 21A - 21E2|88D8 - 68C3J88D9 - 100;
108-9). 61 - 68C2|21E3 - 60.
origin: X ray induced in Bl. origin: X ray induced in al ast ho.
discoverer: Dobzhansky, 28h. discoverer: E. B. Lewis, 1942.
references: 1929, Biol. Zentr. 49: 408—19. references: 1945, Genetics 30: 158.
1933, Z, Induktive Abstammungs- Vererbunigslehre genetics: Associated with a reversion of ami. Homo-
64: 269-309. zygous lethal.
Dobzhansky and Sturtevant, 1931, Carnegie last.
Wash. Publ. No. 421: 29-59. cytology: T(2;3)21E2-3;61C2-3,
genetics: Homozygous lethal. origin: X my induced in net ast dp cl.
*T(2;3)A1: 7"ranslocalhn(2;3) from Austin discoverer: E. B. Lewis, 1942.
origin: X my induced. references: 1945, Genetics 30: 158.
genetics: Associated with reversion of ast. Lethal references: Glass, 1933, J. Genet. 28: 69—112.
homozygous and heterozygous with Df(2L)S4 = 1934, Am. Naturalist 68: 107-14.
Df(2L)21 C3-4;22B2-3. genetics: Variegated for bw. Break in 2R near bw
*T(2;3)Ata: Translocation(2;3) Arista and in 3L near centromere.
cytology: T(2;3)40;66F-67A + T(2;3)47;81.
new order: 21 - 40|67A - 81J47 - 60; origin: X ray induced.
61 - 66F|40-47|81 - 100. discoverer: Patterson,
origin: X ray induced. references: Glass, 1933, J. Genet. 28: 69-112.
discoverer: Krivshenko, 1949. 1934, Am. Naturalist 68: 107-14.
synonym: T(2;3)At (symbol preoccupied). genetics: Variegated for bw. Break in 2R near bw
references: 1954, DIS 28: 74-75. and in 3L near centromere.
1955, DIS 29: 73. *T(2;3)bwY*
genetics: Associated with Ata. Homozygous lethal. origin: X ray induced in bw.
T(2;3)B discoverer: Moore, 1929.
cytology: T(2;3)33;S1F (Lewis, 1951, DIS 25: references: Glass, 1933, J. Genet. 28: 69-112.
108-9; 1954, Am. Naturalist 88: 225-38). 1934, Am. Naturalist 68: 107-14.
origin: X ray induced. genetics: Crossing over reduced in 2L, 2RJand base
discoverer: Dobzhansky, 28h. of 3R. Probably breaks in all three arms.
references: 1929, Biol. Zentr. 49: 408-19. other information: Eye color reverted to wild type,
Dobzhansky and Sturtevant, 1931, Carnegie Inst. but translocation remained.
Wash. Publ. No. 421: 29-59. *T(2;3)bwV8
genetics: Homozygous lethal. Crossing over re- origin: X ray induced.
duced in 2L. discoverer: Levy, 1932.
genetics: Variegated for bw. Break in 2R at bw and
*T(2;3)bwR4: Translocation(2;3) brown-Rearranged 3R near p.
cytology: T(2;3)59E2-3;80-81. *T(2;3)bwV30ki2
origin: X-ray-induced derivative of few. origin: X ray induced.
discoverer: Slatis. discoverer: Van Atta, 30kl2.
genetics: Associated with bwR4. genetics: Variegated for few. Complex rearrangement
*T(2;3)bw*i2 with break in 2R near few, near centromere of 2, in
cytology: T(2;3)59D;80C. 2L, and 3L near centromere; also appears to carry
origin: X ray induced. an inversion in 3R.
discoverer: Slatis. *T(2;3)bwV30ki3
genetics: Associated with bwR12. discoverer: Van Atta, 30kl3.
*T(2;3)bw*u references: 1932, Genetics 17: 637—59.
cytology: T(2;3)59E2-3;80. genetics: Variegated for few. Breaks in 2R near c
origin: X ray induced. and bw and in 3R near cu.
discoverer: Slatis. *T(2;3)bwVD: Translocation(2;3) brown-Variegated
references: 1955, Genetics 40: 5-23. Dichaete linked
genetics: Associated with bwR14. origin: X ray induced,
discoverer: Oliver, 29k24.
cytology: T(2;3)59D;80C. references: 1932, Z. Induktive Abstammungs-
origin: X ray induced. Vererbungalehre 61: 447—88.
discoverer: Shi Us. genetics: Variegated for few. Homozygous lethal.
genetics: Associated with bwRls, 7T2;3)4wVD.3; Translocation(2;3) brown-
'3JfcwV*: Translocation(2;3) brown-Variegated Variegated of Demerec
origin: X ray induced. cytology: T(2;3)59D;81F. Also an inversion in 2R.
discoverer: Muller. origin: X ray induced,
references; Glass, 1933, J. Genet. 28: 69-112. discoverer Demerec, 33jl4.
1934, Am. Naturalist 68: 107-14. genetics: Variegates for few and mi but not abb.
genetics: Variegated for bw. Break near bw in 2R Mutant for Dfd. Homozygous lethal. Gives trans-
and in 3L just left of centromere, vection effects with certain pairs of bithorax pseu-
ofrwf information: Similar to T(2;3)bwV* and doalleles (Lewis, 1955, Am. Naturalist 89: 73-89).
T(2?3)hwvs; translocatioo parts interchangeable
among these rearrangements without altering pbe- cytology. T(2;3)59D2-4;80 (Schultz).
notype. origin: X ray induced,
discoverer: Demerec, 33k22.
origin: X ray induced, genetics: Variegates for bw and mi. Homozygous
discoverer: Patterson. lethal.
T(2;3)C discoverer: Roberts, 1964.

origin: X ray induced. genetics: Homozygous viable. Recombination re-
discoverer: Dobzhansky, 28h. duced in 3R.
references: 1929, Biol. Zentr. 49: 408-19. T(2;3)C49
Dobzhansky and Sturtevant, 1931, Carnegie Inst. cytology: T(2;3)22C-D;86E.
Wash. Publ. No. 421: 29-59. origin: X ray induced,
genetics: Break near centromere in chromosomes 2 discoverer: Roberts, 1964.
and 3. New order is 2L + 3L; 2R + 3R. Homozy- genetics: Homozygous viable. Recombination re-
gous lethal. duced in 2L.
T(2;3)C4: Translocatipn(2;3) Crossover T(2;3)CS8
suppressor cytology: T(2;3)40-41;96F; position of breakpoint in
cytology: T(2;3)40-41;94A; position of breakpoint in chromosome 2 with respect to centromere not deter-
chromosome 2 with respect to centromere not determined.
mined. origin: X ray induced.
discoverer: Roberts, 1964. genetics: Homozygous lethal. Recombination re-
genetics: Homozygous lethal. Recombination reduced in 3R.
duced in 3R. T(2;3)C6S
T(2;3)C11 cytology: T(2;3)40-41;75A;80-8l + In(3L)64C;77A;
cytology: T(2;3)40-41;64D;77A; position of break- involvement of chromosome 2 inferred from genetic
point in chromosome 2 with respect to centromere data; positions of heterochromatic breakpoints with
not determined. respect to centromeres not determined,
new order: 21 - 40|77A - 64D|77A - 100; new order: 21 - 4Q|80 - 100;
60 - 40|64D - 61. 60 - 40|75A - 64C|77A - 80|75A -
Tentative. 77A|64C - 61.
origin: X ray induced, Tentative.
genetics: Homozygous viable. Recombination re- discoverer: Roberts and D. Stewart, 1964.
duced in 3L. genetics: Homozygous lethal. Recombination re-
T(2;3)C16 duced in 3L..
cytology: T(2;3)50E;66C;70C. T(2;3)Ct01
new order: 21 - 50E|70C - 66C|50E - 60; cytology: T(2;3)29B;80-81; position of breakpoint in
61 - 66C|70C - 100. chromosome 3 with respect to centromere not deter-
origin: X ray induced, mined.
genetics: Homozygous lethal. Recombination re- discoverer: Roberts, 1965.
duced in 3L and 2R. genetics: Homozygote survives infrequently. Re-
T(2;3)C17 combination reduced in 2L.
cytology: T(2;3)56F;67E. T(2;3)CU1
origin: X ray induced, cytology: T(2;3)40-41;70F + In(3L)62B;79D-E; posi-
discoverer: Roberts, 1964. tion of breakpoint in chromosome 2 with respect to
genetics: Homozygous lethal. Recombination re- centromere not determined; new order therefore am-
duced in 3L and 2R. biguous .
T(2;3)C18 new order: 21 - 40|70F - 62B|79E - 100;
cytology: T(2;3)25B;40;84B. 60 - 4Q|70F - 79D|62B - 61.
new order: 21 — 25B|40 — 60; Tentative.
61 - 84B|25B - 40|84B - 100. origin: X ray induced.
genetics: Homozygous lethal. Recombination reduced in 3L.
duced in 2L. T(2;3)CJ22
T(2;3)C24 cytology: T(2;3)60B;80-81; position of breakpoint in
cytology: T(2;3)S3B;80-81; position of breakpoint in chromosome 3 with respect to centromere not deter-
chromosome 3 with respect to centromere not determined.
mined. origin: X ray induced.
discoverer: Roberts, 1964. genetics: Homozygous viable. Recombination re-
genetics: Homozygous viable. Recombination reduced in 2R.
duced in 2R. T(2;3)CU4
T(2;3)C29 cytology: T(2;3)34D;75F.
cytology: T(2;3)43F;92D. origin: X ray induced.
genetics: Homozygous lethal. Recombination re- T(2;3)C218

duced in 2L. cytology: T(2;3)40-41;70F; position of breakpoint in
T(2;3)C132 chromosome 2 with respect to centromere not deter-
cytology: T(2;3)55E;80-81; position of breakpoint in mined.
chromosome 3 with respect to centromere not deter- origin: X ray induced.
mined. discoverer: Roberts, 1965.
origin: X ray induced. genetics: Homozygous viable. Recombination re-
discoverer: Roberts, 1965. duced in 3L.
duced in 2R. cytology: T(2;3)35D;61A.
T(2;3)CJ49 origin: X ray induced.
cytology: T(2;3)52A;93B. discoverer: Roberts, 1965.
origin: X ray induced. genetics: Homozygous lethal. Recombination re-
discoverer: Roberts, 1965. duced in 3L.
duced in 2R and 3R. cytology: T(2;3)50D;62B + In(2LR)35C-D,52A-B.
T(2;3)C157 new order: 21 - 35C|52B - 50D|62B - 100;
cyto Iogy: T(2;3)41;96D-E + In(2LR)24F; 54F. 60 - 52A|35D - 50D|62B - 61.
new order: 21 - 24F|54F - 4l|96D - 61; origin: X ray induced.
60 - 54F|24F - 41 |96E - 100. discoverer: Roberts, 1965.
origin: X ray induced. genetics: Homozygous lethal. Recombination re-
discoverer: Roberts, 1965. duced in 2R and 3L.
genetics: Homozygous viable. Recombination re- T(2;3)C248
duced in 2L and 3R. cytology: T(2;3)S2C;94D;96B.
new order 60 - 52C|94D - 61;
T(2;3)C164 21 - 52CJ96B - 94D|96B - 100.
cytology: T(2;3)32F;64B. origin: X ray induced.
genetics: Homozygous lethal. Recombination reduced in 2R and 3R.
duced in 2L and 3L. T(2;3)C257
T(2;3)C177 cytology: T(2;3)50F;80; position of breakpoint in
cytology: T(2;3)40-41;62F + T(2;3)56F;79B; posi- chromosome 3 with respect to centromere not deter-
tion of left breakpoint in chromosome 2 with re- mined.
spect to centromere not determined. origin: X ray induced.
new order: 21 - 40|62F - 79B|56F - 4l|62F - 61; discoverer Roberts, 1965.
6 0 - 56F|79B - 100. genetics: Homozygous lethal. Recombination re-
origin: X ray induced. duced in 2R.
discoverer: Roberts, 1965. T(2;3)C259
genetics: Homozygous lethal. Recombination re- cytology: T(2;3)40-41;61E;73A; position of break-
duced in 3L and 2R. point in chromosome 2 with respect to centromere
not determined.
T(2;3)C199 new order 21 - 40|61E - 73A|41 - 60;
cytology: T(2;3)41;93E. 61A - 61E|73A - 100.
origin: X ray induced. Tentative.
genetics: Homozygous lethal. Recombination re- discoverer Roberts, 1965.
duced in 3R. genetics: Homozygous lethal. Recombination re-
T(2;3)C2O2 duced in 3L.
cytology: T(2;3)S6D;89D. T(2;3)C267
origin: X ray induced. cytology: T(2;3)21D;63F;64E + In(3LR)74F;8&D.
discoverer: Roberts, 1965. new order 21A - 21D|64E - 74FJ8SD - 74F[88D -
genetics: Homozygous viable. Recombination re- 100;
duced in 2R. 60 - 21DJ63F - 64E|63F - 61.
T(2;3)C2ll origin: X ray induced.
cytology: T(2;3)40-41;70C; position of breakpoint in discoverer Roberts, 1965.
chromosome 2 with respect to cenfaromere not deter- genetics: Homozygous lethal. Recombination re-
mined. duced in 3L.
origin: X ray induced. T(2;3)C287
discoverer Roberts, 1965. cytology: T(2;3)54F;89F.
genetics: Homozygous lethal. Recombination re- origin: X ray induced,
duced in 3L. discoverer Roberts, 1965.
genetics: Homozygous lethal. Recombination re- genetics: Homozygous lethal. Recombination re-
duced in 2R. duced in 2L and 3R.
T(2;3)C293 T(2;3)C328
cytology: T(2;3)43A;67A;80-81; position of break- cytology: T(2;3)55C;58B;80-81; position of break-
point in chromosome 3 with respect to centromere point in chromosome 3 with respect to centromere
not determined. not determined.
new order: 21 - 43A|67A - 61; new order: 21 - 55C|58B - 60;
60 - 43A|80 - 67A|81 - 100. 61 - 80|(55C - 58B)[81 - 100.
discoverer Roberts, 1965. discoverer: Roberts, 1965.
genetics: Homozygous lethal. Recombination re- genetics: Homozygous viable. Recombination re-
duced in 3L. duced in 2R. The segregant Dp(2;3)C328 =
T(2;3)C304 Dp(2;3)55C;58B;80-81 survives but not the comple-
cytology: T(2;3)48A;83C;100B. mentary deficiency.
new order: 21 - 48A|l00B - 100F; T(2;3)C356
60 - 48A|83C - 100B|83C - 61. cytology: T(2;3)29F;80-81; position of breakpoint in
origin: X ray induced. chromosome 3 with respect to centromere not deter-
discoverer: Roberts, 1965. mined.
genetics: Homozygous lethal. Recombination re- origin: X ray induced.
duced in 3R. discoverer: Roberts, 1965.
T(2;3)C308 genetics: Homozygous viable. Recombination re-
cytology: T(2;3)40-41;84B;94D;99B. duced in 2L.
new order: 21 - 40|94D - 84B|94D - 99B|84B - 61; T(2;3)C591
60 - 40|99B - 100. cytology: T(2;3)28D;69D.
Tentative. origin: X ray induced in oocyte.
origin: X ray induced, discoverer: Roberts and Thomas, 1965.
discoverer: Roberts, 1965. references: Thomas and Roberts, 1966, Genetics 53:
genetics: Homozygous lethal. Recombination re- 855-62.
duced in 3R. genetics: Homozygous lethal. Recombination re-
T(2;3)C309 duced in 2L.
cytology: T(2;3)58D;68F.
origin: X ray induced, *T(2;3)C-K: Translocation(2;3) Curved of
discoverer: Roberts, 1965. Krivshenko
genetics: Homozygous lethal. Recombination re- cytology: T(2;3)52;76;81;86.
duced in 2R and 3L. new order: 21 - 52186 - 100;
60 - 52)81 - 76J81 - 86|?6 - 61.
T(2;3)C3T1 origin: X ray induced.
cytology: T(2;3)54C;64C. discoverer: Krivshenko, 5513.
origin: X ray induced, references: 1956, DIS 30: 74.
discoverer. Roberts, 1965. genetics: Associated with C-K. Homozygous lethal.
genetics: Homozygous lethal. Recombination re- *T(2;3)D
duced in 2R and 3L. origin: X ray induced,
T(2;3)C313 discoverer: Dobzhansky, 28h.
cytology: T(2;3)27B;80-81; position of breakpoint in references: 1929, Biol. Zentr. 49: 408—19.
chromosome 3 with respect to centromere not deter- Dobzhansky and Sturtevant, 1931, Carnegie Inst.
mined. Wash. Publ. No. 421: 29-59-
origin: X ray induced. genetics: Heterozygote short lived and frequently
discoverer: Roberts, 1965. sterile, especially in female. Wings mis-shapen
genetics: Homozygous lethal. Recombination re- and legs short.
duced in 2L. T(2;3)dp: Translocation(2;3) dumpy
T(2;3)C316 cytology.* T(2;3)34D;41A;47E;48A;80;81 +
cytology: T(2;3)25F;80-81; position of breakpoint in In(2)27Dl-2;32D;44C5-6;44F3-12 (Bridges); exis-
chromosome 3 with respect to centromere not deter- tence of break or breaks in chromosome 3 inferred
mined. from genetic data (Muller, 1942; Cooper, Zitnsaering,
origin: X ray induced. and Krivshenko, 1955, Proc. Natl. Acad. ScL U.S.
discoverer. Roberts, 1965. 41: 911-14),
genetics: Homozygous lethal. Recombination re- new order. 21 - 27D1J32D - 34D|41A -
duced in 2L. 44C5J44F3 - 44C6|27D2 - 32D|44F12 -
T(2;3)C3U 47E|(80 - 81)|48A - 60;
cytology: T(2;3)24D;97D. 61 - 8 Q | ( 3 4 D - 41A)|81 - 100.
origin: X ray induced. Extremely tentative; 47E — 48A unaccounted for.
discoverer: Roberts, 1965. origin: Reportedly spontaneous.
discoverer. Nichols-Skoog, 36el6. *T(2;3)hlO0.27l: Translocation(2;3) hairy

synonym: In(2LR)dp. cytology: T(2;3)41;66D14-E1.
references: Morgan, Bridges, and Schultz, 1937, origin: X ray induced.
Carnegie Inst. Wash. Year Book 36: 301. discoverer: Alexander.
Curry, 1939, DIS 12: 46. references: Ward and Alexander, 1957, Genetics 42:
Muller, 1942, DIS 16: 64. 42-54.
genetics: Mutant for dp (dp36°), fcOb3*5'). rl[T(2;3)dp/ genetics: Mutant for h.
Dt(2RjM-S2r*11 is rl], tuf[T(2;3)dp/tuf is extreme *T(2;3)HK: Translocation(2;3) Half of Krivshenko
tuf], and a lethal that is uncovered by Df(2R)Np. cytology: T(2;3)22A;61A.
Associated .with a dominant rough eye mutant. Ho- origin: X ray induced in female.
mozygous lethal. The chromosome 3 segregant discoverer: Krivshenko, 56114.
Dp(2;3)dp = Dp(2;3)34D;41A;80;81 (tentative) sur- references: 1959, DIS 33: 95.
vives in poorly fertile male and sterile female, both genetics: Only the 2LD3P element recovered from
of which have arched wings and low viability; du- the treated oocyte.
plicated for pr. The complementary Df(2LR)dp — T(2;3)Hm: Translocation(2;3) Haltere mimic
Df(2LR)34D;41A is lethal. cytology: Breakpoints unknown.
T(2;3)dpD: Translocation(2;3) dumpy-Dominant origin: X ray induced.
cytology: T(2;3)25A;95B-D (E. B. Lewis). discoverer: Slat is, 49b5.
origin: X ray induced. genetics: Associated with Hm.
discoverer: £. B. Lewis, 1962. T(2;3)Hn: Translocation(2;3) Henna
references: Del Campo, 1963, DIS 38: 32. cytology: T(2;3)53E-54A;77A;94F;96A (E. B.
genetics: Mutant for dp. Homozygous lethal. Lewis),
T(2;3)dp»i; Translocation(2;3) dumpy-warped new order. 21 - 53E|77A - 61;
origin: X ray induced. 60 - 54A|94F - 96A|77A - 94F|96A -
discoverer: Schalet, 1955. 100.
references: Carlson and Schalet, 1955, DIS 29: 71— Tentative.
72. origin: X ray induced,
Carlson, 1958, DIS 32: 117-18. discoverer: Van Atta, 30k.
genetics: Apparently variegated for dp. Homozygous references: 1932, Am. Naturalist 66: 93—95.
lethal. 1932, Genetics 17: 637-59.
T(2;3)Dp-S: Translocation(2;3) with Duplication genetics: Separable from Hn, which is associated
Star with Df(3L)Hn =Df(3L)66A,66B.
cytology: T(2;3)21D4-E1;81F superimposed on *T(2;3)lf"h Translocation(2;3) light-mottled
Dp(2;2)21D2-3;21E2-3. cytology: T(2;3)40B-F;63E-F.
new order: 21A - 21E2J21D3 - 21D4|81F - 61; origin: X ray induced.
60-21El|81F - 100. discoverer: Hessler, 1957.
origin: X ray induced in Dp(2;2)S, ast ast. references: 1958, Genetics 43: 395—403.
discoverer: E. B. Lewis. genetics: Variegated for It.
genetics*. Y-suppressible expression of ast. cytology: T(2;3)40B-F;67E.
T(2;3)DrL: Translocation(2;3) Drop of Lewis discoverer. Hessler, 1957.
cytology: T(2;3)44;89F-90A + Itt(3R)89C}95D-96Bl. references: 1958, Genetics 43: 395-403.
new order 21 - 44|89F - 89CJ96B1 - 100; genetics: Variegated for It.
60 - 44|90A - 95DJ89C - 61. *T(2;3)lt'nS
origin: X ray induced, cytology: T(2;3)40B-F;98C.
discoverer: E. B. Lewis. origin: X ray induced.
genetics: Mutant for Dr, which is probably inde- discoverer: Hessler, 1957.
pendent of rearrangement. references: 1958, Genetics 43: 395—403.
*T(2;3)E genetics: Variegated for It.
cytology: T(2;3)30B;67E (Schultz). *T(2;3)lt>n6
origin: Spontaneous. cytology: T(2;3)26E-F;40B-F;96E.
discoverer: Sturtevant, 1929. new order: 21 - 26E|40B - 2 6 F | 9 6 E - 61;
references: Dobzhansky and Sturtevant, 1931, 60 - 4 0 F | 9 6 E - 100.
Carnegie Inst. Wash. Publ. No. 421: 29-59. origin: X ray induced,
genetics: Homozygous lethal. discoverer Hessler, 1957.
TTZ'W"rf- Translocation(2;3) glass references: 1958, Genetics 43: 395-403.
origin: Gamma ray induced. genetics: Variegated for It.
discoverer: Ives, 63d29.
references: 1965, DIS 40: 35. cytology: T(2;3)40B-F;100F.
genetics: Mutant for 0. origin: X ray induced.
T(2;3)Gt: see T(2;3)Puor discoverer: Hessler, 1957.
references: 1958, Genetics 43: 395—403. *T(2;3)lt*>23

genetics: Variegated for It. cytology: T(2;3)40B-F;62F.
*T(2;3)lf»8 origin: X ray induced.
cytology: T(2;3)40B-F;92B. discoverer: Hessler, 1957.
discoverer: Hessler, 1957. genetics: Variegated for It.
references: 1958, Genetics 43: 395—403. *T(2;3)lt<»24
genetics: Variegated for It. cytology: T(2;3)40B-F;59F;75C.
*T(2;3)lt>nio new order. 21 - 40B|75C - 100;
cytology: T(2;3)40B-F;64E. 60 - 59F|40F - 59F|75C - 61.
discoverer: Hessler, 1957. discoverer: Hessler, 1957.
references: 1958, Genetics 43: 395—403. references: 1958, Genetics 43: 395-403.
genetics: Variegated for It. genetics: Variegated for It.
*T(2;3)ltn>" *T(2;3)lf»27
cytology: T(2;3)40B-F;96F. cytology: T(2;3)40B-F;88E-F.
*T(2;3)lt>»28
cytology: T(2;3)40B-F;64F. cytology: T(2;3)40B-F;97E.
discoverer: Hessler, 1957. discoverer. Hessler, 1957.
7(2:3)1^29
cytology: T(2;3)40B-F;95F. cytology: T(2;3)40B-F;99F.
discoverer. Hessler, 1957. discoverer. Hessler, 1957.
*T(2;3)lt>»30
cytology: T(2;3)40B-F;92E. cytology: T(2;3)40B-F;99C.
discoverer: Hessler, 1957. discoverer Hessler, 1957.
*T(2;3)lf»32
cytology: T(2;3)40B'F;95C-D. cytology: T(2;3)40B-F;97A.
*T(2;3)lt'°1*
cytology: T(2;3)40B-F;98A. cytology: T(2;3)*0B-F;6tB.
discoverer: Hessler, 1957. discoverer. Hessler, 1957.
*T(2;3)lt>»3S
cytology: T(2;3)40B~F;94B. cytology: TX2;3)40B-F;64C-
feneties: Variegated for U. genetics: Variegated for tt.
T(2;3)lt>»ioo
cytology: T(2;3)4QB-F;93D. cytology: T(2;3)40;97F.
discoverer: Hessler, 1957. references: Baker and Rein, 1962, Genetics 47:
iwfwomeo*: 1958* Geoetic* 43: 395—403. 1399-1407.
§ an ©tics: Variegated for It. genetics: Variegated for It. Homozygous lethal.
T(2;3)Mv™d: s e e T(2;3)MV genetics: Eye color variegated; more prominent in

T(2;3)Me: Translocation(2;3) Moir6 male.
cytology: T(2;3)48Cl-2;59D2-3;60F;80-81 (tenta- *T(2;3)Hu: Translocation(2;3) Nude
tive) + In(3LR)69E;9lC + In(3R)89B;97D super- cytology: T(2;3)24;36-37;39-40;73-74;75-76;77-78;81-
imposed on In(3L)63C;72El-2 (Whittinghill, 1937, 8 2;8 5-86; 89-90.
DIS 8: 83); breakpoint in chromosome 3 with re- origin: X ray induced.
spect to centromere not determined; new order discoverer Sutton, 41a27.
therefore ambiguous. genetics: Associated with Nu. Homozygous lethal.
new order: 21 -48Cl|S9D2 - 48C2|60F; T(2;3)pGr: S ee T(2;3)PuOr
61 - 63C|72E1 - 69E|91C - 97D|89B - T(2;3)P: Translocation(2;3) Pale
8l|59D3 - 60F|80 - 72E2|63C - cytology: T(2;3)58E3-F2;60D14-E2;96B5-Cl
69E|91C - 89B|97D - 100. (Morgan, Bridges, and Schultz, 1935, Carnegie
Tentative. Inst. Wash. Year Book 34: 286).
origin: X ray induced in In(3L)P, Me. new order: 21 - 58E3|60E2 - 60F;
discoverer: Muller, 1930. 61 - 96B5|60D14 - 58F2|96CI - 100.
references: Glass, 1933, J. Genet. 28: 104. origin: Spontaneous.
genetics: Mutant for sbd (sbd1), Dp(2;3)Me = discoverer: Bridges, 17J16.
Dp(2;3)59D2-3;60F;80-81 survives. references: 1919, Anat. Record 15: 357-58.
*T(2;3)Me2 1923, Anat. Record 24: 426-27.
origin: X ray induced. Bridges and Morgan, 1923, Carnegie Inst. Wash.
discoverer Moore, 1929. Publ. No. 327: 184-87.
references: Glass, 1933, J. Genet. 28: 69-112. Li, 1927, Genetics 12: 1-58.
genetics: Break in 2L near centromere. Mutant for Kossikov and Muller, 1935, J. Heredity 26: 305-
Me. 17.
*T(2;3)MeS°: Translocation(2;3) Moire of Sytko Bridges, 1937, Cytologia (Tokyo), Fujii Jub. Vol.
discoverer: Sytko. 2: 745-55.
references: Agol, 1936, DIS 5: 7. genetics: Associated with P. Homozygote ordinarily
genetics: Breaks reportedly in 2R and 3R, yet mu- lethal but survives in presence of bw+Y =
tant for Me in 3L. Dp(2;Y)YL;58Fl-59A2;60E3-Fl; lethality therefore
T(2;3)Met: Translocation(2;3) Metatarsi irregular associated with 60D14-E2 breakpoint (Muller, 1942,
origin: X ray induced. DIS 16: 64). Dp(2;3)P = Dp(2;3)58E3-F2;60D14-
discoverer: Jonsson, 56al0. E2;96B5-C1 is viable and fertile; duplicated for
references: 1956, DIS 30: 73. loci of px, M(2)t, crs, bw, mi, abb, pd, 11, 1(2)NS,
genetics: Associated with Met, sp, bs, and ba but not a or M(2)c. Homozygous
*T(2;3)M0 Dp(2;3)P is lethal unless one chromosome 2 is
origin: Spontaneous. Df(2R)P = Df(2R)58E3-4;60D14-E2. Df(2R)P with
discoverer. Imaizumi, 59a. two normal third chromosomes is lethal.
references: 1962, Cytologia 27: 212—28. other information: First translocation recorded in
genetics: Breaks between en and vg in 2R and be- Drosophila melanogaster.
tween st and ss in 3R.
T(2;3)P23: Translocation(2;3) from Pasadena
T(2;3)Mot-K: Translocation(2;3) Mottled of cytology: T(2;3)81F;56F.
Krivshenko origin: X ray induced in Ubx e4.
cytology: T(2;3)41;6QD;80-8t; breakpoint in chromo- discoverer: E. B. Lewis, 49k.
some 3 with respect to centromere not determined; references: 1963, Am. Zoologist 3: 33—56.
association of arms therefore ambiguous. genetics: Gives transvection effect in T(2;3)P23,
new order 21 - 4l|80 - 61; Ubx/bx34* heterozygote.
60F - 60D|41 - 60D|80 - 100. T(2;3)P32
Tentative. cytology: T(2;3)42D-E,89D7-E1 + D((2)41A;44C-D.
origin: X ray induced. new order: 21 - 41A|44D - 60;
discoverer Krivshenko, 54c25. 61 - 89D7|42D - 41Ai44C - 42E|89E1 -
references: 1954, DIS 28: 75. 100.
1955, DIS 29: 76. origin: X ray induced in bx^4e.
genetics: Associated with Mot-K. Homozygous discoverer: E. B. Lewis, 50L
lethal. genetics: Gives transvection effect in T(2;3)P32,
*T(2;3}MV: Translocation(2;3) Variegated of bx34e/ZJbx heterozygote. The segregant
Mickey Dp(2;3)P32 = Dp(2;3)41A;42D-E;44C-D;89D7-El
cytology: T(2;3)43E;75C. survives and is fertile and virtually wild type; du-
origin: Gamma ray induced. plicated for stw, ap, tuf, and en but not pr or ltd.
discoverer: Mickey, 54d. *T(2;3)Pu: Translocation(2;3) Punch
synonym: T(2;3WVS4d. cytology: T(2;3)40F-41Al;70D~E + T(2;3)57B5~
references: 1963, DIS 38: 30. C1;79F.
new order: 21 - 40F|70E - 79F|57C1 - 60; 7(2,3)$*: Translocation(2;3) Star of Muller

61 - 7OD|41A1 - 57B5|79F - 100. cytology: T(2;3)21E2-3;79D2-E1 superimposed on
Tentative. In(2L)22Dl-2;33F5-34Al + In(2R)42A2-3;58A4-Bl.
origin: X ray induced, new order: 21A - 21E2J79E1 - 100F;
discoverer: Oliver, 28k4. 60F - 58B1 |42A3 - 58A4|42A2 -
references: Muller, 1930, J. Genet. 22: 326. 34Al|22D2 - 33F5|22D1 - 21E3|79D2 -
Oliver, 1932, Z. Induktive Abstammungs- 61A-
Vererbungslehre 61: 484. origin: X ray induced in In(2L)Cy +In(2R)Cy.
genetics: Associated with Pu. Homozygous lethal. discoverer Muller, 1928.
references: Painter and Muller, 1929, J. Heredity
T(2;3)PuGr: Translocation(2;3) Punch-Grape 20: 287-98.
cytology: T(2;3)57C;81F (Lewis, 1956, DIS 30: Muller, 1930, J. Genet. 22: 335-57.
130). Morgan, Bridges, and Schultz, 1936, Carnegie Inst.
origin: X ray induced, Wash. Year Book 35: 293.
discoverer Muller, 291. genetics: Mutant for S; also carries Cy.
synonym: T(2;3)Gr; Translocation(2;3) Grape;
T(2;3)pOr: Translocation(2;3) pink-Grape, *T(2;3)Sa: Translocation(2;3) Salmon
references: Glass, 1933, J. Genet. 28: 69—112. origin: X ray induced.
1934, Am. Naturalist 68: 107-114. discoverer: Van Atta, 30kl.
genetics: Mutant for Pu. Homozygous lethal. references: 1932, Am. Naturalist 66: 93—95.
1932, Genetics 17: 637-59.
*T(2;3)Pur*: Translocation(2;3) Punch-reversed genetics: Associated with Sa. Homozygous lethal.
cytology: T(2;3)40F-41A;70D-E + T(2;3)57B5- Break in 2L between pr and centromere and in 3L
C1;79F. near centromere.
new order 21 - 4 0 F | 7 0 E - 79F|57B5 - 41Al|70D - T(2;3)SbV: Translocation(2;3) Stubble-Variegated
61; cytology. T(2;3)41A-C;88;89B superimposed on
60 - 57Cl|79F - 100. In(3R)93D7-El;98F2-6. In(3LR)65;83 induced si-
Tentative, multaneously but was separated from it by recombi-
origin: X-ray-induced derivative of T(2;3)Pu = nation.
T(2;3)40F-41A;70D-E + T(2;3)57B5-C1;79F. new order. 21 - 41A|89B - 93D7|98F2 -
discoverer: Oliver, 32127. 9 3 E l | 9 8 F 6 - 100;
references: 1939, Genetics 24: 82. 61A - 88|89B - 88|41C - 60.
1941, Proc. Intern. Congr. Genet., 7th. p. 228. origin: X ray induced in In(3R)Mo, Sb sr.
genetics: Partial reversal of Pu. Homozygous discoverer E. B. Lewis, 1948.
lethal. references: 1956, DIS 30: 76-77.
T(2;3)Pu*f: Translocation(2;3) Punch-Wine genetics: Variegates for phenotype of deficiency for
cytology: T(2;3)57B-C;80. Sb, which is normal.
discoverer: E. B. Lewis, 55h. T(2;3)sbdlO6: Translocation(2;3) stubbloid
genetics: Mutant for Pu. cytology: T(2;3)22E;89B.
T(2;3)rn: Translocation(2;3) rotund origin: X ray induced.
origin: Probably X ray induced. discoverer: E. B. Lewis.
discoverer: Glass, 1929. *T(2;3)$car: Translocation(2;3) Scarred
references: 1934, DIS 2: 8. cytology: T(2;3)27E;95A + In(3)91F;96A,
Muller, 1953, DIS 27: 106-7. new order 21 - 27EJ95A - 96A|91F - 61;
Carlson, 1956, DIS 30: 109. 60 - 27EJ9SA - 91FJ96A - 100.
genetics: Mutant for rn. Homozygous viable but origin: X ray induced,
sterile in both sexes. Breakpoints near the centro- discoverer Yu, 48h.
meres and probably in right arms of chromosomes 2 references: 1949, DIS 23: 65.
and 3 (Carlson, 1956). About 10 percent of the genetics: Associated with Scar.
progeny of parents heterozygous for T(2;3)tn and
chromosome 2 inversions are nondisjunctional for *T(2;3)SM2: TranslocotioM2;3) Second Multiple
chromosome 2 (Muller, 1953). cytology- T(2;3)21A;40F;80-81 superimposed on
*T(2;3)SL: Translocation(2;3) Star of Lewis In(2L)22Dl-2;33F5-34AI + ln(2LR)22A3-Bl?6QB-C +
cytology. T(2;3)2tE2-3,81 F;88D6-8. In(2R)42A2'3;58A4-Bl; position of breaks in prox-
new order: 21 - 21E2J81F - 88D6J81F - 61; imal heterochroMitin with respect to centromeres
6 0 - 21E3|88D8 - 100. not determined.
Tentative. origin: X ray induced in In(2LR)SMl,
origin: X ray induced, discoverer R. F. Grell, 1953.
discoverer E. B. Lewis, 1940. references: Lewis and liislove, 19S3, DIS 27: 58.
references: 1945, Genetics 30: 137-66. Mis love and Lewis, 1954, DIS 28: 77.
genetics: Mutant for S. genetics: Variegated for It.
other information: Discarded because the T(2;3) im- discoverer: Slat is.
pairs its general usefulness as a chromosome 2 references: 1955, Genetics 40: 5—23.
balancer, described as SM2 in the section on bal- genetics: Associated with bwR5«t
ancers. T(2;3;4)bwV30ki8; Translocation(2;3;4) brown-
*T(2;3)sr4-2: Translocation(2;3) stripe Variegated
cytology: T(2;3)30C;90C-96. origin: X ray induced.
origin: X ray induced. discoverer: Van Atta, 3Okl3.
discoverer: Alexander. references: 1932, Genetics 17: 637-59.
references: I960, Genetics 45: 1019-22. genetics: Variegated for bw. Produces aneuploids
genetics: Mutant for sr. Homozygous lethal. that have Minute bristles.
*T(2;3)sri00.3i2
cytology: T(2;3)40-41;90D2-El. T(2;4)a
origin: X ray induced. cytology: T(2;4)50B2-3;102E (E. B. Lewis).
discoverer: Alexander. origin: X ray induced.
references: Ward and Alexander, 1957, Genetics 42: discoverer: Dobzhansky, 1929.
42-54. references: 1930, Biol. Zentr. 50: 671-85.
genetics: Mutant for sr. Homozygous lethal. 1931, Genetics 16: 629-58.
*T(2;3)sfl00.359: Translacation(2;3) scarlet genetics: Homozygous lethal. Fly hyperploid for
cytology: T(2;3)21C3-5;73A2-3;98F2-4. 2RD4P element survives rarely and is sterile.
new order: 21A - 21C3|73A3 - 98F2|73A2 - 61;
6 0 - 21C5|98F4 - 100. *T(2;4)A6: Translocation(2;4) from Austin
origin: X ray induced. cytology: T(2;4)57F2-3; breakpoint in chromosome 4
discoverer: Alexander. not determined.
references: Ward and Alexander, 1957, Genetics 42: origin: X ray induced.
42-54. discoverer: Patterson, Stone, Bedichek, and Suche.
genetics: Mutant for st. Homozygous lethal. references: 1934, Am. Naturalist 68: 359-69.
T(2;3)Xd: see T(2;3)apXa Patterson, Brown, and Stone, 1940, Texas Univ.
Publ. 4032: 167-89.
*T(Z-3;4)+3 genetics: Homozygous viable but sterile. Fly hyper-
cytology: T(2;3)2W;74F + T(3;4)67C;101B + ploid for the 4P2RD element viable and fertile.
T(3;4)95D-E;97C;101E. *T(2;4)A8
new order: 21A - 21D|74F - 95D|97C - 100; cytology: T(2;4)26F4-27A1; breakpoint in chromo-
60 - 21D|74F - 67CJ101E - 102F; some 4 not determined.
61 - 67C|l01E - 101A; origin: X ray induced.
101A - 101E|95E - 97C|l01E - 102F. discoverer: Patterson, Stone, Bedichek, and Suche.
This new order postulates involvement of two references: 1934, Am. Naturalist 68: 359-69.
fourth chromosomes, but the true origin of the cen- Patterson, Brown, and Stone, Texas Univ. Publ.
tromere to which 95E-97C is attached is unknown. 4032: 167-89.
In larval ganglia 1 metaphases, this element is not genetics: Homozygous viable and fertile. Fly hyper-
visible. ploid for the 2LD4P element viable and fertile.
origin: X ray induced. *T(2;4)A23
discoverer: Stern, Schaffer, and Heidenthal. cytology: T(2;4)58F; breakpoint in chromosome 4
synonym: JR3(+). not determined.
references: 1946, Proc. Natl. Acad. Sci. U*S. 32: origin: X ray induced.
26-33. discoverer: Patterson, Stone, Bedichek, and Suche.
Stern, MacKnight, and Kodani, 1946, Genetics 31: references: 1934, Am. Naturalist 68: 359-69.
598-619. genetics: Homozygous viable but sterile.
Kodani and Stern, 1946, Science 104: 620-21 *T(2;4)A27
(«g.). cytology: T(2;4)4OD1-F1; breakpoint in chromosome
genetics: Variegates for ci. Homozygous lethal. 4 not determined.
T(2;3;4)+3/ci has greater interruption of wing ori gin: X ray induced.
veins than ci/ci. T(2;3;4)+3/M(4) is normal, sup- discoverer: Patterson, Stone, Bedichek, and Suche.
porting the postulated involvement of two fourth references: 1934, Am. Naturalist 68: 359-69.
chromosome's. Burdette, 1940, Texas Univ. Publ. 4032: 157-63.
*T(2;3;4)hw*S8; Tmnslocaiion(2;3;4) brown- Patterson, Brown, and Stone, 1940, Texas Univ.
Rearranged Publ. 4032: 157-63.
cytology: T(2;3;4)59D;6S;1QIC. genetics: Homozygous lethal.
new order: 21 - 59O|65 - 61; *T(2;4)A29
60 - 59DJ101C - 102; cytology: T(2;4)47A4-5; breakpoint in chromosome 4
?|65 - 100. not determined.
101A to C lost. origin: X ray induced,
origin: X ray Induced.. discoverer. Patterson, Stone, Bedichek, and Such®.
references: 1934, Am. Naturalist 68: 359—69. fertile according to Bridges and Brehtne (1944, The
Patterson, Brown, and Stone, 1940, Texas Univ. Mutants of Drosophila melano&aster, Carnegie Inst.
Publ. 4032: 167-89. Wash. Publ. No. 552: 202).
genetics: Homozygous lethal. *T(2;4)AS2
*T(2;4)A30 cytology: T(2;4)36B; breakpoint in chromosome 4
cytology: T(2;4)53B2-C1; breakpoint in chromosome not determined,
4 not determined, origin: X ray induced.
origin: X ray induced. discoverer: Patterson, Stone, Bedichek, and Suche.
discoverer: Patterson, Stone, Bedichek, and Suche. references: 1934, Am. Naturalist 68: 359—69.
references: 1934, Am. Naturalist 68: 359—69. genetics: Homozygous viable and fertile.
Burdette, 1940, Texas Univ. Publ. 4032: 157-63. T(2;4)A53
Patterson, Brown, and Stone, 1940, Texas Univ. cytology: T(2;4)36El-3; breakpoint in chromosome 4
Publ. 4032: 167-89. not determined.
genetics: Homozygous viable and fertile. origin: X ray induced.
*T(2;4)A34 discoverer: Patterson, Stone, Bedichek, and Suche.
cytology: T(2;4)56F6-7; breakpoint in chromosome 4 references: 1934, Am. Naturalist 68: 359-69.
not determined. Burdette, 1940, Texas Univ. Publ. 4032: 157-63.
origin: X ray induced. Patterson, Brown, and Stone, 1940, Texas Univ.
discoverer: Patterson, Stone, Bedichek, and Suche. Publ. 4032: 167-89.
references: 1934, Am. Naturalist 68: 359—69. genetics: Homozygous viable and fertile.
Burdette, 1940, Texas Univ. Publ. 4032: 157-63. T(2;4)astv: Translocation(2;4) asteroid-variegated
Patterson, Brown, and Stone, 1940, Texas Univ. cytology: T(2;4)21E2-3;101.
Publ. 4032: 167-89. origin: X ray induced.
genetics: Homozygous viable. Either acts as or discoverer. E. B. Lewis, 1940.
carries a dominant suppressor of Pu (Oliver, 1943, references: 1945, Genetics 30: 137—166.
Anat. Record 87: 461). genetics: Variegates for S, ast, and ci. Homozygous
*T(2;4)A35 lethal. Fly with 4D2P element in place of one
cytology: T(2;4)26E; breakpoint in chromosome 4 chromosome 2 survives and has extremely rough
not determined, eyes. 4D2P is deficient for l(2)gl and net and pre-
origin: X ray induced. sumably for al, ex, and ds. Fly hyperploid for com-
discoverer: Patterson, Stone, Bedichek, and Suche. plementary 2LD4P also survives.
references: 1934, Am. Naturalist 68: 359—69. T(2;4)b
genetics: Homozygous viable. cytology: T(2;4)25E;102C15-Dl (Schultz and E. B.
*T(2;4)A40 Lewis). Metaphase chromosome 4 twice normal
cytology: T(2;4)49F3-50Al; breakpoint in chromo- size.
some 4 not determined. origin: X ray induced.
origin: X ray induced. discoverer. Dobzhansky, 1929.
discoverer: Patterson, Stone, Bedichek, and Suche. references: 1930, Biol. Zentr. 50: 671—85.
references: 1934, Am. Naturalist 68: 359—69. 1931, Genetics 16: 629-58.
Burdette, 1940, Texas Univ. Publ. 4032: 157-63. genetics: ci not affected. Homozygous viable and
Patterson, Brown, and Stone, 1940, Texas Univ. fertile. Fly hyperploid for 2LD4P element sur-
Publ. 4032: 167-89. vives; short and thick with flattened abdomen,
genetics: Homozygous viable and fertile. bulging eyes, and curved wings; both sexes sterile.
*T(2;4)A43 Duplicated for M(2)z and dp but not cl, ey, or av
cytology: T(2;4)22C; breakpoint in chromosome 4 (Morgan, 1946, DIS 20: 88).
not determined, *T(2;4)bwK2S; Translocation(2;4) brown-
origin: X ray induced. Rearranged
discoverer Patterson, Stone, Bedichek, and Suche. cytology: T(2;4)59D;101E.
references: 1934, Am. Naturalist 68: 359—69. origin: X-ray-induced derivative of bw.
Patterson, Brown, and Stone, 1940, Texas Univ. discoverer Slatis.
Publ. 4032: 167-89. references: 1955, Genetics 40: 5—23.
genetics: Homozygous viable and fertile. 2LD4P genetics: Associated with bw&2S.
element not recoverable in hyperploid; therefore *T(2;4)c
trans location probably more complex than given. cytology: Metaphase chromosome 4 about twice
*T(2;4)A45 normal size.
cytology: T(2;4)36D; breakpoint in chromosome 4 origin: X ray induced.
not determined, discoverer: Dobzhansky, 1929.
origin: X ray induced. references: 1930, Biol. Zentr. 50: 671—85.
discoverer: Patterson, Stone, Bedichek, and Suche. 1961, Genetics 16: 629-58.
references: 1934, Am. Naturalist 68: 359—69. genetics: Hotnozygote nearly lethal; wings do not
genetics: Homozygous lethal according to Patterson, expand, and fly dies early. Break in 2L between
Stone, Bedichek, and Suche (1934); viable and dp and b, but close to dp. Male hyperploid for
2LD4P element poorly viable and sterile. No varie- references: 1929, Biol. Zentr. 49: 408-19.
gation for ci + (Stern). 1929, Proc. Natl. Acad. Sci. U.S. 15: 633-38.
T(2;4)d 1930, Genetics 15: 347-99.
cytology: T(2;4)55E-F (Lewis, 1956, DIS 30: 130); genetics: Homozygous lethal. Break in 3L between
breakpoint in chromosome 4 not determined. D and th.
origin: X ray induced. *T(3;4)A1: Translocation(3;4) from Austin
discoverer: Dobzhansky, 1929. cytology: T(3;4)89A6-Bl;102B; breakpoint in chro-
references: 1930, Biol. Zentr. 50: 671-85. mosome 4 inferred from Painter's fig. 40 (1935).
1931, Genetics 16: 629-58. origin: X ray induced.
genetics: Homozygote nearly lethal; fly is short discoverer: Patterson, Stone, Bedichek, and Suche.
lived and has inflated wings. No viable aneuploid references: 1934, Am. Naturalist 68: 359—69.
product. Painter, 1935, Genetics 20: 301-26 (fig.).
genetics: Homozygous viable and fertile according
T(3;4)85C to Patterson, Stone, Bedichek, and Suche (1934);
cytology: T(3;4)85C; breakpoint in chromosome 4 homozygous lethal according to Bridges and
not determined. Brehme (1944, The Mutants of Drosophila melano-
discoverer: E. B. Lewis. gaster, Carnegie Inst. Wash. Publ. No. 552: 203).
references: Pipkin, 1959, Texas Univ. Publ. 5914: T(3;4)A2
69-88. cytology: T(3;4)94A3-4;101F (Brown).
T(3;4)86D discoverer: Patterson, Stone, Bedichek, and Suche.
cytology: T(3;4)86D2-3;1Q1F. references: 1934, Am. Naturalist 68: 359—69.
origin: Neutron induced in bx34e e4. Painter, 1935, Genetics 20: 301-26 (fig.).
discoverer: E. B. Lewis. Brown, 1940, Texas Univ. Publ. 4032: 11-64.
references: Grell, 1959, Genetics 44: 421—35. genetics: Homozygous viable and fertile. Fly hyper-
1959, Genetics 44: 911-22. ploid for 3RD4P element survives.
genetics: Homozygous viable and fertile. *T(3;4)A3
T(3;4)86D/ci has ci effect; enhanced by low tem- origin: X ray induced.
perature; tends to be suppressed by extra Y chro- discoverer: Patterson, Stone, Bedichek, and Suche.
mosome. Venation of homozygote and haplo-4 is synonym: T(3;4)A60.
ci+. references: 1934, Am. Naturalist 68: 359—69.
T(3;4)88B Painter, 1935, Genetics 20: 301-26 (fig.).
cytology: T(3;4)88B; breakpoint in 4 not determined, genetics: Homozygous lethal. 3R broken between e
origin: X ray induced in Ubx. and ca.
discoverer E. B. Lewis. *T(3;4)A4
references: Grell, 1959, Genetics 44: 421—35. cytology: T(3;4)80-81;101.
genetics: Homozygous lethal. Has no position effect origin: X ray induced.
on d. discoverer: Patterson, Stone, Bedichek, and Suche.
T(3;4)89E Painter, 1935, Genetics 20: 301-26 (fig.).
cytology: T(3;4)89E2-3;101F. genetics: Homozygous lethal.
origin: X ray induced in ss bx Su(as)2. *T(3;4)A5
discoverer: E. B. Lewis. cytology: T(3;4)92A5-6; breakpoint in chromosome 4
references: Grell, 1959, Genetics 44: 911—22. not determined.
genetics: Associated with bxd101. Homozygous origin: X ray induced.
lethal. T(3;4)89E/ci has a ci effect; enhanced by discoverer: Patterson, Stone, Bedichek, and Suche.
low temperature; tends to be suppressed by extra Y references: 1934, Am. Naturalist 68: 359-69.
chromosome. Burdette, 1940, Texas Univ. Publ. 4032: 157-63.
T(3;4)104: see T(3;4)f genetics: Homozygous lethal.
*T(3;4)684 *T(3;4)A8
cytology: T(3;4)67;101; breakpoints roughly esti- cytology: T(3;4)75B4-5;102Dl-3 (Brown).
mated from fig. of Dubinin and Sidorov (1935). origin: X ray induced.
origin: X ray induced. discoverer Patterson, Stone, Bedichek, and Suche.
discoverer: Dubinin and Sidorov. references: 1934, Am. Naturalist 68: 359—69.
references: 1934, Biol. Zh. (Moscow) 3: 307—31. Painter, 1935, Genetics 20: 301-26 (fig.).
1935, Biol. Zh. (Moscow) 4: 555-68 (fig.). Brown, 1940, Texas Univ. Publ. 4032: 11-64.
genetics: Position effects on both h and ci. Burdette, 1940, Texas Univ. Publ. 4032: 157—63.
T(3;4)o genetics; Homozygous viable and fertile.
cytology: Metaphase chromosome 4 about one-half *T(3;4)A9
length of 3L. cytology: T(3;4)87E3-Fl;102F.
origin: X my induced. origin: X ray induced.
discoverer: Dobzhansky, 29h. discoverer: Patterson, Stone, Bedichek, and Suche.
references: 1934, Am. Naturalist 68: 359-69. *T(3;4)A24

Painter, 1935, Genetics 20: 301-26 (fig.)- cytology: T(3;4)99;102B-C; inferred from Painter's
Brown, 1940, Texas Univ. Publ. 4032: 11-64. fig. 48 (1935).
T(3;4)A12 discoverer: Patterson, Stone, Bedichek, and Suche.
cytology: T(3;4)73Cl-2;102C; breakpoint in chromo- references: 1934, Am. Naturalist 68: 359—69.
some 4 roughly estimated from Painter's fig. 15 Painter, 1935, Genetics 20: 301—26 (figO-
(1935). genetics: Homozygous viable and fertile.
discoverer: Patterson, "Stone, Bedichek, and Suche. cytology: T(3;4)82B3-Cl;101A-D.
references: 1934, Am. Naturalist 68: 359-69. origin: X ray induced.
Painter, 1935, Genetics 20: 301-26 (fig.). discoverer: Patterson, Stone, Bedichek, and Suche.
Burdette, 1940, Texas Univ. Publ. 4032: 157-63. references: 1934, Am. Naturalist 68: 359—69.
Pipkin, 1959, Texas Univ. Publ. 5914: 69-88. Painter, 1935, Genetics 20: 301-26 (fig.).
genetics: Homozygote poorly viable and fertile. Fly Brown, 1940, Texas Univ. Publ. 4032: 11-64.
hyperploid for 3LD4P element survives. Burdette, 1940, Texas Univ. Publ. 4032: 157—63.
T(3;4)A13 genetics: Homozygous viable and fertile.
cytology: T(3;4)67E3-4;102D-E; breakpoint in chro- T(3;4)A28
mosome 4 roughly estimated from Painter's fig. 14 cytology: T(3;4)94D3-4;102 (E. B. Lewis); break-
(1935). point in chromosome 4 estimated from Painter's
origin: X ray induced. fig. 44 (1935).
discoverer: Patterson, Stone, Bedichek, and Suche. origin: X ray induced.
references: 1934, Am. Naturalist 68: 359—69. discoverer: Patterson, Stone, Bedichek, and Suche.
Painter, 1935, Genetics 20: 301-26 (fig.). references: 1934, Am. Naturalist 68: 359—69.
Patterson, Brown, and Stone, 1940, Texas Univ. Painter, 1935, Genetics 20: 301-26 (fig.).
Publ. 4032: 167-89. Pipkin, 1959, Texas Univ. Publ. 5914: 69-88.
Pipkin, 1959, Texas Univ. Publ. 5914: 69-88. genetics: Homozygous viable and fertile.
genetics: Homozygous lethal. Fly hyperploid for *T(3;4)A3Q
3LD4P element survives. cytology: T(3;4)96E5-Fl;102B-C; breakpoint in chro-
some 4 roughly estimated from Painter's fig. 47
*T(3;4)A14 (1935).
cytology: T(3;4)80;101. origin: X ray induced.
origin: X ray induced. discoverer: Patterson, Stone, Bedichek, and Suche.
discoverer: Patterson, Stone, Bedichek, and Suche. references: 1934, Am. Naturalist 68: 359-69.
references: 1934, Am. Naturalist 68: 359—69. Painter, 1935, Genetics 20: 301-26 (fig.).
Painter, 1935, Genetics 20: 301-26 (fig.). Pipkin, 1959, Texas Univ. Publ. 5914: 69-88.
*T(3;4)A20 genetics: Homozygous lethal. Fly hyperploid for
cytology: T(3;4)89A; 101F; breakpoint in chromosome 3RD4P survives.
4 roughly estimated from Painter's fig. 41 (1935). *T(3;4)A31
origin: X ray induced. cytology: T(3;4)80;101.
discoverer: Patterson, Stone, Bedichek, and Suche. origin: X ray induced.
references: 1934, Am. Naturalist 68: 359—69. discoverer: Patterson, Stone, Bedichek, and Suche.
Painter, 1935, Genetics 20: 301-26 (fig.). references: 1934, Am. Naturalist 68: 359—69.
genetics: Homozygous lethal. Painter, 1935, Genetics 20: 301-26 (fig.).
Brown, 1940, Texas Univ. Publ. 4032: 11-64.
*T(3;4)A22 Burdette, 1940, Texas Univ. Publ. 4032: 157—63.
cytology: T(3;4)61E-F;102B-C; estimated from genetics: Homozygous viable and fertile.
Painter's fig. 11 (1935). *T(3;4)A34
origin: X ray induced. cytology: T(3;4)61F;101F; estimated from Painter's
discoverer Patterson, Stone, Bedichek, and Suche. fig. 10 (1935).
references: 1934, Am. Naturalist 68: 359—69. origin: X ray induced.
1935, Painter, Genetics 20: 301-26 (fig.), discoverer: Patterson, Stone, Bedichek, and Suche.
genetics: Homozygous lethal. 3L broken to the left references: 1934, Am. Naturalist 68: 359—69.
of tu. Painter, 1935, Genetics 20: 301-26 (fig.),
*T(3;4)A23 genetics: Homozygous lethal. 3L broken to left of
cytology: T(3;4)66D5-E1;1O1F.
discoverer: Patterson, Stone, Bedichek, and Suche. cytology: T(3;4)8OB3-C1;1Q2E (Brown). 3L broken
references: 1934, Am. Naturalist 68: 359—69. about one-sixth the distance from centromere to tip
Painter, 1935, Genetics 20: 301-26 (fig.). in roetaphase chromosome.
genetics: Hcsaoxygous lethal in male, viable in fe- origin: X ray induced.
male. discoverer. Patterson, Stone, Bedichek, and Suche.
references: 1934, Am. Naturalist 68: 359—69. T(3;4)A60: see T(.3;4)A3

Painter, 1935, Genetics 20: 301-26 (fig.). *T(3;4)b
Brown, 1940, Texas Univ. Publ. 4032: 11-64. cytology: Chromosome 4 increased to one-half the
Burdette, 1940, Texas Univ. Publ. 4032: 157-63. length of 3L in metaphase figures.
*T(3;4)A37 discoverer. Dobzhansky, 28h.
cytology: T(3;4)86E5-6;101F; breakpoint in chromo- references: 1929, Biol. Zentr. 49: 408-19.
some 4 roughly estimated from Painter's fig. 38 1929, Proc. Natl. Acad. Sci. U.S. 15: 633-38.
(1935). 1930, Genetics 15: 347-99.
origin: X ray induced. genetics: Breakpoint in 3L. near th. Crossing over
discoverer: Patterson, Stone, Bedichek, and Suche. markedly lowered near th and somewhat so at 3L
references: 1934, Am. Naturalist 68: 359-69. tip.
Painter, 1935, Genetics 20: 301-26 (fig.). T(3;4)c
genetics: Homozygous lethal. cytology: T(3,-4)86B-C;l01F (Lewis, 1951, DIS 25:
*T(3;4)A39 108-9).
cytology: T(3;4)94B4-C1;IO1F; breakpoint in chro- origin: X ray induced.
mosome 4 estimated from Painter's fig. 46 (1935). discoverer: Dobzhansky, 28h.
origin: X ray induced. references: 1929, Biol. Zentr. 49: 408-19.
discoverer: Patterson, Stone, Bedichek, and Suche. 1929, Proc. Natl. Acad. Sci. U.S. 15: 633-38.
references: 1934, Am. Naturalist 68: 359-69. 1930, Genetics 15: 347-99.
Painter, 1935, Genetics, 20: 301-26 (fig.). genetics: Homozygous viable and fertile, ci not
Burdette, 1940, Texas Univ. Publ. 4032: 157-63. affected. Crossing over much reduced near break-
genetics: Homozygous lethal. point in heterozygote and even more reduced in
*T(3;4)A43 homozygote in some regions (Beadle, 1932, Proc.
origin: X ray induced. Natl. Acad. Sci. U.S. 18: 160-65).
discoverer: Patterson, Stone, Bedichek, and Suche. *T(3;4)d
references: 1934, Am. Naturalist 68: 359—69. cytology: Metaphase figures show barely detectable
Painter, 1935, Genetics 20: 301-26 (fig.). increase in size of chromosome 4.
genetics: Homozygous lethal. 3R broken near sr. origin: X ray induced.
*T(3;4)A44 discoverer: Dobzhansky, 28h.
cytology: T(3;4)76;99;102D-F; estimated from references: 1929, Biol. Zentr. 49: 408—19.
Painter's figs. 49 and 51 (1935). 1929, Proc. Natl. Acad. Sci. U.S. 15: 633-38.
new order 61 - 76|lO2D - 101; 1930, Genetics 15: 347-99.
1 0 0 - 99|76 - 99|1O2F. genetics: Homozygous lethal, ci not affected
references: 1934, Am. Naturalist 68: 359—69. (Stern). Breakpoint in 3R between ca and M(3)g
origin: X ray induced, and in 4 to the left of M(4) and ey. Apparently,
discoverer: Patterson, Stone, Bedichek, and Suche. 4D3P element can substitute for a normal 3 pro-
Painter, 1935, Genetics 20: 301-26 (fig.), ducing Minute flies. Hyperploids for 3RD4P ele-
genetics: Homozygous lethal. ment probably also survive.
*T(3;4)A45 T(3;4)DI7P: Translocation(3;4) De/fo-7 of
cytology: T(3;4)80;101. Panshin
discoverer: Patterson, Stone, Bedichek, and Suche. discoverer: Panshin.
references: 1934, Am. Naturalist 68: 359-*69. references: 1935, Dolk. Akad. Nauk SSSR 4: 85-88.
Painter, 1935, Genetics 20: 301-26 (fig.)- genetics: Chromosome 3 broken to the right of cu.
phenorype: Homozygous viable but sterile. Mutant for Dl; position effect that weakens domi-
*T(3;4)A52 nance of c a + .
cytology: T(3;4)65D3-F2; breakpoint in chromosome T(3;4h
4 not determined, cytology: T(3;4)79E;102F (Lewis, 1956, DIS 30:
origin: X ray induced. 130),
discoverer. Patterson, Stone, Bedichek, and Sucbe. origin: X ray induced.
references: 1934, Am. Naturalist 68: 359—69. discoverer: Dobzhansky, 28h.
Painter, 1935, Genetics 20: 301-26 (fig.). references: 1929, Biol. Zentr. 49: 408-19.
ph©notype: Hcnsozygous viable and fertile. 1929, Proc. Natl. Ac®d. Sci. U.S. 15: 633-38.
*T(3;4)A56 1930, Genetics 15: 347-99.
cytology: T(3;4)76E2"F3;101F; breakpoint in chro- genetics: Homozygous semi lethal and female sterile.
mosome 4 estimated from Painter's fig. 17 (1935). ci not affected (Stern). Crossing over normal in
origin: X ray induced. betarocygote except oear p.
discoverer. Patterson, Stone, Bedichek, and Suche. T(3;4)f
references: 1934, Am. Naturalist 68: 356~§ Q . cytology: T(3;4)6SD; at least seven bands of chro-
Painter, 1935, Genetics 20: 301-26 (fig.). raomoaa© 4 Inserted into 3L (Lewis, 1956, DIS 30:
genetics: Hornozygous lethal. 130).
CHROMOSOME ABERRATIONS - TRANSPOSITIONS 403

discoverer: Sturtevant, 1930. discoverer: Bishop, 1939.
synonym: T(3;4)104. references: Sutton, 1943, Genetics 28: 99.
references: Beadle, 1933, Z. Induktive genetics: Male and homozygous female viable.
Abstammungs- Vererbungslehre 65: 111—28. Crossing over in region 3E3-15F9 in
genetics: Homozygous lethal. No ci variegation Tp(l)B263-*8/+ heterozygote yields
(Stern). Dp(l;l)B263-48 = Dp(l;l)3E2-3;15F9-16Al;20A2-3,
*T(3;4)H1: Translocation(3;4) from Howard which is heterozygous viable and produces the Bar
University effect. The complementary deficiency is heterozy-
cytology: T(3;4)80-81; breakpoint in chromosome 4 gous lethal.
not determined.
origin: X ray induced, *Tp(l)ct6ai: Transposition(l) cut
discoverer: Pipkin. cytology: Tp(l)7B2-Cl;19;20.
references: 1959, Texas Univ. Publ. 5914: 69—88. new order: 1 - 7B2|(19 - 2O)|7C1 - 19|20.
*T(3;4)H3 Nucleolus organizer included in transposed piece,
cytology: T(3;4)80-81; breakpoint in chromosome 4 origin: X ray induced.
not determined, discoverer: Hannah, 1947.
origin: X ray induced. references: 1949, Proc. Intern. Congr. Genet., 8th.
discoverer: Pipkin. pp. 588-89.
references: 1959, Texas Univ. Publ. 5914: 69-88. genetics: Variegated for ct. Male lethal.
*T(3;4)H5 Tp(l)hill: see ln(l)hill
cytology: T(3;4)96E; breakpoint in chromosome 4 Tp(l)l-272-13: see In(l)l-272~13
not determined.
origin: X ray induced. *TpO)N264-63: Transposition) Notch
discoverer: Pipkin. cytology: Tp(l)3C7-9;13C7-8;19F (Hoover),
references: 1959, Texas Univ. Publ. 5914: 69-88. new order: 1 - 3C7|(13C8 - 19F)|(3C9 -
genetics: Fly hyperploid for 3RD4P survives. 13C7)|l9F - 20.
*T(3;4)H6 origin: X ray induced,
cytology: T(3;4)98A; breakpoint in chromosome 4 discoverer: Demerec, 38e.
not determined. genetics: Mutant for N but not for w, rst, or dm.
origin: X ray induced. s: S ee
discoverer: Pipkin.
references: 1959, Texas Univ. Publ. 5914: 69—88. Tp(3)bxd100: Transpasition(3) bithoraxoid
genetics: Fly hyperploid for 3RD4P survives. cytology: Tp(3)66C;89B5-6;89E2-3.
*T(3;4)H7 new order: 61 - 66C|(89B6 - 89E2)|66C -
cytology: T(3;4)66C; breakpoint in chromosome 4 89B5|89E3 - 100.
not determined, origin: X ray induced.
origin: X ray induced, discoverer: E. B. Lewis.
discoverer Pipkin. references: 1951, Cold Spring Harbor Symp. Quant.
references: 1959, Texas Univ. Publ. 5914: 69-88. Biol. 16: 159-74.
*T(3;4)K: Translocation(3;4) of Kirssanov genetics: Mutant for bxd but not bx, Dp(3;3)bxdl00 »
origin: X ray induced. Dp(3;3)66C;89B5-6;89E2-3 derived by crossing
discoverer: Kirssanov. over in region 66C-89B of Tp(3)bxd100/+ survives
references: 1933, Biol. Zh. (Moscow) 2: 447-50. and is duplicated for bx and the lethal effect of
T(3;4)l-18: Translocation(3;4) lethal Ubx but not for bxd. The complementary
origin: X ray induced, Df(3R)bxdl 00 « Dt(3R)89B5-6;89E2-3 survives in
discoverer: Gloor and Green, 1957. heterozygote and has Ubx phenotype.
genetics: Variegates for ci; mutant for 1(4)18.
T(3;4)P86: Translocation(3;4) from Pasadena
cytology: T(3;4)88B-C;101 (E. B. Lewis), cytology: Tp(3)89E2-3;91C7-Dl;92A2-3.
origin: X ray induced. new order. 61 - 89E2|(91D1 - 92A2)|89E3 -
discoverer: E. B. Lewis. 91C7|92A3 - 100.
TRANSPOSITIONS discoverer. E. B. Lewis.
genetics: Mutant for bxd but not bx or Dl.
Tp(l)303-l: see In(l)303-l Dt(3R)bxdllO ^ Di(3R)91C7-DJ;92A2-3, derived
Tp(l)At: see ln(!)At from crossing over in region 89E-91C in
Tp(l)B363-24'. see/nfl>i?2<sj-2 4 Tp(3)bxd*1 °/ + female, survives in h«terozygote
*Tp(l)B263~48: Trcnsposition(l) Bar and has Dl phenotype. The complementary
cytology: Tp(l)3E2-3;15F9-16Al;20A2-3. Dpf3;3)hxd*10 = Dp{3;3)B9E2~3;9lC7~Dl;92A2-3 is
new order. 1 - 3E2J16A1 - 2QA2J3E3 - duplicated far Dl and acts as a suppressor of Dl in
15F9J20A3 - 20F. Dp(3;3)hxdUG/Dl7 hetero*ygot« (E. B. Lewis).
Tp(3)C285: Transposition^) Crossover origin: X ray induced.

suppressor discoverer: Alexander.
cytology: Tp(3)88F;98B;99B. references: Ward and Alexander, 1957, Genetics 42:
new order: 61 - 88F|98B - 99B|88F - 98B|99B - 42-54.
100. genetics: Mutant for sr. Homozygous lethal.
discoverer. Roberts, 1965.
genetics: Homozygous lethal. Recombination be-
tween st and ca sharply reduced. Tp(3)Vno: Transposition^) Vein off
TP(3)C341 cytology: Tp(3)89E;93F;97A (Nicoletti and Lewis,
cytology: Tp(3)63C;71B;80-81 position of right 1960, DIS 34: 53).
breakpoint in chromosome 3 with respect to centro- new order: 61 - 89E|93F - 97A|89E - 93F|97A -
mere not determined. 100.
new order: 61 - 63C|71E - 80|{63C - 71E)|81 - origin: X ray induced.
100. discoverer: E. H. Grell, 56c.
origin: X ray induced. synonym: In(3R)Vno.
discoverer: Roberts, 1965. references: 1959, DIS 33: 94.
genetics: Homozygous viable. Recombination re- genetics: Associated with Vno. Homozygous lethal.
duced in 3L.
*Tp(3)sr300.240: Transposition^) stripe
cytology: Tp(3)75C;89E;92A. : seeR(l)l
new order: 61 - 75C|(89E - 92A)|75C - 89E|92A -
100. Inserted piece said to be in inverted order but : seeR(l)l
not specified whether with respect to numerical
sequence or centromere. : see R(l)2
SPECIAL CHROMOSOMES
Balancers
Compound Chromosomes
Multiply Marked Chromosomes
X—Y Combinations
Y Derivatives
BALANCERS complete: see FM1

CyO: Curly derivative of Oster
asc constitution: ln(2LR)O, dp / v J Cy pr en2.
sl L 8R si 8
constitution: In(l)sc sc +S, sc sc w°. synthesis: Oster.
properties: Like Base except that B reverted. synonym: Cy,InsO5.
Base references: 1956, DIS 30: 145.
constitution: In(l)scslLsc8R+S, scsi sc8 wa B. properties: More effective suppressor of crossing
synthesis: Muller. over in chromosome 2 than In(2L+2R)Cy; should be
synonym: M-5: Muller-5. superior to SMI as balancer for chromosome 2.
references: Spencer and Stern, 1948, Genetics 33: FMJ: First Multiple
43-74. constitution: In(l)sc8+dl-49, y31d sc8 wa lzs B.
properties: Male and homozygous female viable and synthesis: Schultz and Curry.
fertile; X/0 male poorly viable, variegated for y, synonym: complete.
ac, and probably 1(1)J1. Suppresses crossing over references: Lewis and Mislove, 1963, DIS 27: 57—
in X, but less so than Binsc because Jn(l)S — 58.
In(l)6Al-3;10F10-llAl less effective than properties: Male viable and fertile; homozygous fe-
In(l)dl-49 =ln(l)4D7-El;llF2-4. Routinely used in male viable but sterile because of lzs. Reduces
detection of sex-linked recessive lethals. crossing over in X chromosome. Useful for bal-
Binsc ancing sex-linked recessive sterile or lethal muta-
constitution: In(l)scSlLsc8R+dl-49, s c ^ sc« B. tions.
synthesis: Muller.
properties: Male and homozygous female viable and FM3
fertile. Suppresses crossing over in X chromo- constitution: In(l)FM3, y31d sc8 dm B.
some. synthesis: R. F. Grell, 1954.
Binscy references: Mislove and Lewis, 1954, DIS 28: 77.
constitution: In(l)scSiLsc8R+dl-49, y scs* sc8 v B. properties: Male lethal owing to presence of two
synthesis: Muller. recessive lethals in In(l)FM3, which may be
references: 1952, DIS 26: 113-14. covered by BsYy+ or by y+Y and BSY. Effectively
Muller and Oster, 1963. In Methodology in Basic suppresses crossing over in the X chromosome.
Genetics, W. J. Burdette, ed. Holden-Day, Inc. Useful for balancing sex-linked recessive female-
pp. 249-78. sterile mutants, and in combination with BsYy+ for
properties: Male and Homozygous female viable and balancing sex-linked recessive lethal and male-
fertile. Suppresses crossing over in X chromo- sterile mutants.
some.
Binsn FM4
constitution: In(l)scslLscaR+dl-49, constitution: In(l)FM4, y31d sc8 dm B.
2
sc$l sc& sn^ B. synthesis: R. F. Grell, 1954.
synthesis: Muller. references: Mislove and Lewis, 1954, DIS 28: 77.
properties: Male viable and fertile; homozygous fe- properties: Male viable and fertile; homozygous fe-
male viable but sterile because of snX2. Sup- male viable but sterile because of dm. In(l)FM4 is
presses crossing over in X chromosome. the consequence of the approximate reinversion of
Biny ln(l)dl-49 in In(l)sca+dl-49 and is similar in se-
constitution: In(l)4sc8LscSlR+dl-49, y31d sc- v f B. quence to ln(l)scs but with the insertion of 3C-4F
synthesis: Lindsley. into 11F. Unless this small transposition has
references: Lindsley and Edington, 1957, DIS 31: an abnormally large effect on crossing over
131-32. e.g., see Dp(2;2)C619, recombination might be
Lindsley, Edington, and Von Halle, 1960, Genetics expected to be frequent in FM4/+ heterozygotes
45: 1649-70. and practically normal in FM4/ln(l)sc8 heterozy-
properties: Male lethal because of deficiency for sc. gotes. In FM4/+ heterozygotes, double crossovers
Suppresses crossing over in the X chromosome. with points of exchange inside or outside the 3C to
Used in the recovery of F-suppressed sex-linked 11F region should produce euploid X chromosomes,
recessive lethals. and those with one point of exchange inside and
C(3)x one outside produce complementary duplications
constitution: Probably In(3L+3R)P. and deficiencies for 3C to 4F. The duplication
CIB survives in either sex and exhibits a Confluens
constitution: In(l)Cl, sc 1(1 )C t2 si B. phenotype (E. H. Grell); the deficiency might sur-
synthesis: Muller. vive in the heterozygote as a Notch Minute female
references: 1928, Genetics 13: 279-357. judging from the survival of the slightly smaller
properties: Male lethpl. Suppresses crossing over in Df(l)W*>258-*4 = Df(l)3C3~4;4D2-El. Balancing
X chromosome. Originally used in recovery of sex- properties not well determined. Some lines carry
+
linked recessive lethals; largely replaced by Base wS5t a n d in some y31d replaced with y or B with
for this purpose. / or +.
SPECIAL CHROMOSOMES - BALANCERS 407
FM6 properties: Male viable and fertile; homozygous fe-

constitution: In(l)FM6, y31d sc* dm B. male and X/0 male inviable because of deficiency
synthesis: R. F- Grell, 55i. for bb. Suppresses crossing over in X chromosome.
references: Grell and Lewis, 1956, DIS 30: 71. Used in the recovery of F-suppressed sex-linked
properties: Male viable and fertile, homozygous fe- recessive lethals.
male viable but sterile because of dm. Like FM4 SMI: Second Multiple
except for the presence of the additional In(l)15D- constitution: In(2LR)SMl) al2 Cy en2 sp2.
E;20A-B. Reservations similar to those about the synthesis: R. F. Grell, 1953.
balancing ability of FM4 apply in FM6 to the sali- references: Lewis and Mislove, 1953, DIS 27: 58.
vary chromosome region from IB to 15D. In geno- properties: Viability and fertility of heterozygote
types with a normal recombination rate, FM6 effec- excellent. Reliable balancer for all of chromosome
tively eliminates recombination in FM6/+ heterozy- 2, although there is an occasional double cross-
gotes but yields viable recombinants when hetero- over in 2R if X and 3 are heterozygous for inver-
gous for such inversions as In(l)sc8. Used for bal- sions. Mclntyre and Wright (1966, DIS 41: 141-42)
ancing sex-linked recessive lethal and sterile found no recombination between In(2LR)SMl and
mutations. Does not effectively balance cv or v in al dp b pr en c px sp in females also heterozygous
stocks that are also heterozygous for In(2LR)SMl for what behaves like In(l)sc8 and In(3LR)UbxI3°.
and In(3LR)Ubx13°. *SM2
constitution: T(2;3)SM2, al2 Cy It? en2 sp2.
constitution: ln(l)dl-49+BM1, sc vBM1. synthesis: R. F. Grell, 1953.
properties: Male and homozygous female viable and references: Lewis and Mislove, 1953, DIS 27: 58.
fertile. Effective suppressor of crossing over in X 1954, DIS 28: 77.
chromosome. properties: Not useful as a balancer.
In(l)sc7+AM
constitution: In(l)sc7+AM, sc7. SMS
properties: Male viable and fertile; homozygous fe- constitution: In(2LR)SM5, al2 Cy ltv en2 sp2.
male viable but sterile because of homozygous synthesis: R. F. Grell, 1953.
In(l)AM. Reduces X chromosome crossing over. references: Mislove and Lewis, 1955» DIS 29: 75.
May be used to balance sex-linked recessive lethal properties: Heterozygote usually has good viability
or sterile mutations. and fertility, although may not be so good as SMI.
Most complete balancer for chromosome 2.
TMh Third Multiple
constitution: In(l)scsl+dl-49, scsl v f car. constitution: In(3LR)TMl, Me ri sbd2.
properties: Male viable and fertile; homozygous fe- synthesis: E. B. Lewis,
male viable but sterile because of homozygous references: 1949, DIS 23: 92.
In(l)scs*. Reduces crossing over in X chromo- Lewis and Mislove, 1953, DIS 27: 58.
some. May be used to balance sex-linked reces- properties: Homozygous lethal. Suppresses crossing
sive lethal or sterile mutations. over in chromosome 3.
Insc TM2
constitution: In(l)scsi Lsc8R+dl-49, acsl sc8. constitution: In(3LRyUbxl30> rjhxÔ e «.
synthesis: Muller. synthesis: E. B. Lewis.
properties: Male and homozygous female viable and references: 1952, Proc. Natl, Acad. Sci. U.S. 38:
fertile. Suppresses crossing over in the X chromo- 953-61.
some. 1952, DIS 26: 66.
Inscy properties: Homozygous lethal. Eliminates crossing
constitution: In(l)scslLsc8R+dl-49, y scsi sc8. over in chromosome 3 except at the end of the right
synthesis: Muller. arm. Does not reliably balance mutations in the
properties: Male and homozygous female viable and vicinity of ca. Maclntyre and Wright (1966, DIS 41:
fertile. Crossing over suppressed in X chromo- 141—42) observed about 9 percent double crossing
some. over in the unbroken segment of the left arm from
LVM: Balancer of L. V. Morgan 61C to 74 on the salivary gland chromosome map
constitution: In(3L)P with a lethal in each arm of and 15 percent recombination between the break-
chromosome 3. point at 96A and ca in In(3LR)Ubx*3Q,
M-5: see Base Ubx130 e/ru h tfi at cu sr es Pr ca females that
Muller-5: see Base were also heterozygous for an X-chromosome inver-
S-5 sion behaving like In(l>sc* and In(2LR)SMl; no
constitution: In(l)sc4Lsc8R+S, y sc 4 sc8 w* B. recombination observed in other regions.
synthesis: Lindsley. TM3
references: Lindsley and Edington, 1957, DIS 31: constitution: In(3LR)TM3, y+ ri pP sep bx-»*e e».
131-32. synthesis: E. B. Lewis, 55g.
Lindsley, Edington, and Von Halle, 1960, Genetics references: Mitchell, 1958, Cold Spring Harbor
45: 1649-70. Symp. Quant. Biol. 23: 279-90.
Lewis, 1960, DIS 34: 51. Novitski (1954, Genetics 39: 127-40) as reversed
Tinderholt, 1960, DIS 34: 53-54. acrocentrics, reversed metacentrics, reversed rings,
properties: Stocks exist in which Ser or Sb and Ser tandem acrocentrics, tandem metacentrics, and
are present. With normal X and 2, all of chromo- tandem rings; where applicable this classification
some 3 is effectively balanced; in the presence of was retained and is used in the designation of com-
FM6 and SMS, however, crossing over between y + pounds.
and ri, i.e., in 61A2-65E, is appreciable. Double When the component arms differ in sequence by
crossovers that separate Sb or Ser from inversion something other than whole-arm inversion, the clas-
complex rare, even in presence of FM6 and SM5. sification tandem or reversed becomes ambiguous.
TM6 Furthermore, when the component arms are separable
constitution: In(3LR)TM6, ss~ bx34e e; also exists from each other by a single break, the terms acro-
with UbxtTb. centric and metacentric are descriptive; but when
synthesis: E. B. Lewis and F. Bacher, 66i. elements of the two arms become interspersed, as
properties: Should effectively balance entire third for example by interarm rearrangements, these terms
chromosome, but has not been tested. Has un- lose meaning. Consequently, the more-complex com-
broken regions with genetic lengths of approxi- pounds are given arbitrary symbols.
mately 10, 15, 20, and 30 units. The chromosomal constitution of compounds in
winscy which the chromosome arms remain intact is desig-
constitution: Jn(l)scSiLsc8R+dl-49, y scsl sc8 w. nated: metacentrics, by the sequences of the compo-
synthesis: Muller. nent arms separated by a centerpoint (which repre-
properties: Male and homozygous female viable and sents the centromere); acrocentrics, by the sequence
fertile. Suppresses crossing over in X chromo- of the distal arm separated by an em dash from the
some. sequence of the proximal arm followed by a center-
point; rings (which are derived from acrocentrics or
COMPOUND CHROMOSOMES metacentrics) by origin. In heterozygotes, the gene
Compound chromosomes are monocentric elements content of the component arms is listed according to
in which the material from one chromosome arm is the same conventions, with the genes on the first
represented twice; they contain the entire diploid arm listed in the chromosomal designation followed
complement for the arm involved. They are desig- by those on the second arm. In homozygotes, the
nated by the symbol C followed parenthetically by genes are listed in chromosome map order. Complete
the designation of the involved arm. Gametes of designation of a compound includes its symbol, its
compound-bearing flies generally carry two or no chromosomal constitution, and the gene content of
doses of the chromosome arm. Compound-Jf chromo- its component arms; e.g., C(1)TM2, +-In(l)sc4LENR,
somes, C(l)'s, exist only in females, which unless y cv v sd'y sn g. It should be emphasized that the
special steps are taken, carry a Y chromosome. Such heterozygous gene content of compounds is often
C(l)/Y females produce patroclinous sons, which in- highly unstable owing to homozygosis and changes
herit their X from their father and their Y from their in coupling relations resulting from exchange.
mother, and matroclinous daughters, which inherit In compounds in which elements of the component
both their X'% from their mother and a Y from their arms have become interspersed, it is usually not
father (so-called noncrisscross inheritance). Com- feasible to designate the chromosomal constitution
pound-autosome-bearing flies usually produce no vi- in terms of the component arms; rather, it is de-
able progeny unless crossed to flies carrying com- scribed in terms of the order of chromosome seg-
pounds for the same arm or arms. ments as seen in salivary-gland chromosomes. In
Some compounds have arisen repeatedly from cer- heterozygotes, the gene content is listed in such a
tain genotypes; they were studied collectively but way as to indicate which genes were originally in
not as individual occurrences. In other cases, sim- the different component arms.
ilar compounds of independent origin were studied • = : see C(1)RM
individually. Both general classes of compounds : = : see C(1)DX
and compounds of unique origin are listed. 2L: see C(2L)RM
The two chromosome arms comprising a compound 2R- see C(2R)RM
may join (1) by attachment of the base of one to the 3L: seeC(3L)RM
terminus of the other to form an acrocentric chromo- 3R: see C(3R)RM
some or (2) by attachment of both proximally to a Attached 2L: see C(2L)RM
single centromere to form a metacentric; the ends of Attached 2R: see C(2R)RM
either an acrocentric or a metacentric may join to Attached 3D: see C(3L)RM
form a compound ring. In addition, the component Attached 3R: see C(3R)RM
arms may be in the same sequence or one may be en- Attached-X: see C(1)RM
tirely inverted with respect to the other. Thus the
elements of a compound may pair as a spiral — the C(1)94-2A
tandem configuration or as a hairpin — the reversed constitution: Homozygous for y; originally heterozy-
configuration. Simple compounds may therefore be gous for cv, sn, v, g, and sd. Ring shaped in mi-
classified according to the conventions of totic metaphase. Salivary chromosome analysis
SPECIAL CHROMOSOMES - COMPOUND 409
shows order to be |lA - 5E|lF - 1A-20 - 5E|lF -

20|. com
origin: Spontaneous (although possibly X ray in- constitution: C(iyM3, In(l)AM -ln(l)FM4',
duced premeiotically) derivative of C(l)TR94-2. y2 — y~ sc8 dm £?•.
Apparently arose through an asymmetrical or re- origin: Recombinant between In(l)sc7+ AM element
versed exchange between the IF region near the of C(1)M2 and In(l)AM, y2 in triploid.
centromere and the 5E region near the interstitial synthesis: E. B. Lewis, 55b.
heterochromatin of C(l)TR94-2. synonym: FMA3.
synthesis: Rosenfeld, 1964. references: 1958, DIS 32: 81-82.
properties: Crossing over in region 1F-6A produces properties: Detachment rare; useful in balancing.
a single ring carrying In(l)94-2A — ln(l)lF-2A;5E-
6A. Reversibly convertible to other double-ring
configurations by other types of exchange (e.g.,
Novitski and Braver, 1954, Genetics 39: 197—209).
C(1)A: Compound(l) of Armentrout

constitution: Homozygous for y and probably origi-
nally heterozygous for cv, sn, v, g, and sd. Ring
shaped in mitotic metaphse. Salivary chromosome
analysis shows order to be 11A — 6F2|6F2 —
1AJ20 - 7Al|7Al - 20-|.
origin: Spontaneous stable derivative of C(1)TR94,
which was originally y cv v sd>y sn g. Apparently
arose by a process describable as reversed cross-
ing over in region 6F2-7A1. C(1)NB: Compound(l) of Novitski and Braver
synthesis: Armentrout, 1964. From Novitski and Braver, 1954, Genetics 39: 197-
properties: An apparently completely stable com- 209.
pound-ring-^ chromosome; cannot produce single-
X-chromosome derivatives by heterochromatic ex- C(1)NB: Compound(l) of Novitski and Braver
change. Should be the best of all compound-X constitution: C(1?JB, In(l)dl-49'In(l)sc^LENR;
chromosomes for stock purposes. originally y v f car-y m; In(l)dl-49 and In(l)EN
attached proximally to a single centromere.
CO)DX: Compound(l) Double X origin: Crossover between the heterochromatic short
constitution: C(1)DX, In(l)dl-49 - In(l)sc8-, arm of In(l)EN and the proxima 1 heterochromatin of
y f — y~ sc8 I'. In(l)dl-49.
origin: X ray induced in In(l)dl-49, y w f/In(l)sc8, synthesis: Novitski and Braver.
sc8 B female [stated by Muller to have been references: 1954, Genetics 39: 197—209 (fig.).
In(l)dl-49/In(l)ac8Ly3PI*, but the derivative does properties: Essentially a tandem metacentric hetero-
not carry y 3 J P ] . Was originally y w I — y— sc8 B', zygous for In(l)dl-49. Can exist in a number of dif-
but by double exchange / became homozygous and ferent configurations interconvertible by crossing
B was lost. over. Generates single rings at different fre-
synthesis: Muller. quencies, depending on configuration of the com-
synonym: The symbol ; = . pound.
6
c
Valencia, Muller, and Valencia, 1949, DIS 23: 9 9 -
102. sc
properties: A reversed acrocentric heterozygous for
ln(l)dl-49; since it is very stable, probably be-
cause there is little interstitial heterochromatin, it
is useful in balancing, y w f detachments produced
very rarely. Also produces a low incidence of ho- C(1)RA: Compound(l) Reversed Acrocentric
mozygosis for w. C(l)DX/0 lethal; probably defi- Redrawn from Sandier, 1954, Genetics 39: 923-42.
cient for bb.
C(m2: Compound^) Multiple C(1)RA: Compound(l) Reversed Acrocentric
constitution: C(1)M2, In(l)sc7+AM -In(l)FM4-, constitution: C(1)RA, + — In(l)sc8".
sc7 — y~ sc8 dm B'. origin: Spontaneous from X>YL/In(l)scs either by
origin: X-ray-induced exchange between the proximal exchange between the proximal heterochromatin of
heterochromatin of In(l)sc7+AM and the distal X'YL and the distal heterochromatin of In(l)scs or,
heterochromatin of ln(l)FM4. more likely, by sister-strand union in one of the
synthesis: Lewis, 54h. heterochromatic segments followed by a normal
synonym: FMA2: First Multiple Attached. euchromatic exchange. A frequently recurring
references: 1958, DIS 32: 81. event that seems to require the presence of Y^.
synthesis: Novitski. origin: X ray induced in an attached-X with In(l)sc8

synonym: RA. and In(l)scslLENR attached proximally to a single
references: Novitski, 1954, Genetics 39: 127—40. centromere. Recovered as simultaneous loss of y"*"
Sandier, 1954, Genetics 39: 923-42. from the tip of both arms.
1958, Cold Spring Harbor Symp. Quant. Biol. 23: synthesis: Sandier.
211-23. references: 1957, Genetics 42: 764-82 (fig.).
properties: Yields frequent detachments resulting 1958, Cold Spring Harbor Symp. Quant. Biol. 23:
from exchange between the Y chromosome and the 211-23.
interstitial heterochromatin of the reversed acro- properties: Tetrad distribution abnormal, with one-
centric. Tetrad distribution usually quite abnormal; exchange tetrads being infrequent but no- and two-
one-exchange tetrads infrequent and no- and two- exchange tetrads being frequent. Exchange fre-
exchange tetrads frequent. Exchange frequency in- quency increased by addition of Y or y*YL but
creased by addition of Y or y+YL, but tetrad distri- tetrad distribution remains abnormal.
bution remains abnormal (Sandier, 1954). YL ap-
pended as a second arm to C(1)RA normalizes
tetrad distribution (Sandier, 1958).
C(l)RA60g
constitution: C(l)RA60g, + -In(l)sc8.
origin: A spontaneous exchange between distally
located heterochromatin of In(l)sc8 and proximal
heterochromatin of a normal X. Occurred in a trip-
loid female. C(1)RR2: Compoundfl) Reversed Ring 2
synthesis: Mohler, 60g. From Sandier, 1957, Genetics 42: 764-82.
other information: The reciprocal exchange product, C(J)RR94-2F
Dp(l;f)60£, recovered from same fly. constitution: C(1)RR94-2F, +•+; two X chromosomes
C(1)RM: Compound(T) Reversed Meiacenfric of normal sequence attached proximally to a single
constitution: C(1)RM, +•+; two X chromosomes in centromere and joined distally by a segment of
normal sequence attached proximally to the same heterochromatin.
centromere; exists with many combinations of origin: X-ray-induced derivative of C(1)TR94.
markers. synthesis: Rosenfeld, 1964.
origin: Spontaneous. Recurs regularly by exchange references: Sandier, 1965, Natl. Cancer Inst. Mono-
between heterochromatin of the short arm of one X, graph No. 18: 243-72.
X'YS, or X-YL and that of the base of the long arm properties: Tetrad distribution more nearly normal
of a sister or homolog. than in C(1)RR2.
discoverer: L. V. Morgan, 21bl2. C(1)SB: Compound(l) of Sturtevant and Beadle
synonym: Attached-X; also the symbol •=. constitution: C(1)SB, +'In(l)y4; In(l)y4 and a normal
references: 1922, Biol. Bull. 42: 267-74. sequence attached proximally to a single centro-
1938, Am. Naturalist 72: 434-46. mere.
properties: Recombination with the Y chromosome origin: Recombinant between the uninverted portion
leads to detachments with a frequency of about of In(l)y4 and C(1)RM in a triploid.
10-3 ^ C(1)RM/Y females. Has been extensively synthesis: Sturtevant and Beadle.
used in studies of crossing over (e.g., Anderson, references: 1936, Genetics 21: 554—604.
1925, Genetics 10: 403-17; Beadle and Emerson, Novitski and Sandier, 1956, Genetics 41: 194-
1935, Genetics 20: 192-206; Welshons, 1955, 206.
Genetics 40: 918-36). properties: A reversed metacentric heterozygous for
*C(1)RRJ: Compound(l) Reversed Ring In(J)y4. Meiotic behavior similar to that of a
constitution: C(1)RR1, + - In(l)EN, y— sc— — y; tandem metacentric. Crossing over within inver-
two X chromosomes attached by their normally sion generates single ring, R(l)y4.
distal ends to a common centromere and by their *C(1)TA1: Compound(l) Tandem Acrocentric
normally proximal ends to each other. Marked constitution: C(1)TA1, In(l)sc4 - In(l)EN-YL,
with y. y sc* — y .
origin: Spontaneous derivative of C(1)TR1. origin: X-ray-induced exchange between the prox-
synthesis: Zimmering. imal heterochromatin of ln(l)sc4 and Ys of
synonym: RR. YSXYL.
references: Novitski, 1954, Genetics 39: 127-40. synthesis: Novitski.
*C(1)RR2 synonym: TA.
constitution: C(l)RR2f In(l)scS.In(l)acSlLENR; references: 1954, Genetics 39: 127-40.
originally y~ cv v f-y m car, In(l)&c* and properties: Produces a single centric rod-X chromo-
In(l)scslLElVR attached proximally to a single some and either an acentric ring or a tandem-triple-
centromere and distally at their distal heterochro- X chromosome by recombination between the prox-
matic segments. imal and distal X chromosomes.
SPECIAL CHROMOSOMES - COMPOUND 411
+ car
C(1)TA2: Compound(l) Tandem Aero centric 2

From Sandier and Lindsley, 1963, Genetics 48: 1533-
43.
C(1)TA2 C(1)TM2: Compound(l) Tandem Mefocen/r/c 2
constitution: C(1)TA2, H +•; originally y cv f — From Lindsley and Sandier, 1965, Genetics 51: 2 2 3 -
y f: 45.
origin: X-ray-induced recombinant in
YSX-, y + Ks y cv v f/X'YL, y car-KL female; C(l)TMBs9-h Compound(i) Tandem Mefocenfr/c
origin required triple exchange. with Bar-Stone
synthesis: Sandier and Lindsley. constitution: C(1)TMBS9-1,
references: 1963, Genetics 48: 1533-43 (fig.). Dp(l;l)BsTAG'In(l)sc«LENR; originally
properties: Generates single-X chromosomes like Bs y cv v sdy sn g. In mitotic prophase the se-
C(1)TA1. Tetrad distribution about normal. quence is: the normal X euchromatin, two large
C(l)TA2/0 lethal; probably deficient for bb. heterochromatic segments, a small segment, the
C(1)TM1: Compound(l) Tandem Met acentric centromere, a small segment, the inverted X
constitution: C(1)TM1, +'In(l)scSÊNR, euchromatin.
y Hw f-y+ y i; a normal sequence and In(l)EN origin: X-ray-induced exchange between the prox-
attached proximally to a single centromere derived imal heterochromatin of Dp(l;l)BsTAG and YL of
from R(l)2. X-YL, In(l)sc8LENR.
origin: Product of one crossover between + and synthesis: Lindsley and Sandier, 1963.
R(l)2 and one between In(l)EN and R(l)2 in a synonym: TMXBS 9-1; also designated as
+/R(l)2/In(l)EN triploid. Dp(l;l)BsTRG.
synthesis: Novitski, 1950. references: 1965, Genetics 51: 223—45.
references: Novitski and Lindsley, 1950, DIS 24: properties: Recombination between the arms pro-
90-91. duces a single-ring-X chromosome, R(l)9-1, and an
properties: Single crossover between the arms pro- acentric rod X. Recombination between the Bs
duces s ingle-ring-X chromosome with the same duplication and the homologous region of the in-
structure as R(l)2 and an acentric rod-X chromo- verted arm generates a nontransmissible tandem-
some. Tetrad distribution about normal (Novitski, ring chromosome. Meiotic behavior similar to that
1951, Genetics 36: 267-80; Novitski and Sandier, of C(1)TM2.
1956, Genetics 41: 194-206).
C(1)TM2 C(l)TMBS9-4
constitution: C(1)TM2, +-In(l)sCfÊNR; originally constitution: C(l)TMBS9-4,
y cv v sd-y sn g. In mitotic prophase the sequence Dp(l;l)BsTAG'ln(l)sc«l'ENR; originally
is: the normal X euchromatin, two large hetero- Bs y cv v sd'y sn g. In mitotic prophase the se-
chromatic segments, a small segment, the centro- quence is: the normal X euchromatin, a large het-
mere, a small segment, the inverted X euchromatin. erochromatic segment, a small segment, the centro-
origin: X-ray-induced exchange between the prox- mere, a small segment, the inverted X euchromatin.
imal heterochromatin of a normal X and YL of origin: X-ray-induced exchange between the prox-
X-YL, In(l)ac*ÊNR. imal heterochromatin of Dp(1;1 )BSTAG and YL of
synthesis: Lindsley and Sandier, 1963.
synonym: TMX y. synthesis: Lindsley and Sandier, 1963.
references: 1965, Genetics 51: 223-45 (fig.). synonym: TMXBs9-4; also designated as
properties: Recombination between the arms pro- Dp(l;l)BsTRG.
duces a s ingle-ring-.X chromosome and an acentric references: 1965, Genetics 51: 223-45 (fig.).
rod X. Meiotic behavior similar to that of C(1)TM1; properties: Recombination between arms produces
tetrad distribution about normal. single-ring-J? chromosome, R(l)9-4, and an acentric
rod X. Recombination between the Bs duplication C(1)TR94

and the homologous region of the inverted arm pro- constitution: C(1)TR94, +-ln(l)sc*LENR-; originally
duces a tandem-ring chromosome that may be trans- y cv v sd-y sn g.
missible. origin: Regular but infrequent product of
C(l)TMBs9-4. Formed by exchange between the
duplicated Bs section and the homologous region
of the inverted arm.
synthesis: Sandier and Lindsley.
properties: Produces a centric single-ring-X and
either an acentric single-ring-X or a tandem triple-
ring-X chromosome by crossing over between the
two arms of the compound. Transmission higher
than that of C(1)TR1. Tetrad distribution about
normal.
*C(1)VM: Compound(l) of Valencias and Muller
constitution: C(1)VM, + — In(l)scsl+dl-49'; originally
y ac sc pn w rb cm ct& sn3 oc ras2 v dy g t car —
y acSi lza JEK
origin: X ray induced in +/In(l)scsl+dl-49/YL fe-
male, either by exchange between the proximal het-
C(l)TMBs: Compound(l) Tandem Me/ocentr/c erochromatin of the normal sequence and the distal
with Bar-Stone heterochromatin of In(l)scsi or by sister-strand
From Lindsley and Sandier, 1965, Genetics 51: 223- union in one of the heterochromatic elements ac-
45. companied by normal euchromatic exchange. A
regularly induced product in such females.
*C(1)TR1: Compound(l) Tandem Ring synthesis: Valencia, Muller, and Valencia.
constitution: C(1)TR1, In(l)sc* _ ln(l)EN% references: 1949, DIS 23: 99-102.
y— sc— — y. properties: Essentially a reversed acrocentric in
origin: Spontaneous derivative of C(1)TA1 in which which the proximal element contains In(l)dl-49.
the YL second arm had been replaced by the 4DXP Detachment by crossing over with a Y chromosome
element of T(1;4)B$ = T(1;4)15F9-16A1;16A7- relatively frequent.
Bl;102F. A product of recombination between the C(2L)RtA: Compound(2L) Reversed Mefacentric
duplicated Bs second arm and the homologous re- constitution: 2L'2L; exists with various marker
gion of the distal element of the tandem acrocen- combinations. Two left arms of chromosome 2
tric. attached proximally to a single centromere.
synthesis: Novitski. origin: X ray induced.
references: 1954, Genetics 39: 127-40. synthesis: Rasmussen, 60c.
properties: Seems to be poorly transmissible synonym: 2L: Attached 2L.
(Novitski, 1954). Produces a centric single-ring-X references: 1960, DIS 34: 53.
and either an acentric single-ring-X or a tandem properties: Produces viable progeny only in crosses
triple-ring-JT chromosome by recombination between in which both parents carry C(2L)RM. Usually
the two elements of the compound. carried in stock of constitution C(2L)RM; C(2R)RM.
Such males may produce viable progeny in crosses
to triploid females and to irradiated females in
which new attachments are produced.
C(2R)RM
constitution: 2R'2R; exists with various marker
combinations. Two right arms of chromosome 2
synthesis: Rasmussen, 59k.
synonym: 2JR_: Attached 2R.
references: I960, DIS 34: 53.
properties: Similar to C(2L)RM.
C(3L)RM
constitution: 3L-3L; exists with various marker
combinations. Two left arms of chromosome 3
C(1)TR94: Compound(l) Tandem Ring 94 origin: X ray induced.
From Sandier and Lindsley, 1967, Genetics 55: 645- synthesis: Rasmussen, 59f.
71. synonym: JZ-: Attached 3L.
SPECIAL CHROMOSOMES - MULTIPLY MARKED 413
references: 1960, DIS 34: 53. Brfspl

properties: Produces viable progeny only in crosses constitution: al dp b Bl c px sp.
in which both parents carry C(3L)RM. Usually synthesis: Muller.
carried as C(3L)RM; C(3R)RM. Such males may hes
produce viable progeny in crosses to triploid fe- constitution: h th st pP cu sr eB.
males and to irradiated females in which new synthesis: Bridges, 1923.
attachments are produced. maple
constitution: y ac sc pn w rb cm ct6 ras2 v g2 f car.
C(3R)RM synthesis: Muller.
constitution: 3R-3R; exists with various marker peple
combinations. Two right arms of chromosome 3 constitution: pP ss k ea ro.
attached proximally to a single centromere. synthesis: Muller, 1914.
origin: X ray induced. p/oc
synthesis: Orias and Deal, 581. constitution: y ac sc pn w rb cm ct6 sn3 oc ras2
synonym: 3R: Attached 3R. v dy g f car.
references: Rasmussen, 1960, DIS 34: 53. synthesis: Muller.
properties: Similar to C(3L)RM. references: Valencia, Muller, and Valencia, 1949,
C(4)RM DIS 23: 99-102.
constitution: 4.4; exists with various marker combi- properties: Used for specific-locus mutation
nations. Two right arms of chromosome 4 attached studies.
proximally to a single centromere. res
origin: X-ray induced, constitution: ru h th st pP cu sr ea.
synthesis: E. B. Lewis, synthesis: Bridges, 1925.
properties: Produces haplo-4 and triplo-4 progeny in synonym: rupes.
crosses to normal diplo-4 flies. rucuco
First Multiple Attached: see C(1)M constitution: ru h th st cu sr ea ca.
FMA: see C(1)M synthesis: Bridges, 1926.
RA: see C(1)RA
RR: see C(1)RR rupes: see res
TA: see C(1)TA ruPrica
TMXBS: see C(l)TMBs constitution: ru h th st cu sr ea Pr ca.
sc-tester
MULTIPLY MARKED CHROMOSOMES constitution: sc ec ct6 t2 v g3 si.
synthesis: L. V. Morgan.
Z-ple seple
constitution: dp b pr c px sp. constitution: se ss k ea ro.
synthesis: Muller. synthesis: Muller, 1914.
references: 1925, Genetics 10: 470—507. tester 1
3-ple constitution: y ac pn w rb wy2 g2.
constitution: ru h st pP ss es. synthesis: Muller.
synthesis: Bridges, 1920. tester 2
references: 1927, J. Gen. Physiol. 8: 689—700. constitution: y2 w* cm wy2 g2 car.
S-ple synthesis: Muller.
constitution: b pr vg a sp, tester 3
synthesis: Muller, 1914. constitution: y rb cm ras2 g2.
72-p/e synthesis: Muller.
constitution: at dp b pr en vg c a px bwmi sp. theca
synthesis: Muller. constitution: th cu sr es ro ca.
synonym: twelve-pl. synthesis: Bridges, 1925.
albasp thes
constitution: a/ bpr en vg a sp2. constitution: th st pP cu sr e s .
synthesis: Bridges, 1926. synthesis: Bridges, 1924.
all thrike
constitution: el dp b pr c px sp. constitution: th at cp in ri pP ss bxd k e s .
synthesis: Bridges, 1926.
synonym: apt. twelve-pl: see 12-ple
oll-BI X-6
constitution: at dp b pr Bl c px sp. constitution: sc ec ct6 g2 Bx2 bb.
apl: see all synthesis: L. V. Morgan, 1928.
bleached X-7
constitution: pn w rb cm ct6 mi3 ra&2 v dy g2 f car. constitution: y oc cv ct6 v g2 f.
synthesis: Muller. synthesis: L. V. Morgan, 1928.
4T4 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER
X-8 synthesis: Stern.

constitution: y ec cv cfi v g? f bb. synonym: XY .
X-9 1929, Z. biduktive Abstammungs- Vererbungslehre
constitution: sc ec cv erf v s ^ f car bb1.
51: 253-353.
synthesis: Bridges, 1 9 3 1 .
Kaufmann, 1933, Proc. Natl. Acad. Sci. U.S. 19:
X-p/e 830-38 (fig.).
constitution: sc ev cv ct6 v g2 t
properties: An X chromosome in normal sequence
with YL appended as a second arm. May carry
references: Bridges and Albright, 1926, Genetics varying amounts of the proximal part of Ys between
11: 41-56. the X and the centromere. Males carrying X>YL re-
quire KS in some form for fertility. Chromosome V
X-Y COMBINATIONS shaped in metaphase.
The X and one or both arms of the Y chromosome

may be linked by recombinational events occurring y* su(wahva
in the heterochromatin. Such X-Y combinations are
composed of the X, a centromere (derived from either
X or F), and either YL or Ys or both. In the designa-
tion of such chromosomes, the component elements
are listed in order such that X precedes the centro-
mere (symbolized by a centerpoint), e.g., YSX'YL.
Events that give rise to X-Y attachments are usually
recurring so that the same combinations arise re- From Lindsiey and Novitski, 1959, Genetics 44: 187-
peatedly; however, since the exact points of ex- 96.
change differ, independent occurrence of similar
combinations certainly differ from one another in
heterochromatic content. Since similar X-Y combina- constitution: XYL-YS; originally
a a
tions of independent origin are not ordinarily desig- y2 su(w ) w (bb?)KL-KS.
nated, studied, or maintained as different chromo- origin: X-ray-induced detachment in C(1)RM,
somes and since, for most purposes, it is not impor- y2 su(wa) w* bb/y+Y female.
tant that they be distinguished, general categories synthesis: Parker.
of X-Y combinations are largely described in the en- references: Parker and McCrone, 1958, Genetics 43:
suing section. Where a specific combination has 172-86.
been studied, it is listed with the designation of its Lindsiey and Novitski, 1959, Genetics 44: 187-96
component elements followed immediately by its (fig-).
specific designation, e.g., X'YLC2. properties: Essentially an intact Y chromosome with
The complete designation of an X-Y combination all of the X euchromatin appended distal to KL.
consists of the following items in the order given: Carries all the sex-chromosome material required
chromosomal elements, sequence of the X chromo- for male viability and fertility.
some (if other than normal), gene content. X-Y com- other information: Several detachments of this con-
binations that differ from one another only with re- stitution recovered; numbered 2-10T13, 2-10T15,
spect to mutant genes or euchromatic inversions are 108-9, 112-17, and 129-11.
not described separately because it is considered
that such mutants and inversions can be removed
from or inserted into the component X by euchro-
matic exchange. When X's differ by an inversion y* Y L
with at least one heterochromatic breakpoint, the
chromosomes are described separately since they
must differ in their heterochromatic constitution.
FR1: see Y*X-
Fragment t: see YSX-
sc* co. X: see Y*X-, /nfijsc*
sc*EN co. X: see Y*X', In(l)EN
XY':
constitution: XYL'YS; originally
y2 su(wa) wa (bb?) y +KL-KS.
origin: A recurrent product of exchange between the origin: X-ray-induced detachment in C(1)RM,
proximal heterochromatin of C(1)RM and either arm y2 su(wa) wa bb/y+Y female.
of the Y. Also may result from exchange between synthesis: Parker.
Ys and the proximal heterochromatin of a normal X references: Parker and McCrone, 1958, Genetics 43:
or the interstitial heterochromatin of C(l)RA. 172-86.
SPECIAL CHROMOSOMES - X-Y COMBINATIONS 415
properties: Essentially an intact y+Y chromosome KS. Carries all the sex-chromosome material re-
with all of the X euchromatin attached to YL distal quired for male viability and fertility.
to y +. Interstitial position of y+ shown by recovery other information: Two detachments of this constitu-
of y + reattachment; also interstitial y + allele tion recovered; numbered 110-8 and 115-9.
shows strong variegation. Carries all the sex-chro-
mosome material required for male viability and
fertility. constitution: YSX'; originally KS y cv v f (Braver).
origin: Spontaneous from YSX'YL, ln(l)EN,
XV-C2 KS yKL/sc cv v f female.
constitution: X'YL, bb~-KL. synthesis: Novitski.
origin: Recombination between Ys proximal to bb + synonym: FR1: Fragment 1,
and C(1)RM distal to bb +. references: 1952, Genetics 37: 270—87.
synthesis: Lindsley. Lindsley and Novitski, 1959, Genetics 44: 187—
properties: Like X'YL but deficient for bb; 96.
X'YLC2/0 lethal. Shows unique behavior in double properties: An X in normal sequence with Ys ap-
first-anaphase bridges (Novitski, 1952, Genetics pended distal to 1(1)JI+ and y. Lowers crossing
37: 270-87). over near y.
XY<-, ln(l)EN V ln(1)EN

constitution: X-YL, In(l)EN, yKL. constitution: YSX; ln(l)EN, KS y.
origin: Recombinant from YSX'YL, In(l)EN/In(l)EN origin: Infrequent product of spermatogonial ex-
female. change between Ys and the distal inverted hetero-
genetics: An entirely inverted chromosome with Y^ chromatic segment of In(l)sc8LENR. Also formed
appended as a second arm. by recombination in YSX-, Jn(l)sc8/In(l)EN fe-
males. Specific occurrence described as
Df(l)sc8P7, deficiency subsection.
origin: Recurrent product of recombination between synthesis: Lindsley.
the proximal heterochromatin of C(1)RM and the Y. synonym: sc8ENco. X.
synthesis: Kaufmann. references: 1955, Genetics 40: 24—44.
references: 1933, Proc. Natl. Acad. Sci. U.S. 19: properties: A completely inverted X chromosome
830-38. with Ys appended distally. The distal heterochro-
properties: An X chromosome in normal sequence matin resembles that of In(l)sc8 in mitotic pro-
with Ys appended as a second arm. May carry phase and carries bb+.
varying amounts of the proximal part of YL be-
tween X and the centromere. Males carrying X'YS YSX-, In(l)sc8
require KL in some form for fertility. Chromosome constitution: YSX', In(l)sc8, KS 1(1)J1~ y- ac~ sc*.
J shaped in metaphase. origin: Infrequent product of spermatogonial ex-
change between Ys and the distal inverted hetero-
chromatic segment of In(l)sc8. Also formed by re-
combination in YSX>, In(l)EN/In(l)sc* females.
Specific occurrence described as Di(l)sc889a, de-
ficiency subsection.
synthesis: Sidorov.
synonym: sc'c.o. X.
references: 1940, Bull. Biol. Me'd. exp. URSS 9:
10-12.
1941, Dolk. Acad. Nauk SSSR 30: 248-49.
From Lindsley and Novitski, 1959, Genetics 44: 187 Lindsley, 1955, Genetics 40: 24-44.
96. properties: In(l)sc8 with the distal uninverted
euchromatic region carrying the normal alleles of
Hl)Jli Yt a n d ac replaced by KS, Chromosome re-
constitution: XYS-YL; originally sembles In(l)sc8 in mitotic prophase.
y2 su(w*) w* (bb?) KS-KL y+.
origin: X-ray-induced detachment in C(1)RM,
y2 8u(w*) w* bb/y+Y female, constitution: YSX-YL, KS y-KL.
synthesis: Parker. origin: Recombination between YSX' and X'YL.
references: Parker and McCrone, 1958, Genetics 43: synthesis: Lindsley and Novitski.
172-86. references: 1959, Genetics 44: 187-96.
Lindsley and Novitski, 1959, Genetics 44: 187-96 properties: Contains all of the sex-chromosome ma-
(fig.). terial required for male viability and fertility.
properties: Essentially an intact y*Y chromosome Commonly kept in stock as YSX'YL/O males
with all of the X euchromatin appended distal to crossed to C(l)/0 females.
separated loci known to be included in it separated

by an em dash, e.g., bw+ — ba+. The constitution of
a marked F or F fragment may be designated by
listing its genetic elements in order, with any ambi-
guities in order enclosed within parentheses, e.g.,
KL(bw+ - ba+)-bb+KS. When there is a hierarchy of
ambiguities in order, a hierarchy of parentheses is
y used, as in ((ci + _ spa+)KL)-bb+KS.
L, ln(l)EN 4Y
From Lindsley and Novitski, 1959, Genetics 44: 187— constitution: ((ci+ — spa+)KL)'bb+KS; tentative.
9o. origin: X ray induced.
synthesis: Edmonds on, 1946.
Y^X-YL, ln(1)EN synonym: Tp4'Y: Transposition 4-Y.
constitution: YSX-YL, In(l)EN, KS y-KL y+. references: Muller and Edmondson, 1957, DIS 31:
origin: Recovered as recombinant between the 140-41.
proximal heterochromatin of YSX% In(l)EN and properties: Contains all known loci of chromosome 4
y+Y. linked to the Y chromosome. Results from recombi-
synthesis: Lindsley, 1949. nation between 4Y and YSX-YL suggest that 4 is
references: Lindsley and Novitski, 1960, DIS 24: inserted into or appended to YL. Two doses of
84-85. this chromosome in the absence of any other F- or
1959, Genetics 44: 187-96 (fig.). 4-derived material produce viable and fertile flies
properties: Contains all of the sex-chromosome ma- of both sexes.
terial required for male viability and fertility. *B*w+y+Y
Exists without the y+ marker at the terminus of the constitution: Bs pdf+ su(f)+ kz+ —
YL arm; also exists with various combinations of spl+ y+ ac+ KL'bb+ KS; inferred from origin.
sex-linked markers. origin: X-ray-induced deletion of majority of euchro-
matin (dm + through mal+) from the recombinant
Y DERIVATIVES composed of left end of YDXP element of
T(1;Y)148 = T(1;Y)2D;YL, which involves BSY,
The F chromosome consists of a long arm, YL, and and right end of XYL-YS129-16, which carries y +
a short arm, Ys; the long arm is arbitrarily taken as from y+Y between X and YL.
the left arm. There are two general categories of synthesis: Nicoletti.
special F chromosomes, F fragments and marked F references: Brosseau, Nicoletti, Grell, and
chromosomes; in addition, there are marked F frag- Lindsley, 1961, Genetics 46: 339-46.
ments. F fragments are symbolized either YL or Ys properties: Meiotic behavior and viability apparently
plus necessary distinguishing notation, e.g., YS8. normal. Produces Co effect; covers N. Has, in
The F chromosome may be marked by mutating the addition, combined marker characteristics of BSY
genetically demonstrable elements of the F or by and y+Y.
translocating normal or mutant alleles from other
parts of the complement to the F chromosome.
Marked F's are symbolized by combining, without constitution: Bs pdt+ su(f)+ kz+ - dm+ KL-bb+ KS;
intervening punctuation, the symbol for the normal or inferred from origin.
mutant gene of primary marker intent with the symbol origin: X-ray-induced deletion of majority of euchro-
F. If the marker is in the long arm, its symbol pre- matin (rb+ through mal+) from a recombinant
cedes Y (e.g., y+Y) and if it is in the short arm, its carrying left end of FW element of T(1;Y)148 =
symbol follows F (e.g., Ybb). Symbols for marked T(1;Y)2D;YL, which involves BSY, and right end
fragments combine the symbol for the appropriate F of XYL-YS.
arm with that for the marker gene, listed in order. synthesis: Nicoletti.
These notations are separated by a centerpoint when references: Brosseau, Nicoletti, Grell, and
the centromere lies between them (e.g., YLtscSi), Lindsley, 1961, Genetics 46: 339-46.
otherwise they are not separated by punctuation properties: Produces Co phenotype in X/Bsw+Y
(e.g., y+YL). male and X/X/Bsw+Y female. Covers many N
The long arm of the F carries a complex of male deficiencies. B phenotype as in BSY.
fertility factors, KL, and the short arm carries a BSy+Y
normal allele of bb proximally and a complex of male constitution: Bs pdf+ su(f)+ y+ ac + KL-bb+ KS;
fertility factors, KS, distally. The genetic constitu- inferred from origin.
tion of the F chromosome may be designated by origin: X-ray-induced deletion of the euchromatin
listing the above components and the centromere in (kz + through mal*) from the recombinant composed
order from left to right, KL'bb+KS. The constitution of left end of YDXP element of T(1;Y)148 ~
of a F fragment is described in a similar manner. T(1;Y)2D;YL, which involves BSY, and right end
The genetic constitution of a marker segment is des- of XYL-YS129-16, which carries y+ from y+F be-
ignated by listing the symbols of the most widely tween X and F*1.
SPECIAL CHROMOSOMES - Y DERIVATIVES 417
synthesis: Nicoletti. synonym: scs-Y:bw+.

references: Brosseau, Nicoletti, Grell, and references: 1952, DIS 26: 97.
Lindsley, 1961, Genetics 46: 339-46. FR2: seey + F*-
Fragment 2: see y + YL
constitution: Bs pdf+ su(f)+ KL'bb* KS. KDJ1+Y
origin: X-ray-induced deletion of majority of euchro- constitution: 1(1)] 1 + KL-bb+ KS.
matin (including mal + ) from a recombinant carrying origin: Neutron-induced derivative of y+Y.
left end of 4DXP element of T(1;4)BS = T(1;4)15F9- synthesis: Muller.
16Al;16A7-Bl;102F and right end of XYL-YS. references: 1954, DIS 28: 140-43.
synthesis: Brosseau. properties: Like y+Y except that y + and a c + but
synonym: YBS. not 1(1)J1 + deleted.
references: Brosseau and Lindsley, 1958, DIS 32: R(Y)bw +
116. constitution: KL (bw+ — ba+)-bb+ KS; closed to
Brosseau, Nicoletti, Grell, and Lindsley, 1961, form a ring.
Genetics 46: 339-46. origin: X-ray-induced derivative of bw+Yy+.
properties: Causes extreme B phenotype; presence synthesis: Oster and Iyengar.
readily scorable in +/+, B/+, and + but not in B/B synonym: Ycbw+; MYR: Marked Y Ring.
or B. Shown to carry pdi+ (Grell) and su(f)+ (in references: 1955, DIS 29: 159.
BsYy+; Zimmering, 1959, DIS 33: 175-76). Does properties: Ring shaped in m it otic metaphase.
not cover spaPo1. Viability and fertility of X/BSY, Lacks y present in the treated chromosome. In-
X/B^Y/BSY, and X/X/BSY good. Three euchro- troduction of Rfif)bw+ via male into certain strains
matic bands visible in salivary chromosomes results in death of nearly all male progeny during
(Nicoletti and Lindsley, I960, Genetics 45: 1705- early embryogenesis. About 10 percent of strains
22). are subject to such killing of male offspring. Intro-
duction of R(Y)bw+ via female does not result in
constitution: Bs pdf+ su(f)+ KL-bb + KS ac + - death of the sons (Oster, 1964, Genetics 50: 274).
R(Y)L
origin: Recombination between BSY and bw+Yy+. constitution: KLm; closed to form a ring.
synthesis: Brosseau. origin: Spontaneous derivative of y+Y.
references: 1958, DIS 32: 115-116. synthesis: Muller.
Brosseau, Nicoletti, Grell, and Lindsley, 1961, synonym: Ycl; YLc.
Genetics 46: 339—46. references: 1948, DIS 22: 73-74.
properties: Ring shaped in mitotic metaphase and
about same length as the X chromosome (Hannah).
constitution: J3S pdf+ su(t)+ KL-bb+ KS ac+ yîd Carries all of KL. bb/R(Y)L is bobbed and
1(1)J1+; inferred from origin, bb*/R(Y)L dies; therefore R(Y)L lacks bb locus.
origin: X-ray-induced recombinant between BsYy +
and In(l)scs, y31d sc8 in a female. *R(Y)L14
references: Brosseau, Nicoletti, Grell, and constitution: KL", closed to form a ring.
Lindsley, 1961, Genetics 46: 339-46. origin: X-ray-induced derivative of bw+Yy+.
bw+Y synthesis: W. K. Baker.
constitution: KL (bW+ - ba+)-bb+ KS. synonym: Yc^-14.
origin: Aneuploid segregant from T(Y;2)bw+Y = references: Baker and Spofford, 1959, Univ. Texas
T(Y;2)YL;58F1-59A2;6OE3-F1 (Gersh, 1956, DIS Publ. 5914: 135-54 (fig.),
30: 115; Nicoletti). properties: Ring shaped in mitotic metaphase.
synthesis: Dempster. Lacks y + , bw+, bb+, and KS present in treated
syn on y m: y": bw +. chromosome.
references: Muller, 1942, DIS 16: 64. R(Y)L15
1951, DIS 25: 119. constitution: KL", closed to form a ring.
1955, DIS 29: 146. origin: Spontaneous derivative of bw+Yy+.
properties: A section of 2R carrying bw+ inserted synthesis: W. K. Baker.
into YL proximal to KL (Baker, 1955, DIS 29: synonym: YcL-15.
101—3). Inserted segment known to carry normal references: Baker and Spofford, 1959, Univ. Texas
alleles of bw, mr, or, Fo, Pin, bs, and 6a but not Publ. 5914: 135-54 (fig.),
px, hv, crs, M(2)l, orM(2)c. Male with two bw+Y properties: Ring shaped in mitotic metaphase.
chromosomes lethal. Lacks y+, bw+, bb+, and KS present In original
y chromosome.
constitution: KL (bw+ - fca+>66 + KS ac+ - t(l)Jl + R(Y)Lbb +
(Baker, 1955, DIS 29: 101-2). constitution: KL'bb+; closed to form a ring.
origin: Recombination between Ys of bw+Y distal to origin: X-ray-induced derivative of bw+Yy+.
KS and YL of y+Y in C(l)RM/y+Y/bw+Y female, synthesis: W. K. Baker,
synthesis: Cooper. synonym: YcL:bb+.
references: Baker and Spofford, 1959, Univ. Texas synthesis: Brosseau.

Publ. 5914: 135-54 (fig.), references: I960, Genetics 45: 257-74.
properties: A medium-sized ring in mitotic meta- properties: A series of y+Y chromosomes that carry
phase. Lacks y+, bw+, and KS present in treated a male-sterilizing mutant in KL; e.g., y+mslY
chromosome. carries ms(Y)L,l. Brosseau (1960) described a
R(Y)Sbw + number of complementing male-sterilizing mutations
constitution: (bw + — ba+)-bb+ KS; closed to form a in YL.
ring. Order of elements inferred from origin. y+Y
origin: X-ray-induced derivative of bw+Yy+. constitution: l(l)Jl + - ac+ KL-bb+ KS.
synthesis: W. K. Baker, origin: X ray induced in spermatogonial cell of
synonym: YcS:bw+bb+. In(l)sc8/Y male.
references: Baker and Spofford, 1959, Univ. Texas synthesis* Muller.
Publ. 5914: 135-54 (fig.). synonym: sc^'Y.
properties: Ring shaped in mitotic metaphase. references: 1948, DIS 22: 73-74.
Lacks y + and KL from treated chromosome. Shows properties: Tip of In(l)scs including 1(1)J1+, y+,
some somatic and germinal instability. and ac+ but not sc transferred to tip of YL distal
SC'YL: seeYL-scsl to KL. Since in metaphase YL appears to be as
scSENc.o. Y B-2: see Y^-y + B2 long as X (Hannah), some heterochromatin derived
scSENc.o. Y T-0: see Y^-y + TO from In(l)sc8 must be carried distally on YL. De-
sc8-Y: see y+Y tachment studies show that bb+ from In(l)sc8 has
sc8-Y:bw+: see bw+Yy+ not been transferred to YL (Parker). Has dominant
see yvssy Hw effect that produces one or more humeral hairs
.o. Y EY80: see Y in X/y+Y male and X/X/y+Y female and one or
more hairs in second and third posterior cells of
constitution: 1(1)J1 + - scVl-bb+ KS; tentative. wing of X/y+Y/y+Y male (Schultz).
origin: Spontaneous recombinant from
In(lLR)scVl/Y male. y + Yms
synthesis: Muller. constitution: l(l)Jl + ~ ac+ KL-bb+ ms(Y)S.
references: 1948, DIS 22: 73-74. origin: X-ray-induced derivative of y+Y.
properties: Small two-armed chromosome in mitotic synthesis: Brosseau.
metaphase. Survives in combination with C(1)DX references: 1960, Genetics 45: 257-74.
and therefore probably carries bb+ and the nucle- properties: A series of y+Y chromosomes that carry
olus organizer. a male-sterilizing mutant in KS; e.g., y+Yms2
Tp4-Y: see 4Y carries ms(Y)S2. Complementing male-sterilizing
Transposition 4-Y: see 4Y mutations in Ys have been described by Brosseau
w+y+Y (1960).
constitution: kz + - spl + y + ac + KL-bb+ KS; y + YL
inferred from origin and supposed constitution of constitution: 1(1)J1 + - ac+ KL-bb+.
Bsw+y+Y. origin: Spontaneous product from sc cv v f/YsX'YL,
origin: X-ray-induced derivative of Bsw+y+Y. In(l)EN, KS y-KL y+ female.
synthesis: Nicoletti. synthesis: Novitski.
references: Brosseau, Nicoletti, Grell, and synonym: FR2: Fragment 2.
Lindsley, 1961, Genetics 46: 339-46. references: 1952, Genetics 37: 270—87.
properties: Like Bsw+y+Y but with stiff)+ and Bs properties: Has subterminal centromere and ex-
missing. tremely short second arm in mitotic metaphase.
w+Y Constitution confirmed by analysis of detachments
constitution: kz+ — spl + KL-bb+ KS; inferred from with C(1)RAI all of which appear to result from ex-
origin. Also associated with T(Y;2)w+Y = change between the intersitial heterochromatin of
T(Y;2)YL;22D (Schultz) in which the break in YL the compound and the fc6+-bearing short arm of
is distal to the kz + — spl insertion. y+Y1* (Sandier, 1954, DIS 28: 153-54).
origin: Spontaneous in C(l)RAYL/w+y+Y female. yS3!Y
Seems likely that the w+y+Y, which was an X-ray- con#itution: 1(1)11+ y53i ac+ KL-bb+ KS.
induced derivative of Bsw+y+Y, was different from origin: X-ray-induced derivative of y+Y.
the one described here and was already trans- synthesis: Luning, 53i.
located with chromosome 2. references: 1953, DIS 27: 58.
synthesis: Nicoletti. properties: Like y+Y but with a mutant allele of y.
references: Brosseau, Nicoletti, Grell, and
Lindsley, 1961, Genetics 46: 339-46. constitution: 1(1)J1+ yP59 ac+ KL-bb+ KS.
properties: Produces Co effect; covers N. origin: Spontaneous derivative of y+Y.
y+msY synthesis: Perkovic, 59h.
constitution: l(l)Jl + ~ ac + ms(Y)L'bb+ KS. references: Meyer, 1959, DIS 33: 97.
origin: X-ray-induced derivative of y+Y. properties: Like y+Y but with a y 2 -like allele of y.
SPECIAL CHROMOSOMES - Y DERIVATIVES 419
*yvS6Y synthesis: Crew and Lamy.

constitution: 1(1)J1+ yvS6 a c + KL'bb+ KS. synonym: SC'YL.
origin: X-ray-induced derivative of y+Y. references: 1940, J. Genet. 39: 273-83.
synthesis: C. Hint on and Schmidt. Pontecorvo, 1940, DIS 13: 74.
synonym: scav-Y. properties: Described as an acrocentric rod the size
references: 1956, DIS 30: 121. of Y1-1 in meta phase (Crew and Lamy, 1940). Later
properties: Like y+Y but with strong variegation for described as an asymmetrically V-shaped element
y. May be associated with unanalyzed rearrange- with the shorter arm the size of YL (Pontecorvo,
ment. 1940).
YB9: see fisy
Ybb constitution: KL-bb+ sc^l _ 1(1)J1+; presence of
constitution: KL-bbKS. 66 + tentative,
origin: Spontaneous. origin: Recombinant between Ys and distal hetero-
synthesis: Bridges, 1926. chromatin of In(l)scsi in Base male.
properties: General symbol for a Y chromosome synthesis: Parker,
carrying a mutant allele of 66, e.g., bbY. Such Y synonym: scsl'YL*2.
chromosomes occasionally encountered in crosses references: Parker and McCrone, 1958, Genetics 43:
of 66 + lines to 66. 172-86.
Ybb-
constitution: KL'KS.
synthesis: Schultz, 33k8. constitution: jRX'66 + scsi - I(1)J1+.
properties: A Y chromosome deficient for a section origin: Recombinant between Ys and distal hetero-
of Ys including 66+ but not KS. X/Ybb~ male fer- chromatin of In(l)scSl in Base male.
tile. Ys about one-third normal length in meta- synthesis: Lindsley.
phase. synonym: scsic.o. Y EY80.
Y:bw + : see bw+Y properties: Resembles a normal Y in mitotic pro-
*Ymal + phase.
constitution: KL*sw + — su(f)+ bb+ KS; inferred from Yi-.Y+B2
origin. constitution: KL-ac+ - 1(1)] 1 +.
origin: X-ray-induced deletion of majority of euchro- origin: Recombinant between Ys and distal hetero-
matin [l(l)Jl+ through car+] from YSX-YL, chromatin of In(l)sc8LENR.
In(l)EN. synthesis: Lindsley.
synthesis: E. H. Grell. synonym: scaENc.o. Y B-2.
references: Brosseau, Nicoletti, Grell, and references: 1955, Genetics 40: 24—44.
Lindsley, 1961, Genetics 46: 339-46. properties: Lethal in combination with
Ymal+2 In(l)sc*LscaR or bb1; bobbed in combination with
constitution: KL'Sw+ — su(f)+ bb+ KS; inferred from 66. Unique among such recombinants between Ys
origin. and distal heterochromatin of In(l)sc8 or In(l)scsi
origin: X-ray-induced deletion of majority of euchro- in lacking 66 + . Resembles normal Y in mitotic pro-
matin [l(l)Jl+ through car+] from YSX-YL, phase.
In(l)EN.
synthesis: £. H. Grell. YL.y+TO
references: Brosseau, Nicoletti, Grell, and constitution: KL'bb+ ac + - 1(1)J1+.
Lindsley, 1961, Genetics 46: 339-46. origin: Recombinant between Ys and distal hetero-
Ycbw+: see R(Y)hw + chromatin of In(l)sc*Z*ENR.
Yd-, see R(Y)L synthesis: Lindsley.
: seeR(Y)Lbb+ synonym: sc8ENco. Y T-0.
: see R(Y)Sbw + references: 1955, Genetics 40: 24—44.
YM3 properties: Resembles a normal Y in mitotic pro-
constitution: KL. phase.
origin: Spontaneous derivative of bw+Yy+. other information: One of a number of similar recom-
synthesis: W. K. Baker. binant Y chromoeomes recovered from In(l)sca or
references: Baker and Spofford, 1959, Univ. Texas In(l)sc8ÊNR males.
Publ. 5914: 135-54 (fig.). YL.y3M
properties: A large acrocentric chromosome in mi- constitution: K!JL-6&+ scsi - y3M 1(1}J1+; inferred
totic metaphase. Lacks y+, bw\ bb\ and KS from supposed constitution of Y^"scsi.
present in original chromosome. origin: Spontaneous mutation in YLtscS1.
YL. SC S? synthesis: Muller.
constitution: KL'bb* s c s * -i(i)ji+; presence of references: Muller and Valencia, 1947, DIS 21: 70.
66 + tentative, properties: Like YL'SCS1 but with y^-like allele
origin: Recombinant between Ys and distal hetero- of y.
matin of Jn(l)scSl. R(Y)L
YSy
constitution: -66 + KS; tentative. constitution: -bb+ KS ac + - 1(1)J1+; tentative.
origin: Spontaneous derivative of bw+Yy + recovered origin: X-ray-induced derivative of bw+Yy+.
from R(l)l/bw+Yy+ male. synthesis: W. K. Baker.
synthesis: W. K. Baker, synonym: F s :y + fc6 + -7.
synonym: Ys:bb+-8. references: Baker and Spofford, 1959, Univ. Texas
references: Baker and Spofford, 1959, Univ. Texas Publ. 5914: 135-54 (fig.),
Publ. 5914: 135-54 (fig.). properties: A rod-shaped chromosome about twice
properties: A small two-armed chromosome in mi- the length of chromosome 4 in mitotic metaphase.
totic metaphase. Lacks y+, bw+, and KL present Lacks bw+ and KL present in treated chromosome.
in chromosome of origin.
Y9:bb+-8: see constitution: KS bb+-bb+ KS; tentative.

YSy+5 synthesis: Stern.
constitution: -bb+ KS ac +- l(l)Jl +; tentative. synonym: F ".
origin: Spontaneous derivative of bw*Yy + recovered references: 1929, Z. Induktive Abstammungs-
from R(l)l/bw+Yy+ male. Vererbungslehre 51: 253-353.
synthesis: W. K. Baker, properties: V-shaped chromosome in mitotic meta-
synonym: Y$:y*bb -5. phase with both arms the length of Ys.
references: Baker and Spofford, 1959, Univ. Texas
Publ. 5914: 135-54 (fig.). constitution: KS bb + -bb+ KS; inferred from probable
properties: Large acrocentric chromosome in mitotic mode of origin.
metaphase. Lacks bw+ and KL present in original origin: Spontaneous product from X'Ys/y+Y male.
chromos ome. synthesis: Muller.
YSy + 6 properties: V-shaped chromosome in mitotic meta-
constitution: -bb+ KS ac+ - 1(1)J1+-, tentative. phase with both arms the length of Ys.
origin: X-ray-induced derivative of bw+Yy+.
synthesis: W. K. Baker. constitution: KS bb+'bb+ KS; inferred from probable
synonym: Ys:y+bb*-6. mode of origin.
references: Baker and Spofford, 1959, Univ. Texas origin: Spontaneous product from X-Ys/y+Y male.
Publ. 5914: 135-54 (fig.). synthesis: Muller.
properties: Two-armed chromosome in mitotic meta- references: 1948, DIS 22: 73-74.
phase. Lacks bw+ and KL present in treated chro- properties: Like YS-YS2.
mosome. Y": see Y*-YS
CYTOLOGICAL MARKERS
7: Chromosome 7 4: Chromosome 4
See X, this subsection. In mitotic configurations chromosome 4 is a dot-
2: Chromosome 2 like element that is separated into two segments of
In mitotic figures, chromosome 2 is less than grossly unequal size by a sometimes visible centric
twice the length of the X and slightly smaller than constriction. Claimed to lack heterochromatic mate-
chromosome 3. It is a V-shaped element with two rial, but can be involved in rearrangements that pro-
centrally located heterochromatic segments presum- duce variegated position effect [e.g., T(l;4)wmS =
ably separated by the centric constriction; the heter- T(l;4)3C3-4;101Fl-2\, and shows incorporation of
ochromatic segments are late replicating, according tritiated thymidine in cells in which only heterochro-
to thymidine incorporation studies (Barigozzi, matic regions of the other chromosomes are labeled
Dolfini, Fraccaro, Raimondi, and Tiepolo, 1966, (Barigozzi, Dolfini, Fraccaro, Raimondi, and
Exptl. Cell Res. 43: 231-34). In early prophase Tiepolo, 1966, Exptl. Cell Res. 43: 231-34). In
there is often a long achromatic gap separating the salivary gland chromosomes, the longer right arm is
euchromatic portion of one arm from the heterochro- associated with the chromocenter. The shorter left
matin. Kaufmann (1934, J. Morphol. 56: 125-55) re- arm is occasionally discernible within the chromo-
ported the gap to be in 2L. center; it has been demonstrated genetically by
Hinton (1942, Genetics 27; 119-27) stated that trans locations between it and the X chromosome;
both the constriction and the centromere are located e.g., T(l;4)w""l* (Panshin and Khvostova, 1938,
in the region between the chromosome 2 breakpoints Biol. Zh. (Moscow) 7: 359-80) and T(l;4)wmA
of T(1;2)N264-59 = T(l;2)3C8-9;40F and (Griffen and Stone, 1940, Texas Univ. Publ. 4032:
T(l;2^j264-23 = T(1;2)3C8-9;41A, a segment con- 201-7).
taining one or two bands but comprising about 15 4L: Left arm of chromosome 4
percent of the metaphase length of 2. See 4, this subsection.
4R: Right arm of chromosome 4
2L: Left arm of chromosome 2 See 4, this subsection,
See 2, this subsection. 4-s'im: Chromosome 4 from Drosophila simulans
2R: Right arm of chromosome 2 Chromosome 4 of D. simulans was introduced into
See 2, this subsection. an otherwise D. melanogaster genome by Muller and
3: Chromosome 3 Pontecorvo (1940, Nature 146: 199-200). Pheno-
A V-shaped element in mitotic figures that is typic effects of 4-sim were described by Muller and
slightly larger than chromosome 2 but less than Pontecorvo (1942, Genetics 27: 157) and Pontecorvo
twice the length of the X chromosome. In prophase, (1943, Proc. Roy. Soc. Edinburg B 61: 385-97,
there are two medial heterochromatic segments sepa- 1943, J. Genet. 45: 51—66). 4 sim/4 apparently
rated by a constriction that presumably marks the normal in phenotype; 4~&im/ci has occasional wing
position of the centromere; these segments comprise vein interruption; 4-sim/dw has more extreme d
about the proximal one-fifth of each arm at meta- phenotype than 4/rt* (Uphoff, 1949, Genetics 34:
phase, and according to tritiated thyraidine incorpo- 351—27); no dominance of spaCat in 4-simfspaCat.
ration studies (Barigozzi, Dolfini, Fraccaro, Homozygous 4-sim/ 4-aim has fair viability with
Raimondi, and Tiepolo, 1966, Exptl. Cell Res. 43: slight morphological peculiarities, e.g., body flat-
231—34), they are late replicating. tened, trident heavy, and eyes reduced. Male geni-
3L: Left arm of chromosome 3 talia said to be a little like those of D. simulans.
See 3, this subsection. Homozygous female fertile, but male sterile, Testes
3R: Right arm of chromosome 3 well developed; meiosis occurs but no motile sperm
See 3, this subsection. are produced. 4-sim/M(4) is Minute and male sterile;
421
4-sim/M(4) is not Minute but is male sterile. Muller hB

and Pontecorvo (1942) suggest a gene necessary for The second from the centromere of four segments
male fertility of D. melanogaster is deleted from discernible in the proximal heterochromatin of XL
Df(4)M and absent from 4-sim. (see X, this subsection; also Cooper, 1959, Chromo-
Comparisons of melanogaster and simulans sali- soma 10: 535—88).
vary chromosomes were published by Kerkis (1936, hC
Am. Naturalist 70: 81-86), Horton (1939, Genetics The third from the centromere of four segments
24: 234-43), and Patau (1935, Naturwissenschaften discernible in the proximal heterochromatin of XL
23: 537-43). According to Horton (1939) 4-sim (see X, this subsection; also Cooper, 1959, Chromo-
differs from 4 by a relatively long inversion, which soma 10: 535-88).
includes at least from 102B1-2 through 102E1-2 and hD
probably through 102E3, 4 and 5. A darkly-staining The distalmost of four segments discernible in the
terminal ring is at the tip of the chromosome. proximal heterochromatin of XL (see X, this subsec-
Slizynski (1941, Proc. Roy. Soc. Edinburg B 61: tion; also Cooper, 1959, Chromosoma 10: 535—38).
95—106) identified the short left arm of 4-sim; it LA
lacks a dark band present in the middle of the arm in The proximalmost of three discernible segments of
D, melanogaster. In melanogaster-simulans hybrids YL (see Y, this subsection; also Cooper, 1959,
the fourth chromosomes do not pair in salivary gland Chromosoma 10: 535—88).
cells; however, Slizynski found one nucleus of LB
sim-4/4 in a melanogaster background in which the The middle of three discernible segments of YL
inverted segment was paired. In triplo-4 larvae with (see Y, this subsection; also Cooper, 1959, Chromo-
one 4-sim, the melanogaster 4's are paired and the soma 10: 535—88).
4-sim is unpaired. 4 tends to have the tip of 4R
LC
stuck into the chromocenter; 4-sim, however, always
The distalmost of three discernible segments of
has its tip free of the chromocenter.
YL (see Y, this subsection; also Cooper, 1959,
Chromosoma 10: 535—88).
BkA: Block A
Muller, Raffel, Gershenson, and Prokofyeva- NO: Nucleolus Organizer
Belgovskaya (1937, Genetics 22: 87-93) defined it The region in the proximal heterochromatin of the
as the distal and major portion of the segment of X and the short arm of the Y chromosome where the
proximal heterochromatin of the X chromosome lo- nucleolus is organized. The nucleolus is visible in
cated between the right breakpoints of In(l)scL8 and interphase, and its relation to the NO may be seen
In(l)sc8. This region was considered to be a unit or in early prophase; it may also be seen associated
the product of a single genetic unit not subject to X- with the chromocenter in salivary gland prepara-
ray-induced breakage. Subsequent work by Kaufmann tions. The nucleolus is formed at the constriction
(1954. In Radiation Biology, A. Hollaender, ed., between hB and AC in the proximal heterochromatin
McGraw-Hill, Inc., Vol. 1, pp. 627-711), and in par- of the X chromosome and at the constriction one-
ticular that of Cooper and Krivshenko with Dp(l;{) third the way from the centromere along the short
derivatives, suggests that breakability in Xh is dis- arm of the Y. Homozygous deficiency for the nucle-
tributed uniformly over its mitotic length; therefore olus organizer is lethal. Ritossa and Spiegelman
the block concept of heterochromatic structure no (1965, Proc. Natl. Acad. Sci. U.S. 53: 737-45)
longer seems valid. showed that the amount of DNA complementary to
ribosomal RNA in a cell is directly related to the
BkB: Block B number of nucleolus-organizing regions present;
Muller, Raffel, Gershenson, and Prokofyeva- they believe that the NO is the chromosomal site of
Belgovskaya (1937, Genetics 22: 87-93) claimed it ribosomal RNA synthesis. The ribosomal RNA-com-
to be proximal to BkA in the region of Xh between plementary DNA comprises 0.27 percent of the total
the right breakpoints of In(l)scL8 and In(l)sc8. DNA of a haploid genome; on the basis of the amount
Gershenson [l940, Vid. Akad. Nauk SSSR., (Kiev) of DNA per cell and the molecular weight of ribo-
3-116] defined it as the segment of the proximal somal RNA in Drosophila, they have calculated that
heterochromatin of XL to the right of the right break- the amount of ribosomal RNA-complementary DNA in
point of In(l)sc8, claimed to comprise 20—30 percent a haploid genome is sufficient to synthesize 130
of Xh, although, according to Cooper (1959, Chromo- molecules each of 18S and 28S ribosomal RNA simul-
soma 10: 535—88), a much smaller proportion of Xh taneously. Ritossa, Atwood, and Spiegelman (1966,
lies between the right breakpoint of In(l)sc8 and the Genetics 54: 819-34) postulated that the NO is the
centromere. Existence of BkB subject to same cytological counterpart of the bb locus, on the basis
doubts as that of BkA, of the demonstration that replacing a normal X with
hA an X carrying 66 reduces the amount of ribosomal-
The proximalmost of four segments discernible in RNA-complementary DNA per cell.
the proximal heterochromatin of XL (see X, this sub- Puffs
section; also Cooper, 1959, Chromosome 10: 535— Localized swellings in polytene chromosomes
88). marking regions of metabolic activity. They are
CYTOLOGICAL MARKERS 423
found in specific regions of the polytene comple- Becker Schultz

ment, and each puff has characteristic times of ap- 44E
pearance and disappearance during development, 46F
which have been described by Becker (1959, Chromo- 47A 47AB
some 10: 654—78) for the puffs in 3L and the distal 47BC
part of XL in the salivary glands. Studies with other 48B 48A
diptera show that the puffing patterns in the polytene 49EF
chromosomes of different tissues are not identical; 50C 50CD
Becker (1959) describes one puff in region 15BC of 50D
Drosophila melanogaster that is present in the an- SIDE
terior but not the posterior portion of the salivary 52BC
gland. The puffing pattern responds to changes in 53-54
cellular environment; e.g., changes in hormonal con- 55B
centration (Becker, 1962, Chromosoma 13: 341—84). 55E
Becker (1959) and Schultz catalogued the regions of 56D 56DE
the salivary gland chromosomes of Drosophila mela- 58BC 58A
nogaster in which puffs may be seen (see following 58F 58DE
tabulation). 59F 60A
60B 60B
Becker Schulfcz 62E 62B-E
1C 1C 63E 63C
2B5-6 2B
2B13-17 63F 64B
2EF 2F 66B
3AB 3A 66B 66DE
3C11-12
3E 3DE 67B 67E
4EF 68BC
7AB 68C 68-69
8B 70C
9B 70C 71DE
9EF 71C-E 72CD
10 10EF 72CD 73C
11B 11BC 74EF 74EF
12E 12-13 75B 75AB
13B 76A
14EF 76D
15BC 15C 78D 78DE
16BC 16A 79DE
16DE 16F 82CD 82BC
21F 21DE 82EF 82EF
22B 22A 83AB 83C
22C 22CD 83EF
23C 23BC 84 BC
23DE 85B
25A-C 25BC 85CD 85D
25D 85EF 85EF
26B 86-87
27-28 88D 88D
29-30 88EF 88EF
33B 33B 89B 89BC
33E 33E 90BC 90C
34A 34A 92A
34C 93B
35AB 93D
36F 36EF 94
37 37B 95F
38B 96E
38F 39B 97BC 97B
40B 98B 98B
42A 42AB 98F 99B
43E 43-44 99EF
SA
The proximal and smaller of two discernible seg-
ments of Ys (see Y, this subsection; also Cooper,
1959, Chromosoma 10: 535—88). LC LB LA SA SB
SB
The distal and larger of two discernible segments
of Ys (see Y, this subsection; also Cooper, 1959,
t t
ceniromere NO
Chromosoma 10: 535-88).
•XL- -\XR Y: Y chromosome

Redrawn from Cooper, 1959, Chromosoma 10: 535—88.
-Xh- —I
hD hC hB hA Y: Y chromosome
•NBA* In mitotic figures the Y chromosome appears as an
f t entirely heterochromatic element; tritiated thymidine
NO centromere
studies (Barigozzi, Dolfini, Fraccaro, Raimondi, and
X: X chromosome Tiepolo, 1966, Exptl. Cell Ress. 43: 231-34) show
Redrawn from Cooper, 1959, Chromosoma 10: 535-88. it to be late replicating. The Y is a two-armed chro-
mosome, Ys being about two-thirds the length of YL.
X: X chromosome At metaphase, the Y is usually somewhat shorter
Also known as chromosome 2. Present in one dose than the X chromosome. The position of the centro-
in male and two doses in female. In mitotic figures mere is indicated by a constriction. The short arm
the X is virtually a rod-shaped element with a quasi- is subdivided by the constriction associated with
terminal centromere and a minute second arm desig- the nucleolus organizer into a distal segment, SB,
nated as the right arm, XR. The left arm, XL, is about two-thirds and a proximal segment, SA, one-
divided into a distal euchromatic or isopycnotic third the length of Ys. In early prophase the nucle-
region, Xe, in which the chromatids are usually sep- olus may sometimes separate SA from SB by a con-
arated and a proximal heterochromatic or heteropyc- siderable distance. YL is divided into three seg-
notic region, Xh, in which the chromatids are not ments of about equal size by a pair of constrictions;
separated. The relative lengths of these subdivi- the more distal of which is the more constant land-
sions of XL vary according to mitotic stage; in early mark. From the centromere the segments are desig-
prophase the isopycnotic region is longer, but by nated LA, LB, and LC (Cooper, 1959, Chromosoma
metaphase the two regions are of about equal length. 10: 535—88). In salivary gland preparations,
Tritiated thymidine incorporation studies (Barigozzi, Prokofyeva-Belgovskaya (1937, Genetics 22: 94-
Dolfini, Fraccaro, Raimondi, and Tiepolo, 1966, 103) observed a small collection of bands that she
Exptl. Cell Res. 43: 231-34) demonstrate that the attributed to the Y. Nicoletti and Lindsley (1960,
heterochromatic region is late replicating. The prox- Genetics 45: 1705—22) found no evidence of bands
imal heterochromatin of XL is subdivided by con- attributable to the Y chromosome in a study of
strictions into four segments of about equal length; T(l;Y)'s. A series of complex structural elements in
these segments are designated from proximal to primary spermatocyte nuclei, whose formation de-
distal hA, hB, hC, and hD (Cooper, 1959, Chromo- pends on the presence of the Y, have been postu-
soma 10: 535—88). The constriction between hB and lated to represent a highly modified state of the Y
hC marks the position of the nucleolus organizer, chromosome (Meyer, Hess, and Beerman, 1961, Chro-
NO; in early prophase it may be a very long gap oc- mosoma 12: 676—716). The Y chromosome carries a
cupied by the nucleolus. The polytene X consists of normal allele of bb and two complexes of factors es-
just over 1000 bands of which 25—30 correspond to sential to male fertility; KL is on YL and composed
the region that is heteropycnotic in the mitotic X. of five cistrons, kll-kl5; KS is on Ys and composed
The length of Xh is large compared with that of Xe of two, ksl and k&2. Brosseau (1960, Genetics 45:
when measured at metaphase or estimated by relative 257—74) showed the order of genetically demon-
frequency of involvement in X-ray-induced rearrange- strable factors on the Y to be A/5 kl4 kl3 kl2 Ml •
ments, but small when measured in salivary-gland bb+ ksl ks2. Addition of F's to a normal chromo-
chromosomes or by crossing over. Most sex-linked some complement suppresses variegated position ef-
genes are in Xe, only the locus of bb and possibly fects, and removal of the Y from the male enhances
that of su(f) being in Xh. variegation (1933, Gowen and Gay, Proc. Natl.
X e : euchromatin of X chromosome Acad. Sci. U.S. 19: 122-26). Two or more Y's
See X, this subsection. added to the normal complement cause variegation of
Xh: heterochromatin of X chromosome otherwise self-colored eyes (Cooper, 1956, Genetics
See X, this subsection. 41: 242-64).
X L ; Left arm of X chromosome YL: Long arm of Y chromosome
See X, this subsection. See Y, this subsection.
XR: Right arm of X chromosome Ys: Sfiorf arm of Y chromosome
See X, this subsection. See Y, this subsection.
DEPARTURES FROM DIPLOIDY
The diploid chromosome complement of Drosophila slash bars and nonhomologous chromosomes by semi-
melanogaster may be designated X/X; 2/2; 3/3; 4/4 colons. When two homologous chromosomes are at-
for females and X/Y; 2/2; 3/3; 4/4 for males. Addi- tached to the same centromere, components are
tion to or subtraction from either of these comple- listed without separation, e.g., XX, XY, and 44.
ments of one or more chromosomes produces a depar- diploid metafemale
ture from diploidy. The non-diploid constitutions constitution: X/X/X; 2/2; 3/3; 4/4; sex chromosome
are designated by a name but not a symbol except as constitution may also be XX/X.
included in the name, e.g., X/0 male. Constitutions source: Produced by triploid and compound-X-bear-
are described by listing their component chromo- ing females. May result from two-X gametes pro-
somes, homologous chromosomes being separated by duced by nondisjunction.
discoverer: Bridges,
synonym: superfemale.
references: 1921, Science 54: 252—54.
1922, Am. Naturalist 56: 51-63 (fig.).
1925, Am. Naturalist 59: 127-37.
Morgan, Bridges, and Sturtevant, 1925, Bibliog.
Genet. 2: 153-62 (fig.).
properties: Wings crumpled or incised on inner
margin. Rear legs often malformed. Viability low,
usually less than 0.5 percent. Flies die mostly in
late larval and pupal stages; at 25°C, puparium
formation delayed 1—2 days (Brehme, 1937, Proc.
Soc. Exptl. Biol. Med. 37: 578-80). Survivors
sterile; two fertile metafemales were apparently
mosaic for triploid tissue [Rolfes and Hollander,
1961, J. Heredity 52: 61-66 (fig.)]. Larval ovaries
from metafemales transplanted into sterile diploids
have produced a few progeny (Beadle and Ephrussi,
1937, Proc. Natl. Acad. Sci. U.S. 23: 356-60).
Crossing over between the X chromosomes appears
to be infrequent.
other information: The term metafemale instead of
diploid metafemale superfemale was suggested by Stern (1959r Lancet
From Bridges, V922, Am. Naturalist 56: 51-63. 12: 1088).
425
discoverer: Bridges, 20/.

references: 1921, Science 54: 252—54.
Genet. 2: 153-62 (fig.).
Bridges, 1939. In Sex and Internal Secretions,
E. Allen, C. H. Danforth, and C. A. Doisy, eds.
The Williams and Wilkins Co. pp. 15-63.
properties: Large-bodied fly with coarse bristles,
roughish eyes, and scalloped wing margins. Small
hairs on surface of wing more sparsely distributed
than in diploids. Usually has sex combs and a
mixture of male and female genitalia; genitalia may
be malelike or femalelike. Addition of sections of
X chromosome shifts intersexes toward femaleness
[Dobzhansky and Schultz, 1934, J. Genet. 28:
349-86 (fig.); Pipkin, 1940, Univ. Texas Publ.
4032: 126—56]. Addition of sections of the second
or the third chromosome has not resulted in a shift
in sexuality (Pipkin, 1947, Genetics 32: 592-607;
1960, Genetics 45: 1205-16). Fung and Gowen
reported that a triploid line producing intersexes
hap Io-4 with preponderantly female genitalia carries sev-
From Bridges, 1922, Am. Naturalist 56: 51-63. eral fourth chromosomes and another triploid line
producing malelike intersexes carries only two
haplo->4 fourth chromosomes.
constitution: X/X; 2/2; 3/3; 4; sex chromosome
constitution may also be X/Y.
source: Produced after occasional loss or nondis-
junction of chromosome 4 during meiosis. Produced
in quantity from crosses of C(4)RM/0 with normal,
or from heterozygous T(2;4) or T(3;4) females.
discoverer: Bridges, 2Oa30.
references: 1921, Proc. Natl. Acad. Sci. U.S. 7:
186-92.
<Stenet..2: 135-43 (fig.).
properties: Minute phenotype caused by deficiency
for M(4). Pale body with prominent trident pattern
on thorax. L5 often does not reach wing margin.
Eclosion delayed 2—4 days. Viability erratic, usu-
ally below 80 percent of normal. Usually sterile.
Male tends to be more viable and fertile than fe-
male. metamale
From Bridges, 1922, Am. Naturalist 56: 51-63.
haploid
constitution: X; 2; 3; 4. metamale
source: Recorded as patches of tissue. constitution: X/Y; 2/2/2; 3/3/3; 4/4/4; inferred
discoverer: Bridges. from markers inherited. May also be diplo-4.
references: 1925, Proc. Natl. Acad. Sci. U.S. 11: source: Occurs among progeny of triploid female.
706-10. discoverer: Bridges, 201.
1930, Science 72: 405-6. synonym: supermale.
properties: Eye facets small in haploid patches. A references: 1921, Science 54: 252—54.
haploid foreleg bore no sex comb; the tissue is 1922, Am. Naturalist 56: 51-63 (fig.).
therefore probably female, as expected on basis of Morgan, Bridges, and Sturtevant, 1925, Bibliog.
balance theory of sex determination. Genet. 2: 153-62 (fig.).
intersex properties: Male has small body and spread wings.
constitution: X/X; 2/2/2; 3/3/3; 4/4/4; presence of Late hatching, poorly viable, and completely
Y and number of fourth chromosomes variable. sterile.
source: Regularly found among progeny of triploid
females. constitution: Y/Y; 2/2; 3/3; 4/4.
DEPARTURES FROM DIPLOIDY 427
source: One-fourth the progeny from crosses be- progeny of crosses between males and females that
tween certain compound-X-bearing females (e.g., carry C(4)RM (E. B. Lewis).
C(1)RM/Y) and normal males. tetraploid
properties: Dies as embryo (Li, 1927, Genetics 12: constitution: X/X/X/X; 2/2/2/2; 3/3/3/3;
1—58). Cleavage nuclei abnormally distributed and 4/4/4/4.
blastoderm not formed, according to Poulson (1940, source: Seen on a few occasions as a tetraploid
J. Exptl. Zool. 83: 271—325). According to Scriba daughter of a triploid female or as a patch of tetra-
(1964, Zool. Jahrb. Abt. Anat. Ontog. Tiere 81: ploid gonial tissue in an otherwise diploid female.
435—90), migration of cleavage nuclei to surface of Extensive attempts to produce tetraploid males
egg is normal, blastoderm formation irregular, and have failed.
germ band development frequently incomplete. discoverer: Bridges.
references: 1925, Am. Naturalist 59: 127-37.
nullo-X nullo-Y Morgan, 1925, Genetics 10: 148-78.
constitution: 2/2; 3/3; 4/4. properties: Recognized by production of progeny
source: One-fourth the progeny of crosses such as that are almost exclusively triploids and inter-
C(l)RM/0 females with YsX-YL/0 males. sexes.
properties: Most embryos die after 10—12 cleavages triplo-4
(von Borstel and Rekemeyer, 1958, Nature 181: constitution: X/X; 2/2; 3/3; 4/4/4. Sex chromo-
1597-98). Embryology like that of nullo-X (Scriba, some constitution may be X/Y; that for chromo-
1964, Zool. Jahrb. Abt. Anat. Ontog. Tiere 81: some 4 may be 44/4.
435-90). source: Product of nondisjunction of chromosome 4.
superfemale: see diploid metafemale Regular product of cross between C(4)RM and
supermale: see metamale normal diplo-4 flies.
discoverer: Bridges, 21bl3.
references: 1922, Am. Naturalist 56: 51-63.
Genet. 2: 21 (fig.), 135-43.
properties: Phenotypic departure from normal very
slight. Body darker than normal and trident pattern
subdued. Eyes small. Body and wings elongate.
Preferential segregation of the different fourth
chromosomes in triplo-4's described by Sturtevant
(1936, Genetics 21: 444-66).
triploid
constitution: X/X/X; 2/2/2; 3/3/3; 4/4/4. Sex
chromosome constitution may also be X/X/X/Y,
XX/X, or XX/X/Y. Triploids from stocks kept for
several generations usually carry only two fourth
chromosomes, i.e., diplo-4 triploids.
source: Spontaneous from unreduced eggs; incidence
increased by treatment with cold (Bauer, 1946,
tetra-4 Z. Naturforsch. 1: 35-38; Gloor, 1950, DIS 24: 82)
above: tetra-4; below: diplo-4 or with colchicine (Braungart and Ott, 1942, Sci.
From Grell, 1961, Genetics 46: 1173-83. Counselor 8: 105; Schultz). Produced in relatively
high frequency by triploid females and by
tefro-4 c(3)G/c(3)G females (Gowen, 1933, J. Exptl. Zool.
constitution: X/X; 2/2; 3/3; 4/4/4/4. Sex chromo- 65: 83-106).
some constitution may also be X/Y; that for chro- discoverer: Bridges, 1920.
mosome 4 may be 44/4/4 or 44/44. references: 1921, Science 54: 252-54.
references: Morgan, Bridges, and Sturtevant, 1925, 1922, Am. Naturalist 56: 51-63 (fig.).
Bibliog. Genet. 2: 135-43. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
Li, 1927, Genetics 12: 1-58. Genet. 2: 135-43.
Bridges, 1935, Tr. Dinam. Razvit. 10: 463-74. properties: Eye facets larger and hairs on surface of
Grell, 1961, Genetics 46: 1177-83 (fig.). wings more sparsely distributed than in diploid,
properties: Viability reduced; usually dies in em- giving eyes and wings a coarse texture; bristles
bryonic or larval stage. Wings of survivors longer also coarse. These characteristics are diagnostic
and more pointed than normal. for three sets of autosomes and result from in-
source: Synthesized as females homozygous for creased cell size. Body thickset. Ventral bristles
T(1;4)W"4 + T(1;4)BS formed by recombination in between first and second pairs of legs often miss-
region 3C4-15F8 between T(l;4)w™5 = T(1;4)3C3- ing. Discernible from diploid with practice. Fer-
4;10IFl-2 and T(1;4)BS = T(1;4)15F9-16A1;16A7- tility poor owing to production of aneuplold
B1;IO2F (Grell, 1961). Also recovered among classes of gametes. Since, during first meiotic
division equal numbers of chromosomes tend to go XXY female

to each pole, euploid gametes are produced with constitution: X/X/Y; 2/2; 3/3; 4/4. Sex chromo-
lower than expected frequencies; also gametes some constitution may also be X/XY or XX/Y.
with one X and two sets of autosomes and with two source: Product of either primary or secondary non-
X's and one set of autosomes far outnumber those disjunction in either male or female. Also pro-
with one X and one set of autosomes or two X's duced from cross of an XF-bearing parent with a
and two sets of autosomes (Bridges and Anderson, normal-X-bearing parent. Condition usually found
1926, Genetics 10: 418-41). Triploids that in compound-X-bearing female.
carry an attached X (attached X triploids) are discoverer: Bridges.
more fertile and produce a higher proportion of references: 1916, Genetics 1: 1-52.
triploid progeny than free X triploids. Triploids properties: Phenotype and fertility like those of
are of necessity female and their progeny include normal female. Nondisjunction of X chromosomes
metafemales, metamales, intersexes, triploid and in X/X/Y much higher than in X/X female; about 4
diploid females, and diploid males. Crossing over percent exceptions with two normal X chromosomes
is markedly increased in triploids; Sturtevant and much higher if X's are heterozygous for inver-
(1951, Proc. Natl. Acad. Sci. U.S. 37: 405-7) has sions (Sturtevant and Beadle, 1936, Genetics 21:
mapped chromosome 4 in diplo-4 triploids. B, Bl, 554-604).
Bx, Cy, D, Dfd, H, Hw, J, L2, Me, and Sib are clas- XXYY female
sifiable in a single dose in triploids. Dl, G, N, constitution: X/X/Y/Y; 2/2; 3/3; 4/4. Sex consti-
bv?Vl, pXi s, and all Minutes are recessive in a tution may also be XX/Y/Y, X/XY/Y, or XY/XY.
single dose. Two doses of D, Dl, G, H, bwvl, Px, source: A common product of crosses such as
and Sb produce an extreme phenotype, whereas two YSX-YL/Y male by X/YSX-YL or X/X/Y female,
doses of M or Me are lethal (Schultz, 1934, DIS 1: or X/Y/Y male by XSX-YL/X, X/X/Y, or
55). C(1)EM/Y female.
triploid met a female discoverer: Stern.
constitution: X/X/X/X; 2/2/2; 3/3/3; 4/4/4; third references: 1929, Biol. Zentr. 49: 261-90; 727.
4 may be absent. Cooper, 1956, Genetics 41: 242-64.
source: Found among progeny of tetraploid female properties: Eye color mottled to varying degrees.
(Morgan). Also produced by nondisjunction of sex Posterior and middle legs often malformed. Fer-
chromosomes in C(1)RM/In(l)scs/Y triploid tility and viability reduced. Gametes preponder-
(Frost). antly X/Y in constitution owing to the regular seg-
discoverer: L. V. Morgan. regation of both the X's and the Y's at the first
references: 1925, Genetics 10: 148—78. meiotic division.
Frost, 1960, Proc. Natl. Acad. Sci. U.S. 46: 4 7 - XYY male
51. constitution: X/Y/Y; 2/2; 3/3; 4/4. Sex chromo-
properties: Coarse eyes, wing texture, and bristles. some constitution may also be XY/Y.
Resembles triploid except body smaller and eyes source: About one-fourth the progeny of crosses
more bulging. Inner wing margins often incised. such as X/X/Y female by X/Y male, C(1)RM/Y
Using exceptional triploid females as a standard, female by YSX-YL/Y male, and X/YSX-YL female
Frost (1960) determined that triploid metafemales by X/Y male.
have 25 percent viability. From 24 to 54 percent discoverer: Bridges.
lay eggs (1 to 150 eggs), and about 11 percent of references: 1916, Genetics 1: 1—52.
the eggs develop into adults. properties: Phenotype normal; usually fertile, but
X0 male with certain normal Y chromosomes completely
constitution: X; 2/2; 3/3; 4/4. sterile (R. F. Grell). The two Y chromosomes tend
source: Product of primary nondisjunction of the sex to separate at the first meiotic division, to a de-
chromosomes in either father or mother in cross of gree depending on the source of the y ' s and the X
X/Y male with X/X female. Forms one-fourth the (Grell, 1958, Proc. Intern. Congr. Genet. 10th. Vol.
progeny of crosses such as X/X female by 2: 105).
YsX-YL/0 male or C(l)RM/0 female by X/Y male. Xyyy male
discoverer: Bridges. constitution: X/Y/Y/Y; 2/2; 3/3; 4/4. Sex chromo-
references: 1916, Genetics 1: 1—52. some constitution may also be XY/Y/Y.
properties: Male morphologically normal but entirely discoverer: Stern.
sterile. No motile sperm produced. Primary references: 1929, Biol. Zentr. 49: 261-90.
spermatocyte nuclei lack the morphological ele- Morgan, Bridges, and Schultz, 1934, Carnegie Inst.
ments characteristic of normal male; these ele- Wash. Year Book 33: 274-80.
ments replaced by needle-shaped crystals, which Cooper, 1956, Genetics 41: 242-64.
are found in the nucleus, the cytoplasm, and extra- properties: Morphologically normal male, but with
cellularly (Meyer, Hess, and Beerman, 1961, Chro- mottled eyes as in XXYY female. Almost entirely
mosome 12: 676—716). Nebenkern and axial fila- sterile; Cooper (1956) suggests that the few off-
ment differentiation during spermiogenesis ab- spring may result from X/Y/Y cysts produced by
normal (Kiefer, 1966, Genetics 54: 1441-52). mitotic loss of a Y chromosome.
NONCHROMOSOMAL INHERITANCE
sigma: carbon dioxide sensitivity stabilized state yield only one-fifth as many infec-
origin: Spontaneous. tious units as nonstabilized flies; however, all
discoverer: L'Heritier and Teissier, 1937. progeny of stabilized females are sensitive, as are
references: 1937, Comp. Rend. 205: 1099—1101. part of the progeny of stabilized males. Progeny of
1938, Comp. Rend. 206: 1193-96, 1683. stabilized females are also stabilized. In contrast,
1945, Publ. Lab. Ecole Norm. Super. Biol, (Paris) the sensitive progeny of stabilized males are non-
1: 35-74. stabilized. Several viral mutations that affect trans-
L'Heritier, 1948, Heredity 2: 325-48. mission or replication have been studied; the D.
1951, Cold Spring Harbor Symp. Quant. Biol. 16: melanogaster mutant, ref, inhibits multiplication of
99-112. most viral strains.
1958. In Advances in Virus Research, Vol. 5, SR: Sex Ratio
K. M. Smith and M. A. Lauffer, eds. Academic origin: Artificially inoculated into D, melanogaster
Press, Inc., pp. 195-245. from Si?-bearing D. willistoni or D, nebulosa.
L'Heriter and Plus, 1963. In Biological Organiza- references: Poulson and Sakaguchi, 1961, Genetics
tion at the Cellular and Supercellular Level, 46: 890-91.
R. J. C. Harris, ed. Academic Press, Inc., pp. Sakaguchi and Poulson, 1962, Ann. Rept. Natl,
59-71. Inst. Genetics (Misiraa, Japan) 12: 18-19; 19-21.
phenotype: Flies anesthetized with carbon dioxide 1963, Genetics 48: 841-61.
are paralyzed and die, whereas normal flies recover Poulson, 1963. In Methodology in Basic Genetics,
in a short time. The cause of carbon dioxide sensi- W. J. Burdette, ed. Holden-Day Inc., pp. 404—24.
tivity is a virus or virus like particle whose diam- pheaotype: Females with SR produce few or no male
eter is 180 tafi, as estimated from filtration experi- progeny; SR is transmitted only from mothers to
ments and 45 m/i from X-ray target experiments. daughters. The Si? agent is infective and may be
Carbon dioxide-sensitive strains may be divided established from artificial inocula. The SR condi-
into two types: stabilized and nonstabilized. Arti- tion is always associated with presence of small
tificial inoculation regularly leads to nonstabilized treponemalike spirochetes in hemolymph of af-
condition. In this state, males do not transmit sen- fected females. Degree of stability of the infection
sitivity to progeny but females do transmit it to a differs among D. melanogaster strains. Male dies
part of their progeny. Some flies of a nonstabilized as embryo. Triploid intersexes not killed by S/?
strain achieve the stabilized state. Flies in the nor are transformed females QC/X; tra/ tra).
429
WILD TYPE STOCKS
Austin Samarkand
Started at the University of Texas before 1929. Stock derived from wild flies captured in 1936 at
Viability and fertility good. Samarkand, Uzbek Republic in Asiatic USSR
[.Dubinin, Sokoloff, and Tiniakov, 1937, Biol. Zh.
Canton-S: Canton-Special (Moscow) 6: 1007—54J. Original stock contained a
Derived from wild flies collected in Canton, Ohio. low frequency of inversions in 3R; chromosomes
Selected by Bridges. Contains a recessive for mul- probably are now all of standard sequence. Ives re-
tiple thoracic and scute liar bristles, which overlaps ports that females of his lines of Samarkand are dis-
wild type in most flies but appears sporadically in tinguishable from Oregon-R females in that they
strains partly derived from Canton-S. Bridges found have no faint trident on the thorax and there is al-
that salivary chromosomes were normal. ways a well-defined black band on seventh (most
posterior) dorsolateral abdominal segment.
Lausanne-S: Lausanne-Special Stephenville
Stock derived from wild flies collected in 1938 by Derived from wild flies captured at Stephenville,
Bridges at Lausanne, Wisconsin. Has short posterior Texas in 1935. Salivary chromosomes probably
scutellar bristles. Salivary chromosomes normal, normal. Fertility and viability good.
according to Bridges. Swedish-b
Stock established by Bridges from flies collected
Oregon-R near Stockholm in 1923. Slight abnormality of ab-
Stock derived from wild flies collected in 1925 or dominal banding and position of scutellar bristles.
earlier by D. E. Lancefield at Roseburg, Oregon. Salivary chromosomes homozygous for Dl^Rfiw-R.
Stock contains a slight ebony allele, a branching of Swedish-c
the posterior crossvein (in chromosome 2), and an Derived by Bridges from Swedish-b in 1938. Body
occasional scooped wing. Salivary chromosomes color lighter than that of Swedish-b. Homozygous for
homozygous for Dt(2R)Ore-R. deficiencies in tips of 2L and 2R.
Oregon-R-C Urbana-S: Urbana-Specral
Selected by Bridges in 1938 from Oregon-R. Body Selected by Bridges from flies collected at Urbana,
color not so dark as that of Oregon-R. Homozygous Illinois. Body color somewhat lighter than standard
for wild type. Salivary chromosomes normal.
431
Cytogenetic Maps
All loci with published genetic positions are listed in order in the following table. Mutants
are included even when the published position seems unreasonable. A few mutants are included
that have been placed according to their cytological positions alone (e.g., on chromosome 4).
Clusters of mutants with similar pheriotype appear as the result of assigning locus names to
pseudoalleles and of failure to test for allelism. In this table an asterisk indicates that, accord-
ing to our records, no mutant allele is known to exist at the locus. Where known the cytological
positions of mutants are given; some imprecision has undoubtedly arisen from investigators'
having assigned numbers to bands on Bridges' original map instead of using the revised maps,
which are used in the table.
CYT0GENET1C MAP - CHROMOSOME 1 433
CHROMOSOME 1
Locus Symbol Name Phenotype
-0.4 * 1(1)6 lethal (1) 6 Lethal

o *dar darky Fly small, melanized; male sterile
o *double double Postverticals doubled; wings small
o *dwp dwarp Body small, pale; wings warped; homozygous
lethal
o 1(1)5 5 a lethal(l) 55a Lethal
o 1(1)7'e lethal(l) 7e Lethal; modifies dor
o 1(1 )sc lethal(l) scute Lethal
o *l(l)v306 lethal(l) Y-suppressed lethal
variegated 306.
*l(l)X10 lethal(l) X ray Embryonic lethal
induced 10
0 *pld pallid Body, wings pale
0 su(s) suppressor of sable Suppresses s and v
0.0 1(1 )J< lethal(l) of Lethal
Jacob s-Muller
0.0 y yellow Body yellow; bristles brown
0.0 ac achaete Postdorsocentrals missing; hairs sparse
0.0 Hw Hairy wing Extra bristles, hairs on wings; homozygous
female sterile
0.0 sc scute Bristles missing
0.0 svr silver Body silvery; bristles, trident dark
0.0 brc brachymacrochaetae Bristles small
0.0 *cc ch lorotic Body small, greenish yellow
0.0 *clv-l cloven thorax 1 Thorax cleft
0.0 *cpl cupola Fly small; wings short, canopied; male
s o.o fs(l)N female sterile(l)
sterile
Female sterile
of Nasrat
* 1(1X2 5 lethal(l) Q5 Lethal
l(l)Q 77 lethal(l) Q77 Lethal
GJ 1(1)Q217 lethal(l) Q217 Lethal
o
_
> •
*mul multiple Eyes rough, oval; female sterile
*saw sawtooth Marginal wing hairs clumped
*tdd tiddler Body small
m
4i *ge genitalless External male genitalia absent or deformed;
AMI
male sterile
o.i *l(l)ne lethal(l) non- Pupal lethal
ro*2>î3TO
evaginated
M(l)Bld Minute (1) Blond Bristles fine; late hatching; male lethal
*su(b) suppressor of black Suppresses b
ommatidia Ommatidia disarranged
su(wa) suppressor of Darkens wa
white-apricot
*ctt contorted Wings short; eyes rough; female sterile
1(1 )BN2 lethal(l) EN2 Lethal
*l(l)Q20 lethal(l)Q20 Lethal
XDQ212 lethal(l)Q212 Lethal
* 1(1 >et lethal(l) ring Larval lethal
gland rudimentary
0.3 *l(l)te lethal(l) tracheae Larval lethal
enlarged
stm stubarista Aristae, antennae stubby; bristles short
dor deep orange Eyes orange; female sterile
0.4 *mwi misheld wings Wings spread; eyes ovoid

0.4 *pop popeye Eyes small, buigxiig, rough; male sterile
-0.4 tw twisted Abdomen twisted; body small
0.5 *gtd giantoid Head, body large; late hatching
0.6 rey rough eye Eyes small, rough
0.6 rsi reduced size Body small
0.6 *srd small round Body small; eyes small, round
-0.6 br broad Wings short, broad
0.7 *dw-sc dwarf with scute Body small
0.8 mk murky Body small, dark; eyes dull red; female
sterile
-0.8 pn prune Eyes brownish purple
0.9 *fc faulty chaetae Bristles short, thin
0.9 & giant Larva, pupa, adult large
0.9 *ovi ovioculus Eyes small, oval, rough; wings spread; male
sterile
Pgd Phosphogluconate Affects phosphogluconate dehydrogenase
dehydrogenase mobility
*sbs stubs Wings abnormal; eyes small; female sterile
kz kurz Bristles short, thin; postscuteliars missing
*fb fine bristle Bristles thin
ws waisted Abdomen constricted anteriorly
e(bx) enhancer of bi- Eyes variegated, enhances most bx alleles
thorax
z zeste Male normal; female eyes yellow
*bsc bent scutellars Bristles bent
* 1(1)1 lethal(l) 1 Lethal
lethal(l) zwl Lethal
I(l)zw4 lethal(l) zw4 Lethal
*abe abnormal eye Eyes rough
1(1)Q219 lethal(l) Q219 Lethal
I(ljzw3 lethai(l) zw3 Lethal
I(l)zw6 lethal(i) zwo Lethal
misproportioned Abdomen deformed; wings short; bristles
thin
1.3 swollen antenna Antennae swollen; female sterile
CYTOGENETIC MAP - CHROMOSOME 1 435
1.4 *crrn cramped Antennae short; aristae crooked; sterile

1.4 *1(1)Q9 lethal(l) Q9 Lethal
1.4 I(l)zw5 lethal(l) zw5 Lethal
1.4 *pte pterygion Wings short, drooping; abdomen large
1.4 *ves vestigium Wings, eyes abnormal
1.5 *l(l)Q40 lethal(l) Q40 Lethal
1.5 *l(l)TS-56 lethal(l) 56 of T. Embryonic lethal
Shiomi
1.5 *smh smaller thinner Body small; bristles thin
1.5 w white Eyes, Malpighian tubules white
1.6 dwg deformed wings Wings broad; bristles fine; eyes small; male
sterile
1.6 1(1)3C3 lethal(l) 3C3 Lethal
1.7 rst roughest Eyes rough, bulging; body small; male
sterile
2 1(1 )v139 lethal(l) variegated Y-suppressed lethal
139
2.2 swb strawberry Eyes large, rough, patchy colored
2.3 vt verticals Verticals absent
2.4 fla flateye Body, eyes small
2.5 *cpw canopy wing Wings short, broad; veins incomplete; male
sterile
2.5 * l(l)mt lethal(l) midget Larval lethal
2.8 1(1)Q218 lethal(l) Q218 Lethal
3 1(1)EN12 lethal(l) EN12 Lethal
3 *we wee Body small; eyes rough; bristles fine;
wings spread
3.0 Ax Abruptex Wings short, arched; veins incomplete
3.0 Co Confluens Veins thick, deltas at margin
3.0 fa facet Eyes rough
3.0 spl split Eyes small, rough; many bristles doubled,
some missing
-3.0 N Notch Wings cut; veins thick, deltas at margin;
male lethal
•—3.0 nd notchoid Wings cut; veins thick
3.1 *im interrupted margin Margin nicked, extra veins; female sterile
3.2 *s/c stiff chaetae Bristles short, stiff
3.3 rud ruddle Eyes reddish brown
3.6 *mtb matt brown Eyes dull; wings spread; male sterile
3.6 sic slim chaetae Bristles thin, short
3.7 *sth small thin Body small; bristles short, thin; female
sterile
4.4 TV raven Body small, dark; eyes dark; wings short
\ 4.5 A Abnormal abdomen Tergites and sternites frayed; abdominal
bristles missing
1 4.5 *tta reduced tarsi Tarsi short; body, eyes, wings small; male
sterile
1 4.5 Sc Scotched eye Eyes rough; male lethal
*— 4.6 dm diminutive Bristles, body small, slender; female
sterile
5 *su(dx) suppressor of Suppresses dx

deltex
5 M(1)3E Minute(l) in 3E Bristles fine; late hatching; male lethal
5.4 cho chocolate Eyes brown
5.5 *mf macro fine Bristles short, thin; body small
5.5 *Z Zerknittert Wings crumpled
5.5 ec echinus Eyes large, bulging, rough; wings short,
broad
5.6 *te tenerchaetae Bristles short, fine; eyes dark
5.7 Oce Ocellarless Ocellars missing
5.8 *l(l)TS-45 lethal(l) 45 of T. Embryonic lethal
Shiomi
5.9 *e(g) enhancer of garnet Enhances g
6 H1)C lethal(l) C Lethal
6.7 mo micro-oculus Eyes small; wings narrow
6.8 amb amber Body pale yellow; bristles short, thin; male
sterile
6.8 M(1)4BC Minute(l) in 4BC Bristles fine; late hatching; male lethal
6.8 Qd Quadroon Tergites broadly banded
6.9 bi bifid Veins fused; wings short, spread
7.3 lac lacquered Body glistens; bristles, eyes, wings ab-
normal
7.3 peb pebbled Eyes rough
7.5 rb ruby Eyes ruby
8 *l(l)ts lethal(l) tempera- Lethal at 23°C
ture sensitive
8.0 dow downy Bristles short, slender; male sterile
8.0 *l(l)trs lethal(l) tracheae Larval lethal
stretched
8.7 *mib miniature bristles Bristles short, thin; body dark; male sterile
10 1(1)EN9 lethal(l) EN9 Larval lethal
10 1(1 )ml lethal(l) melanoma- Larval lethal
like
11 *lzl lozengelike Eyes rough
Locus Symbol Nor Phenotype
11.0 rg rugose Eyes rough; wings thin, margins frayed

11.3 *apx apexless Wings broad, blunt, incised
11.5 *rgt reduced pigment Male fifth tergite pale; male sterile
12.5 bo bordeaux Eyes dark wine
12.8 omm omma toreductum Some peripheral ommatidia missing; eyes
rough
13 *Ch Curled blistered Wings curled, blistered with px
13.1 *1(1)Q1 lethal(l) Ql Lethal
13.3 *dvw divergent wings Wings spread in male
13.3 *dw( dwarfoid Fly small
13.4 1(1)EN13 lethal(l) EN13 Semilethal
13.6 ex curlex Wings warped, bent upward
13.7 cv crossveinless Crossveins absent
13.9 *dla deformed antennae Antennae, aristae abnormal; eyes, wings,
bristles small; female sterile
14 lethal(l) jawless Larval lethal
14 *M(l)30 Minute(l) 30 Bristles fine; late hatching; male lethal
14.3 *mur murrey Eyes red purple; bristles small; sterile
14.4 *rmp rumpled Wings unexpanded; bristles deranged
14.6 *stb short bristle Bristles short, thin
15.0 TUX roughex Eyes small, rough; male sterile
15.2 Ext Extras Extra veins; male lethal
15.2 *pra prawny abdomen Body slender; wings short
16.0 *sct scooped thickvein Wings short, scooped; veins thick; male
sterile
16.3 vs vesiculated Wings warped, wrinkled, blistered, spread
16.5 *cb club Wings clubbed; sternopleurals absent
17.0 dx deltex Veins thick, deltas at margins
17.5 *lem lemon Body, wings yellow; bristles black; sterile
17.5 ov oval Eyes oval, rough
17.5 *tmc tonomacrochaetae Bristles thin; abdomen pale
17.9 *lCl)sd lethal(l) scheiben Larval lethal
defekt
17.9 shl shifted L3 incomplete, shifted toward L4
18 *dep depressed Wings turned down at tips
18 *Eyl Eyeluf One or both eyes small
18.0 *tnt "tent Wings drooping; male sterile
18.7 !(1)Q82 lethal(l) Q82 Lethal
18.9 cm carmine Eyes ruby
19.1 Fl Female lethal Female lethal
19.3 scp scooped Wing tips upturned
19.5 *tyb-2 tiny bristle 2 Bristles small, thin
20.0 ct cut Wings cut, scalloped; eyes kidney shaped
20.1 bis bistre Eyes brown
20.2 *kf kinked femur Body small; femurs kinked; wings unex-
pended
20.2 *tre triangle eye Eyes triangular; female sterile
20.3 KDQ6 7 lethal(l) Q67 Lethal
20.4 *stu small tumoroid Body small; pseudotumors present
20.5 HDQ2J lethal(l) Q21 Lethal
20.7 *lgh long haired Bristles long; body small; male sterile
20.9 postverticalless Postverticals absent; thoracic hairs sparse;

female sterile
20.9 *sht short tarsi Legs short
21.0 sn singed Bristles twisted, short; hairs kinked; fe-
male sterile
21.1 *sch slender chaetae Bristles thin, short
21.3 * 1(1)8 lethal(l) § Larval lethal
21.6 dfw deflected wing Wings upheld, spread; female sterile
21.7 l(l)mya lethal(l) myosphe- Embryonic lethal
roid
21.7 *rdb reddish brown Eyes reddish brown; male sterile
21.9 *bwl bow-legged Wings, bristles, legs short; male sterile
22.0 *SCT scruff Hairs and bristles absent or doubled
22.4 shm short macros Bristles short; male sterile
22.6 spx split thorax Longitudinal furrow on thorax
22.7 *ha hair bristles Body small; bristles thin, short
^"z 22.7 *l(l)Q30 lethal(l) Q30 Lethal
23 *depl depressedlike Wings drooping; fly dark
23 *dis distorted eye Eyes rough
23 lix little isoxanthop- Little isoxanthopterin in testis sheath
K >
terin
23.1 ii goggle Eyes protruding; female sterile
23.1 pt platinum Body, bristles pale; male sterile
23.1 *smb slim bristle Bristles thin, short; female sterile
23.1 oc ocelliless Ocelli absent; female sterile
23.2 ptg pentagon Trident dark
23.4 ccw concave wing Wings short, narrow
23.8 *ch-b chilblained-b Tarsi fused
24 *jyx jaunty x Wings upcurved at tips
** > 24.2 1(1)Q244 lethal(l) Q244 Lethal
24.3 *dd displaced Head shortened; antennae sunken; eyes de-
formed
~P Z£-=\ 24.6 *bre bright eye Eyes bright red; wings short; abdomen large
24.6 *svs shortened veins Wings abnormal; female sterile
25 *sml small Body small; wings short; eyes small, rough,
bulging
25 *tbd tiny bristleoid Bristles short, thin
25.1 *e/r elliptical rough Eyes oval, rough; wings broad
25.4 tea rearranged tergites Tergites abnormal
cssw 25.6 *smp small pallid Body small, pale; female sterile
25.9 dlv deltoid veins Extra veins; body, eyes, wings small
26 *asx ascutex Scutellar groove shallow; body pale
26.5 * K1X23 lethal(l)Q3 Lethal
27 *Lg Large Body large; homozygous lethal
27.1 *con condensed Abdomen, thorax, wings short; male sterile
27.2 *ddl displacedlike Head shortened; antennae sunken; eyes de-
formed; male sterile
27.3 *dss disturbed segmen- Abdominal segments deformed; eyes small;
tation female sterile
27.3 tar tarry Legs black or spotted
27.5 t tan Body tan
27.7 amx almondex Eyes reduced; female sterile
27.7 Iz lozenge Eyes narrow, ovoid, irregular surface; fe-
male sterile
27.8 *tha thin arched Wings arched or drooping; bristles short,

thin
28.1 dvt divers Wings short, dark; body small
28.3 *opb opaque broad Wings short, broad, opaque; female sterile
28.6 *ke kidney eye Eyes small, rough, kidney shaped; sterile
28.6 *1(1X>7 lethal(l) Q7 Lethal
29 *pi& pigmy Body small, melanotic
29.0 *me focal melanosis Lethal
29.6 fin finer Body small; bristles short, thin; male
sterile
29.8 *sto stocky Body short, stocky; male sterile
29.9 sma smaller Body small
30 su(Cbx) suppressor of Suppresses Cbx
Contrabithorax
31 *flw flap wing Wings spread, curled; eyes bulging
32 *aw awry Wings upcurled, wavy, convex, opaque
32 *e(we) enhancer of white Enhances some w alleles; female sterile
eosin
32 ny notchy Wing tips nicked
32.4 *pat patchytergum Abdominal pigmentation patchy; wings
spread; male sterile
32.5 *df defective Head bristles near ocelli missing
32.6 elm clumpy marginals Marginal wing hairs bent; bristles stiff
32.8 KDQ54 lethal(l) Q54 Lethal
32.8 *sbt shorter bristles Bristles short, thin; wings spread
32.8 ras raspberry Eyes dark ruby
32.9 ww wider ring Wings short, broad
33 *brd broadened Wings expanded
33.0 *csk costakink Eyes, wings small
33.0 *osh outshifted Wings short
33.0 *wgv wing variance Wings held abnormally; male sterile
33.0 V vermilion Eyes scarlet; ocelli colorless
33.2 dWK dwarfex Body small; wings coarse
33.2 *1(1X>26 lethal(l) Q26 Lethal
33.4 sbr small bristle Bristles small
33.5 dsh dishevelled Thoracic hairs deranged; female sterile
33.5 l(l)Q66 lethal(l) Q66 Lethal
33.5 *tny thorny Body deformed; eyes small, rough; wings
crumpled; male sterile
33.7 *dft deformed terga Tergites deformed
33.7 slm slim Body small; abdomen narrow
34.0 l(l)Q211 lethal(l)Q211 Lethal
34.3 *stt spotty Abdomen spotted; male sterile
34.7 *rdp reduplicated Legs malformed, branched
34.9 *fnc fine chaetae Bristles short, fine; male sterile
35.7 *ano anomogenitals Bristles, hairs sparse; male genitalia ab-
normal; male sterile
35.8 *tb tiny bristle Bristles short, fine
36 tyl tiny like Bristles short, thin, stubbleiike
— 36.1 m miniature Wings small

36.2 dy dusky Wings small, dark
36.3 *M(l)k Minute(l) k Bristles fine; late hatching; male lethal
36.3 *shl shorter legs Body small; legs short
36.3 trb thread bristle Bristles short, thin; female sterile
36.4 *gr gracile Body small; male sterile
37.1 *twt twirled tips Wings unexpended, tips twisted
37.2 KW52 lethal(l) Q52 Lethal
37.2 *ob oblique Wings truncated; veins abnormal
37.2 *plw pale wing Body, wings, bristles pale yellow
38.0 *gli glide Wings spread; male sterile
38.2 1(1JQ58 lethal(l) Q58 Lethal
38.3 *alo alopecia Abdominal hairs sparse; pigmentation light
38.3 *1(1JQ45 lethal(l) Q45 Lethal
- 38.3 fw furrowed Eyes furrowed; head, scutellum shortened;
bristles short, gnarled
38.5 *aw-b awry-b Wings upcurled, wavy, convex, opaque
38.9 *wtw water wings Wings short, broad, blistered
39.0 *shb shortened bristles Bristles short, thin; female sterile
39.8 *brw broader wing Wings broad, round; male sterile
40.1 1(1 X? 2 02 lethal(l) Q2O2 Lethal
40.3 crt crumpled tips Wing tips crumpled
40.8 *som sombre Body, eyes dark
40.8 *ups upright scutellars Scutellars upheld; male sterile
41 *tu-53 tumor-53 Adult melanotic tumors; wings abnormal
41.0 up upheld Wings upheld
41.1 pun puny Body small; wings short
41.1 *taw tawny Head, thorax dark; abdomen pale
-41.9 wy wavy Wings wavy
42 *kk kinky Bristles bent
42.0 *c!v-2 cloven thorax 2 Thorax cleft; wings reduced
42.0 eb ebonized Body dark; wings short; female sterile
42.1 *tht thickset Body short, stocky
42.5 *swy swarthy Body dark
43 1(1)ENU lethal(l) EN11 Lethal
43.0 s sable Body dark; trident prominent
43.2 *bla bladderwing Wings small, deformed, inflated; female
sterile
43.3 cop copper Eyes brownish red
43.9 *ten tenuis chaetae Bristles short, thin
44.4 & garnet Eyes purplish ruby
44.5 t(UQ59 lethal(l)Q59 Lethal
44.5 ty tiny Bristles, body small; female sterile
- 44.6 *dyb dusky body Body dark; eyes brown; female sterile
45 *Bxd Beadexoid Wings long, narrow, ragged
45 *cb( clubfoot Legs short; wings warped
45.2 na narrow abdomen Abdomen long, cylindrical
45.3 *$1b slim body Body narrow
45.6 *abt abnormal tergites Abdomen, eyes, wings deformed
45.7 *smn small narrow Abdomen narrow; fly weak

46.1 *stp silver tips Bristles thin, pale; male sterile
47.5 *shp shrimp Body small
47.8 *lme lame Legs, wings deformed
— 47.9 Pi pleated Wings pleated
48.0 *thb thin bristle Bristles thin
48.1 rm rimy Eyes brownish red; wings pleated
48.1 *twg twisted genitals External genitalia positioned abnormally
48.4 *sge shifted genitals Genitalia, anal plates rotated; male sterile
48.6 sla slimma Body narrow; female sterile
48.7 mgt midget Body small
1
i 2>
i>
\ 48.9
49.1
49.3
49.7
*thm
1(1)Q76
vb
thv
thin macros
lethal(l) Q76
vibrissae
thick vein
Bristles short, thin
Lethal
Vibrissae tufted
Wing veins thick
r l(l)EN10a lethal(l) ENlOa Pupal lethal
/ 50
50 *op opaque Wings opaque, divergent, convex
50 Tul Turneduplike Wings curled, wrinkled
50.1 *hpa hyperantenna Antennae large; head, eyes deformed; fe-
male sterile
50.1 sld slender Body small, slim
50.6 *Stp-l Strapped in chromo- Male wings incised with Stp-2
some 1
51.2 *slw slope wing Wings spread
51.5 *exl exiguous Fly small, dark
— 51.5 sd scalloped Wing margins scalloped; veins thickened
51.6 Bg Bag Wings inflated; veins abnormal; male lethal
51.6 tc tiny chaetae Bristles short, fine
51 9 l(l)Q70 lethal(l) Q70 Lethal
I *
I 51.9 *smt small thorax Thorax, head small
52.0 *mch minute chaetae Bristles short, fine; body small
52.3 *drw droopy wing Wings drooping; body small; male sterile
52.3 *unr unexpanded irreg- Wings somewhat unexpanded
ular
52.4 *ber berry tail Abdomen narrow; male genitalia defective;
male sterile
52.5 *us undersized Body small
52.6 *msc melanoscutellum Scutellurn dark
52.9 *brb broad abdomen Abdomen broad; thorax, wings short
53 *fi frail Wings small, thin; bristles fine
53.0 HDQ63 lethal(l) Q63 Lethal
53.5 *pyp polyphene Wings spread; eyes small, rough; female
sterile
53.5 si small wing Wings short, blunt; eyes large
53.5 *sln slimmer abdomen Body small; abdomen narrow; female sterile
54.0 me microchaete Bristles, hairs irregular; eyes rough; wings
short
54.2 nrs narrow scoop Wings narrow, short, scooped
54.4 *rdt reduced thorax Head, thorax small; wings short; male
sterile
54.4 un uneven Eyes small, rough
54.5 ace acclinal wing Wings upheld, sloping back
54.5 r rudimentary Wings truncated; female sterile
55 if inflated Wings inflated, small; veins abnormal
55.5 *St Stumpy Body short; bristles short, thin; male lethal
56 cs creased Wings creased longitudinally
56 *de deacon Body, wings narrow; eyes flat
56 *l(l)tr lethal(l) tracheae Larval lethal
ramified
56.5 std staroid Eyes small, oval, rough; bristles short;
male sterile
M(l)o Minute(l) o Bristles fine; late hatching; male lethal
*1(1)Q18 lethal(l) Q18 Lethal
l(l)Q214 lethal(l)Q214 Lethal
I(l)v451 lethal(l) variegated Y-suppressed lethal; male sterile
451
forked Bristles short, bent
fine lash Bristles thin; eyes small
Splotched Wing hairs disarranged
late hatching Body large, develops slowly
Bar Eyes small, narrow
pod foot Tarsi swollen
deranged Thoracic hairs deranged
Enhancer of Bar Enhances B; homozygous lethal
fluff Bristles short, fine
Shaker Fly trembles when etherized
lethal(l) Q23 Lethal
bulging Eyes rough, bulging
rabbit Acrostichal hairs irregular
convex wing Wings short, arched
dwarf unexpanded Body small; semilethal
lethal(l)Q24 Lethal
side wings Wings rooflike; male sterile
vacuolated Wings blistered
splay wing Wings short; eyes small; male sterile
tumorous Tumors
straight abdomen Abdomen long, narrow, straight
lethal(l) EN10 Lethal
lethal (1) tracheae Larval lethal
lacking
Turned-up wing Wings curled, wrinkled
outstretched small Wings spread, eyes small, or both
eye
Beadex Wings long, narrow, margins excised
heldup Wings upheld
fused L3, L4 partly fused; wings spread; female
sterile
59.8 *bk buckled Wings divergent
59.8 *rdm reduced macros Bristles thin, short
59.9 *bkl buckledlike Wings divergent
60 abw abnormal wings Wings small, upturned; L5, crossveins ab-

sent
60.1 *crk crooked setae Bristles thin, short
60.1 obi oblique wings Wings spread
60.1 *ton tonochaetae Bristles short, thin; female sterile
60.3 sts streaked sterni Sternites striped
60.7 *thl thick legs Legs short, swollen; wings small, broad
60.8 *pph polyphenic Body small; eyes bright; wings abnormal
60.8 *sby small body Body small, pale
61.1 *smd smalloid Body small
61.5 *coc collapsed ocelli Ocelli small, fi^t
61.9 *meg magaoculus Kyes large, rough; wings affected
62.0 *srb smaller body Body small; bristles fine
62.5 car carnation Eyes dark ruby
62.7 M(l)n Minute(l) n Bristles fine; late hatching; male lethal
62.9 RD(1) Recovery Alters sex ratio with RD(2)
Disrupter(l)
63 lo folded Wings unexpanded; halteres shriveled
63 pub pubescent Bristles short, fine; abdomen pale; male
sterile
63 Zw Zwischenferment Affects electrophoretic mobility of glucose
6-phosphate dehydrogenase
63.1 unp unexpended Wings unexpanded
63.4 *1(1)X27 lethal(l) X-ray- Embryonic lethal
induced 27
63.9 *kno knobbyhead Head, eyes abnormal
64.0 sw short wing Wings spread, incised; veins irregular; eyes
small, rough
64.1 me I melanized Body dark; eyes dull red
64.4 wa warty Eyes rough
64.5 tuh-1 tumorous head in Asymmetric head growths with tuh-3
chromosome 1
64.7 *mdg midgoid Body small, pale
64.8 mal maroonlike Eyes purple; lacks xanthine dehydrogenase
65.6 *cf cleft Wings small; veins abnormal; male sterile
65.7 *1(W8 lethal(l) Q8 Lethal
65.7 ot outheld Wings spread; hairs sparse; male sterile
65.9 su(f) suppressor of Suppresses some i alleles; dilutes wa
forked
65.9 *unc uncoordinated Leg movements uncoordinated; wings up-
held, curled
66 *Hv Hooked veins Veins branched; eyes small, rough; female
lethal
66 *l(l)w lethal(l) white Male normal; female lethal
66.0 bb bobbed Bristles small; abdomen etched
66.1 *(m fine macros Fly small; bristles short, thin
67.9 rtt refringent Wings yellow, iridescent; female sterile
68.1 It little fly Body small; abdomen narrow, tumorous
68.9 *sme smaller eye Body small; eyes small, dark; male sterile
70 *bottl bordered Wings small; veins ragged
CHROMOSOME 2
*OCT ochracea Eyes light, darken with age

*tg telegraph L2 interrupted or thin
K2)gl lethal(2) giant Late larval lethal
net net Extra wing veins
M(2)21Cl-2 Minute (2) in 21C1-2 Bristles short, tiny; homozygous lethal
al aristaless Aristae reduced; scutellars divergent
ex expanded Wings wide, large
ds dachsous Wings short, broad; crossveins close
S Star Eyes small, narrow, rough; homozygous
lethal
ast asteroid Eyes small, rough; wing veins incomplete
*l(2)Sp9a lethal(2) of Lethal
Speiss 9a
*Stp-2 Strapped in Male wings incised with Stp-1
chromosome 2
2.3 s/ir shrunken Body small, dark
3 Su(S) Suppressor of Star Suppresses 5
3.2 K2)Sp7 lethal(2) of Lethal
Speiss 7
3.8 shv short vein L2, L3 incomplete
4.0 ho heldout Wings spread
4.3 *vst vestar Wings small, straplike; female sterile
5 fs(2)B female sterile(2) Female sterile; ovaries tumorous
of Bridges
5.0 rub rubroad Eyes rough, kidney shaped; wings broad
6 ECS) Enhancer of Star Eyes rough; enhances some S and ast
alleles
*P1 Pearl Pearl-like nodes in wings; eyes small,
rough; homozygous lethal
6.1 Cy Curly Wings upcurled; homozygous lethal
8.3 K2)ay lethal(2) ay Lethal
8.4 *sq square Wings truncated
10 *Dt Detached L2 incomplete
10.5 ang angle wing Wings upheld, spread

r- 11.0 ed echinoid Eyes large, rough
DD tot -
- 12 G Gull Wings large, spread, curved; homozygous
r
O E
J - 12.0 ft fat
lethal
Abdomen short, fat; thorax broad; wings
short, broad
h r- 12.9 M(2)z Minute(2)z Bristles fine; late hatching; homozygous
_/
lethal
- 13 dw-24F dwarf in 24F Body small; abdomen narrow
- 13.0 dp dumpy Wings truncated; thoracic bristles whorled
r\> 14 *pw pink wing Eyes ruby; wings short, crumpled
15 * 1(2)1076 lethal(2) 1076 Lethal
15 l(2)cg lethal(2) with Lethal
c omb gap
> 15.0 *M(2)S1 Minute (2) of Bristle fine; late hatching, homozygous
>
OD - Schultz 1 lethal
16 tkv thick veins Veins thickened, branched
16.0 Sk Streak Dark streak down thorax; homozygous lethal
j — 16.5 c/ clot Eyes dark maroon
17 Pi pied Eyes small, rough; wings arched; male
sterile
o -
t\)
— 21.9 spa spade Wings short, broad, pointed at tips

22.0 *fs(2)E2 female sterile(2) of Female sterile
Edmonds on 2
<©:
— 22.0 Sp Sternopleural Extra sternopleurals; homozygous lethal
22.9 lys lysine Accumulates lysine
23.4 cui curvi Wings upcurved
24 *cr.b cream b Dilutes we
m > 24.0 gt-4 giant 4 Body large; late hatching
27.3 re red cells Normal; pigments fat cells of lys fly
28 *ms(2)E3 male sterile(2) of Male sterile
Edmondson 3
rv) ^ 29.5 tu-48 tumor 48 Abdominal melanotic tumors
Q» = 30 *flp flipper Fly dwarfed, dark; wings unexpanded;
sterile
30.3 l(2)pm lethal(2) polymorph Larval or pupal lethal
O > 31.0 d dachs Tarsi four jointed; eyes small, rough
32.0 *l(2)Spl4 lethal(2) of Speiss Lethal
14
33 *fy fuzzy Hairs irregular
35.0 *l(2)Spl lethal(2) of Speiss Lethal
55a 1
i
Edmondson 9
36 corr corrugated wing Wings wavy, short
)
00
o -
o > >
03 I
O z
<oz
m -
36 *Si Ski Wing tips upturned with si-3

36 *ter terraced Eyes seamed horizontally
10
38 *Sq Squat Wings, thorax, head short, broad; homozy-
gous lethal
39 *fol folded wings Wings folded
39. 3 da daughterless Female produces no daughters
40 *pg prong Extra crossveins
40 *wd wavoid Wings waved
"——I
41. 0 J Jammed Wings narrow
43 M(2)e Minute(2) e Bristles fine; late hatching; female sterile;
m > homozygous lethal
43.7 *Cpt Clipt Bristles short; homozygous lethal; male
sterile
43.8 l(2)bl lethal(2) bluter Late pupal lethal
44 an ancon Wings, legs short
CD 44.0 ab abrupt L5 incomplete
> 44.0 *ms(2)2 male sterile(2) 2 Male sterile
o - 44.7 *apb apterblister Wings notched, spread, blistered; short
lived
45 * oph ophthalmopedia Eyes kidney shaped
47 *l(2)pup lethal(2) pupal Pupal lethal
m - 47.0 nub nubbin Wings small, curved; margins interrupted;
veins missing
47.5 *fs(2)E3 female sterile(2) of Female sterile; wings narrow, curved
Edmonds on 3
47.9 *ms(2)E4 male sterile(2) of Male sterile
Edmondson 4
03
O
o» -
<•> -n - " — a
CD 1
— 48.2 rk rickets Legs flattened, bent; wings unexpanded

— 48.5 b black Body black
48.5 *is(2)E4 female sterile(2) of Female sterile
Edmondson 4
48.7 Coi Coiled Wings curled
h - 48.7 j j aunty Wings upturned
49 *l(2)Sp2b lethal(2) of Speiss Lethal
2b
49 l(2)Sp9b lethal(2) of Speiss Lethal
9b
1(2)3 9a lethal(2) 39a Lethal
I(2)H lethal(2) of Usually pupal lethal; few survivors
Humphrey
lm limited Sternites abnormal; bristles sparse; female
sterile
el elbow Wings extended, bent; L5 shortened;
halteres reduced
*l(2)Sp6b lethal(2) of Speiss Lethal
6b
Adh Alcohol dehydro- Affects alcohol dehydrogenase electropho-
genase retic mobility
*ls(2)E5 female sterile(2) of Female semisterile
Edmondson 5
*E(H) Enhancer of Hair- Enhances H
less
Su(H) Suppressor of Hair- Suppresses H; homozygous lethal
less
pu pupal Wings unexpanded
Sco Scutoid Scutellars absent; homozygous lethal
id reduced Bristles reduced; female sterile
* baton baton Abdomen elongated; eyes lobed
pys polychaetous Extra bristles
tyr-1 tyrosinase 1 Tyrosinase activity low
cm cream underscored Dilutes we and P
ref refractaire Inhibits growth of carbon dioxide sensitivity
virus
ck crinkled Wings flimsy, wavy; bristles stubby
rdo reduced ocelli Ocelli small, colorless
l(2)Bld lethal(2) from Lethal
Blond
53.2 Tit Tuft Extra bristles

-53.5 M(2)H Minute(2) from Bristles fine; late hatching; homozygous
CD ^ T(Y;2)H lethal
_ > — 53.9 hk hook Bristle tips bent; eyes rough; wings diver-
gent
O = 54 sple spiny legs Leg hairs irregular
54 *M(2)m Minute(2) m Bristles fine; late hatching; homozygous
lethal
54.3 bri bright Eyes bright red
o» - Edmondson 6
- 54.5 pr purple Eyes purplish ruby
U) -n r- 54.6 *Cd Coiledex Wings upcurved; homozygous lethal
> 54.7 th roughish Eyes rough
oo > > j 54.8 Bl Bristle Bristles short, blunt, thick; homozygous
lethal
>
CD Edmondson 5
o> -
Edmondson 6
Edmondson 7
54.9 Alu Alula Alula fused with wing; wings buckled
54.9 esc extra sex combs Male has extra sex combs; sterile
54.9 *Jag Jagged Wings jagged; eyes small, rough
54.9 wi witty eye Eyes variegated, rough; extra vibrissae
present
55 *inb incised balloon Wings spread, margin incised
55 *1(2)1323 lethal(2) 1323 Lethal
55 l(2)CTC lethal(2) cryp- Pupal lethal
tocephal
55 *pads pads Wings unexpanded
55 SD Segregation Dis- Morphologically normal; segregation ab-
torter normal in male
55 tri trident Trident dark
55 *uex unextended Wings unexpanded
55.0 l(2)55i lethal(2) 55i Larval lethal
- 55.0 It light Eyes yellowish pink; ocelli colorless
p 55.1 il rolled Wing edges rolled; L4 interrupted; eyes

small, rough
- 55.1 *l(2)Sp9c lethal(2) of Speiss Lethal
9c
55.1 *I(2)Sp9d lethal(2) of Speiss Lethal
9d
- 55.1 l(2)Spll lethal(2) of Speiss Lethal
11
l J - 55.1 l(2)Spl5 lethal(2) of Speiss Lethal
15
- 55.1 M(2)S2 Minute(2) of Bristles fine; late hatching; body pale;
Schultz 2 homozygous lethal
- 55.1 stw strjw Hairs yellow; bristle tips pale
— 55.2 ap apterous Wings, halteres missing; bristles sparse;
sterile
55.2 *Cu Curl Wings curled
55.2 *fs(2)E7 female sterile(2) of Female sterile; eggs collapse
Edmondson 7
- 55.2 msf mis formed Eyes misshapen; wings short, crumpled;
legs short
— 55.3 pk prickle Acrostichals whorled; costal wing hairs
g -
slanted anteriorly
— 55.3 tk thick Legs thick; wings short, broad
— 55.5 tut tufted Tuft of hairs between eyes and antennae
Edmondson 8
55.7 bur burgundy Eyes dull brown
55.8 *ae aeroplane Wings spread; halteres drooping
o 55.9 *tj tarsi irregular Tarsal segments fused, swollen
56 l(2)hst lethal(2) histolytic Pupal lethal
56 ltd lightoid Eyes yellowish pink
56 *mbs miniature blistered Wings small, blistered; bristles bent
56 *Rw Rough wing Wings notched; veins irregular; female
sterile
56.6 ta tapered Wings narrow, pointed; male sterile
56.8 *awu augenwulst Eyes indented anteriorly
57 *dil specific dilutor Dilutes bw, we, we2, wbl
57 *M(2)38b Minute(2) 38b Bristles fine; late hatching; homozygous
lethal
57 *ms(2)E9 male sterile(2) of Male sterile
Edmondson 9
57 * Tyr-2 Tyrosinase 2 Tyrosinase activity low
57.1 *buo burnt orange Eyes bright orange-brown
57.1 SO sine oculis Ocelli absent; eyes small
57.5 cn cinnabar Eyes bright red
57.5 Ps Pigmentless Female abdominal pigment reduced; homo-
zygous lethal
57.6 *ls(2)El female sterile(2) of Female sterile; ovaries rudimentary
Edmondson 1
58 put puff Wings inflated, warped
58.5 bio bloated Wings spread, crumpled, blistered
58.6 *smk smoky Body dark
59.5 *Np Notopleural Bristles short, blunt; wings short, broad;

60.1 *at arctus oculus Eyes small, narrow
60.5 arch arch Wings downcurved
60.5 ix inters ex Female changed into sterile intersex
60.7 ad arcoid Wings arched, broad; crossveins close
60.8 chl chaetelle Bristles, body small; wing venation ab-
normal
61 whd withered Wings warped or shrunken
12
61.5 *l(2)Sp8 lethal(2) of Speiss Lethal
Q
O
62 *upw upward Wing tips upturned
62.0 che cherub Wings short, downcurved; male sterile
62.0 en engrailed Scutellum cleft; bristles hooked; veins in-
terrupted, branched
Edmonds on 8
62.7 *Eye Eyeless dominant Eyes small; homozygous lethal
in chromosome 2
63 *lnd Indented Eyes kidney shaped
63.3 spt spermatheca Spermathecae abnormal
63.5 twl twirl Wings curled
63.6 *sps spastic Lethal; survivors weak
to — 64.3 Go Gold tip Bristle tips pale, curved; homozygous
lethal
64.7 l(2)a lethal(2) a Lethal
65 *Bkd Blackoid Body black
65 *M(2)40c Minute(2) 40c Bristles fine; late hatching; homozygous
lethal
65.2 po pale ocelli Ocelli colorless
65.3 *1(2)SP18 lethal(2) of Speiss Lethal
18
65.5 ms(2)l male sterile(2) 1 Male sterile; female fertile
Edmondson 10

00
66.7 sea scabrous Eyes large, rough
67 *Str Stretched wings Wings held out; homozygous lethal
o> —
67.0 vg vestigial Wings vestigial; halteres small
rC 67.1 1(2 )C lethal(2) of Curry Late larval lethal
9»
68 *ms(2)Ell male sterile(2) of Male sterile
All
Edmondson 11
o 6>- 68 *ts telescope Abdominal segments long
_
-si
Co 68.2 *ms(2)E12 male sterile(2) of Male sterile
— T
Edmondson 12
m —
69.7 wx waxy Wings opaque, small; male sterile
0)
-
fv
(i)
> U>
>
— CD
Ill
-I
00
i >
>
> > tfk
I'll
>
5>
u>
at —
—
CD »
>
IVI
O •y
Mil
rn >
ii >
•7-
> 6
- *
CD
V>
en ?>
O _
••^>
—
-|i-
CD u» -
a> >
II
•>
If o» -
• -
m IT

~~
o <6
70 l(2)mr2 lethal(2) with Lethal
AIA
1-4
(Jl
" 0) — morula-2
d> — 70 Wings curled, dark, waxy; hpmozygous
T > U Upturned
lethal
no 70.8 Pfd Pufdi Wings spread, blistered
11-6
E 71.0 bat bat Wings extended, bent back

t VIIV
o $
71.1 eg comb gap Sex combs large: L4 interrupted; female
sterile
71.2 *dr droopy Wings spread, drooping
71*. 5 sf safranin Eyes brown
72 1(2 )me lethal(2) meander Larval lethal
72 *M(2)d Minute(2) d Heterozygote normal; Minute with M(3)d;
probably homozygous lethal
72.0 Lobe Eyes small, kidney shaped
72.0 N-2G Notch 2 from Gallup Wings cut; homozygous lethal
72.3 kn knot L3, L4 close or fused; crossveins abnormal
72.5 ch chubby fly short, thick
73.0 dke dark eye Eyes dark
74 6P gap L4 thin or interrupted
74 scrp scarp Eyes flattened, furrowed
74 *Su(t) Suppressor of Reduces expression of f; homozygous lethal
forked
75.5 c curved Wings downcurved, thin
76 *Wr Wrinkle Wings wrinkled, blistered
77.3 Amy Amylase Affects amylase electrophoretic mobility
77.5 M(2)S7 Minute(2) of Schultz Bristles fine; late hatching; homozygous
7 lethal
79 pw-c pink wing c Eyes light; wings short, blunt
brrA
cn
CJl
CD ^•.?3
o> -
HMJMMM
O <P 5S
r tl.-..-.4
m
80 it fringed Wing margins snipped

80. 5 tu-bw tumor with brown Larval and adult melanotic tumors
81 ij four jointed Tarsi four jointed; crossveins close
together
81 ri roof wings Wings sloped
82 *Off Off Bristles missing; eyes large, creased,
rough
82 wt welt Eyes small, narrow, horizontally seamed
83 abr abero Abdomen abnormal; eyes rough; bristles
sparse
83 nw narrow Wings long, narrow, pointed
83. 1 a dp adipose Adult fat bodies abnormal, hypertrophied
86. 5 *E(f) Enhancer of forked Bristles short, twisted; enhances f
87. 5 *M(2)b Minute(2) b Bristles fine; late hatching; homozygous
lethal
90 dsr disrupted Extra veins; wings warped
90 l(2)56a lethal(2) 56a Lethal
>£T
*5
m
91.5 sm smooth Abdomen shrunken, denuded; sterile

92.3 M(2)173 Minute(2) 173 Bristles short, thin; homozygous lethal
93.2 rw raised wing Wings upheld
93.3 hy humpy Thorax ridged; wings truncated
94 *su(B) suppressor of Bar Modifies B
96.2 cv-2 crossveinless on Crossveins absent
chromosome 2
97 Pu Punch Eyes purple; homozygous lethal
99 Elp Ellipse Eyes small, facets fused
99 l(2)Su(H) lethal(2) from Sup- Lethal
pressor of Hair-
less
99.2 a arc Wings broad, bent; crossveins close
100 *fs(2)K female sterile(2) of Female sterile
Kikkawa
100 *pkh pinkish Dilutes we
100.5 px plexus Wings have network of extra veins
101.0 pa patulous Wings spread
101.2 M(2)l Minute(2) / Bristles fine; late hatching; homozygous
lethal
crusterile Male sterile
crs
102 *tit tufts Sternopleurals increased
103.1 Body speckled black, tumorous; homozy-
Frd
Freckled gous lethal
104 *Bsl Wings blistered; extra veins; homozygous
Blisterlike lethal
104 1(2 )bw Lethal
lethal(2) with brown
104.0 hv heavy vein Veins thick, knotted; wings warped

r— 104.5 bw brown Eyes brown
/ r- 104.7 mi minus Bristles small; hairs reduced; female
\JI sterile
J—' 105.2 Su(bwV1) Suppressor of Modifies bw VI
br o wn-Varie ga ted
i
l
i— 105.5 abb abbreviated Bristles small
106.3 sit slight Body small; bristles short, thin
- 106.4 pd purpleoid Eyes maroon
— 106.7 mr morula Eyes rough; bristles reduced in size and
"V — 106.7 11 lanceolate
number; female sterile
Wings narrow at tips, slightly divergent
- 106.9 l(2)ax lethal(2) ax Early larval lethal
- 107.0 or orange Eyes orange
•1
- 107.0 Fo Forkoid Bristles short, bent; homozygous lethal
J — 107.0 *1(2)NS lethal(2) Nova Larval lethal
Scotia
L >— 107.0 sp speck Black speck at wing axil
- 107.2 Px Plexate Wing veins plexuslike; homozygous lethal
\ _ — 107.3 bs blistered Wings small, pointed, blistered; extra veins
- 107.3 Pin Pin Thoracic bristles short, sharply tapered
- 107.4 ba balloon Wings blistered, extra veins
107.6 If Irregular facets Eyes small, narrow, facets irregular
-^ 108 Kr Kriippel Thorax malformed; homozygous lethal
— 108 M(2)c Minute(2) c Bristles fine; late hatching; homozygous
lethal
CHROMOSOME 3
-0.1 su(ve) suppressor of vein- Suppresses ve

CD to > let
0 *she sherry Eyes sherry; sterile
O 0 *tyw tiny wing Wings small; extra bristles
0.0 aa anarista Aristae bare or tufted; wings broad
0.0 *bos bordosteril Eyes brownish red; female sterile
o . - 0.0 *mp micro pt era Wings small; veins irregular; legs abnormal
m U) - 1 0.0 mwh multiple wing hairs Wing cell hairs increased
0.0 rough oid Eyes small, rough
E 1 0.2
TU
ve veinlet L3, L4, and L5 incomplete

92 OB -
1.03 eyr eyes reduced Eyes small
> I 1.4 R Roughened Eyes rough; homozygous semilethal
1 1 A I A I 1 AiA
ro 3 *ven venation Veins thick, branched

> 11 fs(3) female sterile(3) Female nearly sterile; male partially sterile
12
of Gill 2
12 *dw-b dwarf b Body small
CD 14.7 *syn syndrome Eyes brown; wings spread; sterile
— > 11
ro -
O
- 1
1 III
m
<o- 1
> 1
rv>
> i
GJ
>
w >
O I
o> —
o *
m -
WjS*1
17 *rai raisin Eyes brown

-19. 2 jv javelin Bristles, hairs cylindrical
-19. 2 Me Moire Eyes flecked, iridescent; homozygous lethal
19. 6 *Vn Vein L4 incomplete; homozygous lethal
20 l(3)tr lethal(3) Pupal lethal
translucida
L-20.0 *dv divergent Wings spread
R >
"H I
23.0 Hn Henna Eyes brown; homozygous lethal

25 *be~3 benign tumor in Benign melanotic tumors
chromosome 3
25 fs(3)G3 female sterile(3) of Female sterile; oogenesis incomplete
Gill 3
26 *On Open Wings spread
26 suCO suppressor of tan Suppresses t
26.0 se sepia Eyes brown, darken to black with age
26.5 h hairy Extra hairs on scutellum, pleurae, head,
and along veins
27 a (3)2 6 abnormal abdomen Abdomen abnormal
in chromosome
O c
3 26
27 abd abdominal Abdominal bands broken, etched
28.9 *M(3)i Minute(3) i Bristles fine; late hatching; homozygous
lethal
30 cur curvoid Wings diverge, curve down
33 Ixd low xanthine dehy- Affects xanthine dehydrogenase level
drogenase
33.8 *l(3)Spl lethal(3) of Lethal
Spiess 1
m 35.0 *rs rose Eyes purplish pink; often sterile
35.1 gs gespleten Thorax cleft; eyes small
35.5 eyg eye gone Eyes and head small
36.2 *gv grooved Thorax grooved; eyes small; bristles irreg-
ular
36.5 *cr-3 cream in chromo- Eye color diluted, dilutes we
some 3
36.8 Est-6 Esterase 6 Affects esterase electrophoretic mobility
37 *rag ragged Marginal hairs irregularly absent
37 Tt rotated abdomen Abdomen twisted; sterile
37.5 app approximated Crossveins close; tarsi four jointed
38.0 *md melanotic lesions Melanotic lesions throughout body
38.4 *I(3)Spl7 lethal(3) of Lethal
Spiess 17
39 pyd polychaetoid Extra bristles
40.0 tt tilt Wings spread, warped; L3 interrupted
o 40.2 M(3)h Minute(3) h Bristles fine; late hatching; homozygous
lethal
40.4 e(dpv) enhancer of dumpy- Normal; enhances dpv
vortex
o _
40.4 *I(3)Sp6 lethal(3) of Lethal
Spiess 6
-si "H » -
o -
u»
o Ok
>
All
m
1 1 IV I I VI I I V 1 IV
oo OP
0) >
CD
>
O Locus Symbol Name Phenotype
Ly Wing margins excised; homozygous lethal

V 1 VIV I I 1 1 I V 1 V I I I V V I V I
40.5 Lyra
D 40.7 D Dichaete Wings spread; alulae missing; dorsocentrals
fewer; homozygous lethal
n 41.0 *1(3)SP5 lethal(3) of Lethal
CD
Spiess 5
41.3 Su(var) Suppressor of var Modifies variegation of w, rst, fa, sc, spl,
0) iegation nd, and y in variegating rearrangements
41.4 Gl Glued Eyes small, oblong, shiny; homozygous
o > lethal
CD
41.7 fz frizzled Hairs and bristles directed irregularly; eyes
rough
41.7 *l(3)SplO lethal(3) of Lethal
Spiess 10
41.7 *rp rotated penis Male genitalia rotated; eyes rough; male
sterile
42 wk weak Bristles, abdomen small; wings warped
o -
>
II 1 A i l A I A M 1
m
> 1
o > >
> > j
> {
w =
«
CD
m Locus Symbol Name Phenotype
> : 1la 43.0

43,2
43.4
*We
th
*mb
Washed eye
thread.
minus bar
Modifies w, homozygous lethal
Aristae lack side branches
Modifies B
GO u» —
rO —
43.5 *Cm Crimp Wings crimped, ruffled; homozygous lethal
O 43.6 bulge
bul Eyes large, bulging; wing margin thick
44 * vr varnished Eyes small, facets fused; female sterile
44.0 st scarlet Eyes bright red; ocelli colorless
44.3 M(3)S34 Minute(3) of Bristles fine; Jate hatching; homozygous
Schultz 34 lethal
44.4 db dark body Body dark; male lethal
45 *re-6 reduced eyes b Eyes small
45 tra transformer Female transformed into sterile male
45.3 cp clipped Wing margins snipped
46 as ascute Scutellum elevated; wings drooping
46 *je jelly Eyes pink
46 *ml minute like Bristles small; late hatching
46 *mot-36e mottled 36e Eyes mottled, rough; female sterile
46 Pdr Purpleoider Eyes maroon; modifies pd
46 tu-48) tumor 48j Larval and adult melanotic tumors
46.0 *mot-28 mottled 28 Eyes mottled
46.0 *W Wrinkled Wings unexpanded; head spotted
46.3 Aph Alkaline phos- Affects alkaline phosphatase electropho-
phatase retic mobility
46.5 *si-3 ski 3 Wingtips turned up with Si
47 *du dunkel Body dark; wings blistered; female sterile
47 (s(3)Gl female sterile(3) of Female sterile
Gill 1
OD
Ui —
1 1 1IIV
o
>
>
o OD -
_
O) > I
CD I
_>
O >
o -
47 in inturned Hairs and bristles directed toward midline

47 *M(3)S39 Minute(3) of Schultz Bristles fine; late hatching; body small;
39 homozygous lethal
47 Sex Extra sex comb Sex combs may be on all male legs; homo-
zygous lethal
47.0 ri radius incompletus L2 interrupted
47.3 eg eagle Wings spread
47.5 Did Deformed Eyes small or furrowed; homozygous lethal
47.5 *wp warped Wings warped, small, spread
47.6 Ki Kinked Bristles and hairs short, twisted
47.6 roe roughened eye Eyes rough
47.7 drb dark red brown Eyes dark red-brown
47.7 pb proboscipedia Oral lobes tarsuslike or aristalike; female
sterile
47.7 rotund Wings short, round; sex combs absent;
sterile
CP u , Z
03 *. =
47.8 *wz wizened Body small, dark

Antp Antennapedia Antennae leglike; homozygous lethal
48 Pc Polycomb Sex combs on second and third legs of
male; homozygous lethal
<o - 48.0 Msc Multiple sexcomb Sexcombs on second and third legs of male
48.0 Na Nasobemia Antennae leglike; homozygous lethal
<0 =
Q o.
09
CD
o> —
m -
u>
• 48.0 P pink Eyes dull ruby

48.1 dsx double sex Male and female resemble intersexes
48.2 *Kg Kugel Body short, thick
48.3 bod bowed Body small; wings curved downward
48.3 *moo moorish Body black; male lethal
48.5 *com compressed Head flattened ventrally
48.5 tet tetraltera Wings halterelike; mesothorax like meta-
thorax
48.7 by blistery Wings blistered, warped, dusky
49 Est-C Esterase C Affects esterase electrophoretic mobility
49 fs(3Xr5 female sterile(3) of Female sterile; oogenesis incomplete
Gill 5
49.2 Odh Octanol dehydro- Affects octanol dehydrogenase electropho-
genase retic mobility
49.5 *Rst(3)ns Resistance(3) nico- Resistant to nicotine sulfate
tine sulfate
49.7 ma maroon Eyes ruby
50 dn doughnut Light spot in eye
50 *dw dwarf Body small; female sterile
50 *Er Erect Scutellars erect; wings unexpanded
50 *mu mussed Wings thin; thorax arched
50-0 M(3)S31 Minute(3) of Schultz Bristles fine; late hatching; homozygous
31 lethal
,-- j
m
o-
cu curled Wings upcurved; body dark

Hu Humeral Extra humeral bristles
1(3)S1 lethal(3) of Schalet Lethal
1
*ttr tetrapter Halteres winglike
2
w - u. kar karmoisin Eyes bright red
3
mes messy Extra bristles on head and thorax; wings
inflated; sterile
ry rosy Eyes reddish brown; lacks xanthine dehy-
drogenase
l(3)S4 lethal(3) of Schalet Lethal
4
5
pic piccolo Bristles short, fine; tergites abnormal;
sterile
1(3)26 lethal(3) 26 Lethal
6
•n ^
7
CD red red Malpighian Malpighian tubes red; eyes brown
*4 5- tubules
OD >
OD > cv-c crossveinless c Posterior crossveins absent or reduced
> su(Hw) suppressor of Hairy Suppresses specific alleles at various loci;
wing female sterile
56 *sh short winged Wings small
03 = te=2S
O =
Q =
O—
LOCU! Symbol Name PKenotype
jvl javelinlike Bristles and hairs cylindrical

c(3)G crossover sup- Eliminates meiotic recombination
pressor in chromo-
some 3 of Gowen
58.2 sbd stubbloid Bristles short
58.2 Sb Stubble Bristles short, thick; homozygous lethal
58.3 *Two-b Two bristles Two postverticals absent; homozygous
lethal
tuh-3 tumorous head in Head tumors with tuh-1
chromosome 3
ss spineless Bristles hairlike
bx bi thorax Metathorax mesothoracic; halteres enlarged
Cbx Contrabithorax Mesothorax metathoracic; wings small; ve-
nation incomplete
Ubx Ultrabithorax Halteres enlarged; homozygous lethal
bxd bithoraxoid Metathorax mesothoracic; halteres enlarged
pbx postbithorax Posterior metathorax mesothoracic
mfs(3)G male and female Male sterile; female semisterile
sterile(3) of Gill
Rf Roof Wings rooflike
*Su(sc) Suppressor of scute Suppresses sc
Me Microcephalus Kyes small or absent; scutellars curved
*cal coal Body black
*wtl weltlike Eyes seamed, small; female sterile
fl fluted Wings creased, dark
Su(ss) Suppressor of Suppresses ss; homozygous lethal
spineless
sr stripe Dark median stripe on thorax
61 glass Eyes small, diamond shaped, glassy
*gt-3 giant in chromo- Body large; late hatching; male sterile
some 3
k kidney Eyes kidney shaped
*M(3)S35 Minute(3) of Schultz Bristles fine; late hatching; body small;
35 homozygous lethal
*cv-b crossveinless b Crossveins absent
cv-d crossveinless d Posterior crossveins absent
*sprd spread Wings spread
Cu-3 Curl in chromo- Wings curly, thin; homozygous lethal
some 3
66.2 Dl Delta Veins thickened, broad at margin; homo-

zygous lethal
— \~»* i •> i 67.3 *gn glisten Eyes rough
O» - 68.5 *com-d compressed dilap- Fly small; legs and wings defective
idator
69.5 H Hairless Bristles sparse; veins incomplete; homo
zygous lethal
70. 7 e ebony Body black
72 *bn band Dark transverse band on thorax
72. 5 det detached Crossveins detached or absent
O> —
-n -
4^ —
75.5 *dg-a degenerated sper- Spermathecae degenerated

matheca
75.7 cd cardinal Eyes yellowish vermilion
76.2 wo white ocelli Ocelli colorless; modifies we
o»-
77.5 obt obtuse Wings short, blunt; thorax humpy
78 *Gd Gulloid Wings short, blunt, spread; homozygous
CD E
lethal
79.1 bar-3 bar on chromosome Eyes small, narrow
3
79.3 lethal(3) of Spiess Lethal
<o — 2
79.7 Minute(3) w Bristles fine; late hatching; homozygous
lethal
o i 81.6 1(3 )a lethal(3) a Lethal
84.5 Minute(3) beta Bristles fine; late hatching; homozygous
lethal
88 mah mahogany Eyes brown
m -
o=
>
OD *
O =
D >
m >
OD-
^1 >
CD >
>
90 groucho Bristles clumped above eyes

90 *Su(y3P) Suppressor of Partially suppresses y 3P
yellow-3 of
Patterson
I\»Z
90.0 Pr Prickly Bristles short, tips thin, twisted
90.2 1(3 )PR lethal(3) PR Lethal
90.2 *M(3)j Minute(3) j Bristles fine; late hatching; female sterile;
homozygous lethal
<£> 91 tx taxi Wings held out
91.1 TO rough Eyes rough
91.8 !(3)XaR lethal(3) XaR Lethal
92.5 Ser Serrate Wingtips notched; homozygous lethal
93 cmp crumpled Wings small, crumpled, or blistered
93.8 Bd Beaded Margins excised; homozygous lethal
94 *Ble Barlike eye Eyes small, narrow
94.1 *Pw Pointed wing Wing tips narrow; extra veins; homozygous
lethal
95 bf brief Body, bristles small; male sterile
95 *M(3)d Minute(3) d Heterozygote normal; Minute with M(2)d;
O »
homozygote probably lethal
95.4 rsd raised Wings upheld
o» — 95.5 su(pr) suppressor of Suppresses pr; enhances Hw; sterile
purple
97.3 ra rase Bristles and hairs small, irregularly absent
98.3 Lap-D Leucine amino Controls leucine aminopeptidase D electro-
peptidase D phoretic mobility
99.2 Dr Drop Eyes small; homozygous lethal
100 *wdn wings down Wings spread, drooping
<£>
m -
r- 100.7 ca claret Eyes ruby

/ 100.9 *l(3)Spl9 lethal(3) of Spiess Lethal
/ 19
1 101.0 M(3)l Minute(3) 1 Bristles fine; late hatching; homozygous
lethal
101.1 *l(3)Sp9 lethal(3) of Spiess Lethal
n
y
101.4 Acph-1 Acid phosphatase 1 Affects acid phosphatase electrophoretic
mobility
102 Id loboid Eyes small, kidney shaped
102.7 bv brevis Bristles short, stubby; body chunky
102.9 K-pn Killer of prune Normal; lethal with pn alleles
105 *M(3)[ Minute(3) f Bristles fine; late hatching; homozygous
lethal
106.2 *M(3)g Minute(3) g Bristles fine; late hatching; homozygous
lethal
CHROMOSOME 4
Locus 4 Symbol Name Phenotype
ar abdomen rota turn Abdomen twisted, male sterile

Ce Cell Wing veins abnormal, homozygous lethal
0 ci cubitus interruptus L4 interrupted
M(4) Minute(4) Bristles fine; late hatching; homozygous
lethal
0.2 gvl grooveless No groove between thorax and scutellum
1(4)25 lethal(4) 25 Larval lethal
*Scn Scutenick Scutellum short, edge nicked; homozygous
lethal
1.4 bt bent Wings bent back
2.0 ey eyeless Eyes small
1(4)9 Iethal(4) 9 Lethal
3.0 sv shaven Bristles small
spa sparkling Eyes rough, bulging
*As measured in diplo-4 triploids (Sturtevant, 1951, Proc. Natl. Acad. Sci. U.S. 37: 405—
07).
-ARISTA
-ANTENNA
-ORBITALS (A.MJP)
-OCELLAR
JS
:ALS (A,P)
- - PO9TVERTICAL
PROBOSCIS
--HUMERALS (U,L}
-PRESUTURAL
STERMOPLEURALS
» CA,M,P) NOTOPLEURALS (A,P)
SUPRA-ALARS (A,«
> DORSO-CENTRAL3 CA,P)

MESOSTERNAL
_ - POST-ALARS (A,P)
SCUTELLARS (A ( P)
1MM
cf
' I 1 1 1—i 1 1 1 ii,

I 1 MM l>
LEGEND:-
HALTERE'
-- SPIRACLE A ANTERIOR
ABO - ABDOMEN
Cl, C2, C3 - COXAE
PENIS HA--HALTERE
HP--HYPOPLEURA
GENITAL ARCH
HU --HUMERU5
6T
ANAL PLATE L LOWER
-VAGINAL PLATE M MIDDLE
ANAL PLATE
MN--ME5ONOTUM
MS --ME5OPLEURA
MT - -METANOTUM
N - -NECK
COSTA,OR MARGINAL VEIN P POSTERIOR
ALQdLIARY v VEIN PT --PTEROPLEURA
S STERNITE
51,52-THORACIC SPIRACLES
SC - - SCUTELLUM
5T --STERNOPLEURA
T TERGITE
U UPPER
W WING
JS
182 200 2iO 215 27.7 330 3 6S 387 40.7 430 444 505 54.5 56.7 570 595 62i> 660 S6.G
0000*00** 0J 06 06 ft 30 55 75 137 •KEY TO REFERENCE SYSTEM FOR
y ac sc tv.* br pn f f« cm ct m i fw sd r f :B fu t>b s p - f
H-U-^ L SALIVARY CHROMOSOMES

10 20 30 50 56
0
DIVISION^ NUMBERED >M C-?CJ?S V 2C ( i - I O i -
Esc is begins wirh a nujer sjns
SUBDIVISIONS. LETTERED'fU f;ROUPS 0? £ f.A-F)

Each fengins vntr. J sra'p ar.s
;
s ?c* & f ; r A %l C b i l l f C <&T t A I 8 ' C 0 < E IF ' . C M - f 1 A i B i C 1 D AIfti C i 0 t *E At 'B cm E * B " c ! Di E I r A , B Icio [ I r A ! B ICIDIr i ? A B c iol t I F ' A B C I o E ' F A U C IDIEWAI s i c ! a i E i F k i s l c i o l E l F i A ! s f c i 9 i E U a k c b ! t I F L I e i c i e
l
i 314 415 56 819 0 10 M IIH2 12 13 13 14 14 15 55116 !6?r7 17118 ISIJ9 19120
19(20
130 16.5 310 410 485 510 545 550
0.0 13 i
ai S dp J b ii I-X
20 30 40 50 55
2L 2L
IMM;; i»
r ' i n T I 3L
E If \ 8 lool E' I F ! A I Bl C I 0 FFAl B C 0 E» F A I B C I D I £ I F Ai B I d 0 1 E ^ Al S iciOf t I F J A I B I C UE E If I{AIBICID!
f ( A ! S ! C ! D !EEi iFf j
4 ••
3132 13
32133 33134 34 35
34 35 36 36137 37138 36(39 39&0 40!4l
4054
82.0 87* 933 99.2 1005 '04 5
'J wt
SCALE ^ "Oj—H
*>*> 80 90 ICO
2R
4034} 4142 42143
is: I if
431 45
nil
'4SI4f
i
if ks* c i»I c4114?
*rl * I i k i a Ii IF Wife! * 11
AI8!CI D
52 53 53 54
A! BIC
•If:
E iF
54 55
Ai B i c ! s ! E "r
f
55156
ABSCI 3 i Z 1 f ] A:.
56157
£
57!58
.1 t i c ! & :F • • i
58 59 53:80
•>.('
54 41?
35 37 4Q4 422 440 46±£60
cur rt D th St r. .p-f
"T" L
20 30 40 46
3L r
i i ; r '•
.,;•» <! * i
^
!l
,»!rln« »t,
if
M' ')
i 7 ! '
; I i •
?
i|> E 'f ' i i I C I D I C AI B Ir I M - ' • I A i a l c b , L I F U B I ' c I D I . - . f |.*.I
' 'fi
$4*15
l til Wter 69 70 70:7! 71 72 72i73 73l74 " 74175 75176 76jT7 77:78 78179 79-?
•>.. • ! •
©5 70.7 757 80 SC 93
ty ft
m H « CO Pi
-r-
70
»V f A
\wwfipj;
I if A F A 1 S
mwm-mi
£ IP IAI S t i n At 8 I C 151 E i r I i t MM MM
i: a JC 5 1 ;
'#.«; rfcllflfll i } ? •'
87188 9»2 V-?*
to Jmtrmd of Heredity, V©1. 26» N©. 2, Fe&rastiy, 193S REFERENCE MAP OF THE SALIVARY CHROMOSOMES OF DROSOPHILA MELANOGASTEE
Cojryr&kl 1933, by th* dmmkm Gmmit Aumktim.
SALIVARY l
X-CHROMOSOME S 'I
SCALE= •i—t-
BAWD CHARACTER llAAlAAIIAAl IA IAI|»IAIA 11 |AA I I(AI nun AID! Ml II A S A I A H A A AAAl A AlAlAfA Ai UUIIAAA A A A A A
LINE NUMBERING I234-* 69' 1 2 " 123*567-90- 67 §2*45 12-45' n -in?-Hift
K-45-
SUB-DIVISIONS C ! D F A B c D F A B C
D VISIONS 3 3
AA IS I A I A Ml|AlA ! A i A I IAJAI I AAI AAAAI A II I IA A ! A A I I I I A I A I A A 1 f A I t J WAHAA A A I A A I A mi i AA AAAAIA AAA AH A l l AA A ^{JAIAAI A I A A AA AHA 1AA1 AA1 AAAAA AI ii i U AA II AiAtAKI]
fl2*4 - 6 7 ' «»'tt-JK*45- I234547S 12-45 56789- •56« *9lHt234 12-45'769 WHI14* fti?|tf45"89Diielt234 1Z-4J67-9B \2M5 234- 123- 1E456789— 1234- • 2-4*6 II
A C 1 D B c D E c D E F A C D F A B c D F
6 9 10 10
l A f A A l AMA A A I A If AAJI AA A i l A ft A l ^ A i S AAA IA A A y\AI AA A AlAAAAiJAIAAIilAAAA A A A A

•#i«J234 123456-WIHb J2-45K3l-789-*l£eiM- 123456 7MW-HB" 1134567-
C D 1 E F I A B
13!I 14
TOTAL LENGTH =4I4JU

REPRODUCTION 3100 DtAWETEK
! !A I A kAAA AAAA A A A i A 'Mi A t M / ^ A M A Al AHA A I A A A A A I Al A I A A I AiA

:
? !I2J45k» 7ft I23tffc-9Oa 12 45' • 4 2 M - A 7 J2Z34 ' 12945 B8456 123 U .4..: it- 12 M--78 12 3 12 3 45 123 12 12 \Z 34 TOTAL LINES « 1024
p ! r%
B S C D E i F A B A BIC D F A B CD E F
16 7 17 19 19 20 ZO
Supplement to Journd of Heredity, VoL 29, No. 1, January, 1938. REFERENCE M A P OF THE SALIVARY QLAND X-CHROMOSOME OF BROSOPBIIA MELANOGASTER.
C&pyrifkt, 193B, hy thug dm*ncm Genetk A$$m%Mmm.
SALIVARY
2L-CHROMOSOME
SCALE * 5|4 I 1
1
Al if ill At* (Ml At H A H A I I All: A l I A t AHA ti AN! Aiur II A A l l Al I! A IIA 1 1 All! AH 1 Al AAIAI A l A 1 I A! A I! A I I! A 1 A 1 AA1
BAND CHARACTER
a 10 10 6 5
LINE NUMBERING 3 a 9 4 4:
a 0 9 a
D 5 D B C D E F
SUB- DIVISIONS, A B B
DIVISIONS ,..«-.. 21 22 22 23 23 24 24 25 25
A I i IA f S!
6
F
30
I I
ll
I
!Iff
v# * \
A l t I A A! ! II f A h A t f A I A i A t I f A I A t ! f IAI A A HA I I A l l A I I I II A H I A S I A I A tA11 A M U ! I AIAI AAI All A I H A At IA! AA I I I A I I A A 1A I ! A 1 ! if

inn
S 10 I2J 7 al 10
!A B
5
D B D B 1C 0 B I C D B D ^1
3il32 32 33 33 34 34 35 35!
,1
I 1 M* tR«
LENGTH
3100 01 AM.
3?0
1 I MA !A t A l l H ' A M i l l A M 1 Hi IAI AHA IMA I A I hi I A I I I I A l l ! HAM M HI? M l

A" A IA A I*AA I M A M I A I AIJAIAf!
1 If! 80S LINES
if 3l 01 t3 7 5? 6 a! a 10
A B mE F B D E!F A i B
D
36 37 37 38
REVISED REFERENCE MAP OF THE SALIVARY GLAND 2L CHROMOSOME OF DROSOPHILA MbLA\OG.4$TER.

Supplement to Journal of Heredity, Vol. 33, No, 11, November, 1942.
Copyright, 1943, by the Americm Genetic Aitociation
H-
BAHD CHARACTER. SI!! 1 SI 'A 1 M
LJI4E NUMSE8f*JG. •; 2 i-3 3 -6
r
SUB-DIV ISiOMS . F
• <: i ' A * ' îi|ts!.a' tt A • :

ktWVkihlMljMlll HI! ACIlAil AAJ 1 A WWII •Uf
B € D ! E 101£i f 8 , C f
48143
i •; # i"
1 ife t i h: A > t AI: lit A t SIA

IftsaUS;Aia!|AiSI A?
D
0 A
51 53
*i -. TOTAL L E M G T H * 4 4 6 M
? $ *;,
•>*< I
A **"* . ; * . *
*Us
' • * . * -
:,j- ;f.--- 4 *
.•*/'»'• * -
A
A
MAP 4 * TH*
4 W.i
SwppleuraeRt lo Journal of Heredity, Vol. }©» N©» 11, N©w»Wf» If >f
SALIVARY
I I
31.-CHROMOSOME
SCALE- Sju
BAND CHARACTER _ 1AH I Al A I 1 AtAIA I A U At 111 I I t|A f I Si i i t M MA I A I Hill 1 1 1 i A I I I f All II I tAIA l|A IA I /*1t.*
U-HE NUMBERING „»» 3
B Cl 0 F c 0 A c
DIVISIONS —
fl I
11 i
l
AAi I A |A I I i IA Ii t i A l ! A i A i !ft » • / 4 ' *' ' ! I ki i i i A M M IAA I A I i A Ml t A 1
15 5; 5j 6 S3
| s I
A I BiC1 D I E F B c D 1 E iFl A I B E F
G5 66 67168
Ml'IA i !A * A JJis;i II 111 f A I J A M A h A I f l l A l i ^ •» A

6! 1*1 4 € 1 4!
Al 8 C ' D i E I F A • B c In r A !B;C E IF A c F'

69 69 70 71 72 72173
i i
* i
i i
»iU » ' \ I 1 . > * ,-M.ft f ;* ' ' : 4 , f f.

A > 'A ! 1 < A A. ^ ,
A :
B :
C i D ' t n
77 pi
to Journal of Heredity. Vol. 32, !No. 2, February, 1941. REVISED REFERENCE MAP OF THE S
€t»p\fighs, 1941, by the Amtruan timeta ,4ttot>alion.
SALIVARY II % i I
3R - CHROMOSOMES?;^fl *w ' I
I! •• ' I
SCALE ~ 5« F—
I' « I I
Mi if u is »
CHARACTER »AAAAI A t A l A i l S I iiifeiii i IM iMAIii ik t MllllltiAMlAIIIIII an A I A I Aim At AIMS UII i I I UAH A i l t A I I I A I A 1 IAIAH HI IAA 1? A 1 h 11 IA I II IA 11 ! 1 I A I I A I A A I A S! A Si A I A 1 A I i i A S I ? MAM I JUi IfjAUliMIU J §
LINE NUMBERING 4 5 a 14 13 1! 9 13 15 s 15
SUB-DIVISIONS A B D F A B D 8 D A 8 C 0 E A 1B C
DIVISIONS 80! 81 82 82 83 83 8 4 8 4 85 85J86
I '/I
l
NI •«
IA 1 Al IAAII AIAI I HI ASA I I A I si i s i AA I I It I I I i i i S SI I I 11 A i n n n i i i MM i! ih t AimiAini Alii k }AA1IAIA 1 1 h\\ K\ U l in kh I

20 Ii 10 ii 10 13 9 13 221 7 13 4. 71 a 10
F A B D E I F B I C D F A ! B c ID! E F
86 8? 87 88 88189 89190
At I A I I 1 * (At i l l , A t H A ! I A l l f I A t ISA I Ml A I A \ A l l ! ! » l l A t t ii f A I I miAl 1AI I IAIAH kt I t h I 1 A 1 I lAI A I I I M )IA • I A <AM \ \m\t « A ! I * A I AiAMAH
8 5 * 13 14 9 15! 7 13! to1; 14" 16 If

B Dl E i D A B C D i F j A IB
91 91192 93 93:94
n \
!)I III » * $\ 4
I in LEWSTH
Ilfl A I A I A I A A I Ml It A I M MIAA1A » |Al Ii A * H »». i M ?»HAH k t pn p, f"4»*i

6! 12'; I4i W 10? 5 1 IS! It? ill ti if 5; If f f l f i UMCS
i 15
F 8 C F 1 J I B ! C I 0 iEi F i A i B l
95196 96!97 ffliOO
Su|ipiem<>ni lo Journal of Heredity^ Vol. 32, No. 9, Seplerafeer, 1941. REVISED REFERENCE MAP OF TOE SAUVARY GLAND » CHROMOSOME OF m0.$OPMHA MBLAXOGASTSR.
Copyright, 1941, by tht Ammum Gmmk A$$mwtim.
t
1
"I ! I
•={
Ii
UAflll I A A A I A 1 A 1 » ^ A A I A ' ^ lift MIA MAA
ft 5v S2 54S 47 85 ft I E HS I? 12-st • rail «' il
A B iC D : E IF A B I c D
101 101 I0Z 102
Scale • IOJUL. Tota^l bands 137
t,... ... .... .. . , . f lotaI lenfth 46
MAP OF CHROMOSOME 4

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BIOLOGY DIVISION, OAK RIDGE NATIONAL LABORATORY

Carnegie Institution of Washington Publication No. 627

THE MUTANTS OF DROSOPHILA MELANOGASTER

Calvin B. Bridges and Katherine F. Brehme

Carnegie Institution of Washington Publication 552, 1944

Genetic Variations of MUTATIONS

Multiply Marked Chromosomes

DEPARTURES FROM DIPLOIDY

WILD TYPE STOCKS

Vn/T(Y;3)1 GENETIC AND CYTOLOGICAL MAPS

a: arc phenotype: Nearly normal; distinguished by its

origin: Spontaneous. Probably a derivative of aba3.

aba: arc-broad angular *abadb: arc-broad angular dumpy blistered

*a(1)48: abnormal abdomen in chromosome 1 *A(2)51

ab: abrupt cytology: Placed in region between 59E2 and 60B10

abr: abero synonym: Called ac11 by Serebrovsky.

rental enzymes as well as a hybrid enzyme of in- ^ 30 -

evidence of intoxication within 1 hr of being adpfs: adipose-female sterile

origin: Spontaneous. *amy: amethyst

discoverer: Kikkawa. Amys: see Amy2-6

suppressed in X/X/Y female. Homozygous lethal. origin: X ray induced.

often rotated. Ax/+ female shows short L5 in half

origin: Ultraviolet induced.

Bridges and Morgan, 1919, Carnegie Inst. Wash.

B*: Bar of Sfone

bb$: see bb and bbda is a property of the 66 locus and bb }4

references: 1925, Genetics 10: 117—47 (fig.).

origin: Spontaneous, presumably through unequal

discoverer: R. C. King, 49h. cytology: Associated with In(3L)Bit; breakpoints

Male viability impaired; females almost com- origin: Spontaneous.

*Bsl: Blister I ike

*bu: bulging phanotype: Eye color reddish brown; brilliant

phenotype: Homozygote not clearly distinguishable

: brown-Auburn phenotype: Like bw except testis sheath pale

cytology: Associated with In(2LR)bwR2° = references: 1955, Genetics 40: 5—23.

origin: X ray induced. origin: X ray induced.

origin: X ray induced. discoverer: Winchester, 1932.

phenotype: Like bwVl, RK1A. except in the presence of an extra Y.

origin: X ray induced.

origin: X ray induced.

47: 135—99 (fig.)]. Thoracic musculature studied

origin: X ray induced.

origin: X ray induced in ss bx Su(ss)2.

phenotype: cav/ca slightly variegated. Can be *Cb: Curled blistered

*CCY: chlorotic-yellowish discoverer: Glass, 39a28.

c/ + 2; cubitus interruptus-wild-type isoallele

references: 1961, Evolution 15: 239-46. synonym: It: Interntptus.

Clipped wings: see Cli More precisely located by Hannah-Alava in 6E,

*c/v-7; cloven thorax no. 7

*com~d: compres sed-dilapida tor corr; corrugated wing

often diverge. Eyes large and slightly rough- Cream: see Cr

crossveinless; see cv *csk: costakink

origin: X ray induced.

phenotype: Males have notched but unexpended origin: X ray induced.

cytology: Associated with Df(l)ct14cl =

pp. 588-89. origin: X ray induced in In(l)scslLsc8R+dl-49.

cytology: Associated with T(l;2)ct268~33 - cfK; cut of Krivshenko

Cur: see Cu-3 cv-c: crossveinless-c

a(1)48: abnormal abdomen in chromosome 1 A(2)51

l(2)pup: lethai(2) pupal l(2)S3a

Ms wlf% D f<"<•>,far <f»f , cyteAeffy: SaSiary cfere?r2%ea''s apparently ttenm!,