TAPROBANICA, ISSN 1800-427X. October, 2012. Vol. 04, No. 02: pp. 69-76.
© Taprobanica Private Limited, 146, Kendalanda, Homagama, Sri Lanka.
BOLBOCERATINE SCARABS OF GENERA Bolbohamatum KRIKKEN,
1980 AND Bolbogonium BOUCOMONT, 1911 (COLEOPTERA:
GEOTRUPIDAE) FROM CENTRAL INDIA
Sectional Editor: Oliver Hillert
Submitted: 26 March 2012, Accepted: 26 July 2012
Kailash Chandra1 and Devanshu Gupta2
1
Zoological Survey of India, New Alipore, Kolkata 700053, West Bengal, India
E-mail: kailash611@rediffmail.com
2
Zoological Survey of India, Jabalpur 482002, Madhya Pradesh, India
E-mail: devanshuguptagb4102@gmail.com
Abstract
This study includes a taxonomic account of four species of genus Bolbohamatum; B. calanus
(Westwood, 1848), B. phallosum Krikken, 1980, B. marginale Krikken, 1980 and B. laterale
(Westwood, 1848) and one species of genus Bolbogonium; B. insidiosum Krikken, 1977 from Central
India (Madhya Pradesh and Chhattisgarh). The pronotal ornamentation and external male genitalia of
Bolbohamatum species has been diagnosed with the incorporation of an identification key to the
species from Central India. A checklist containing 19 Indian species of both genera (Bolbohamatum
and Bolbogonium) has also been prepared with their distribution in different states of India as well as
outside of India.
Keywords: dung beetles, pronotal ornamentation, external male genitalia, distribution, India.
Introduction
Bolboceratine
scarabs
in
the
family
Geotrupidae are commonly called Earth-boring
dung beetles because adults of most species
provision larvae in earthen burrows with dead
leaves, cow dung, horse dung, or humus. The
family Geotrupidae currently includes 620
species belonging to 68 genera in three
subfamilies; Taurocerastinae, Bolboceratinae
and Geotrupinae (Scholtz & Browne, 1996).
The first comprehensive study of Asian
Bolboceratinae was carried out by Westwood
(1848, 1852), which considered 29 species to
be in one genus Bolboceras. Later, several new
species names were added based on the
materials from tropical and eastern Asia.
Boucomont (1911) proposed Bolbogonium as a
subgenus for Bolboceras. A series of
taxonomic
publications
on
Asian
Bolboceratinae were then made by Krikken
(1977ab, 1978ab, 1979, 1980, 1984), Carpaneto
et al. (1993), Masumoto (1984), Li et al.
(2008), Nikolajev (1979ab, 2003, 2008), Ochi
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BOLBOCERATINE SCARABS OF CENTRAL INDIA
& Kawahara (2002), Ochi & Masumoto (2005)
and Ochi et al. (2010, 2011). Krikken (1977a,b)
raised the subgenus Bolbogonium to the genus
level and described seven new species, along
with producing a key to all ten Asian species.
Subsequently, Krikken (1980) proposed the
genus Bolbohamatum for four species to be
combined with Bolboceras, while also
describing nine new species and discussing the
significance of external male genitalia and
pronotal ornamentation in the accurate
identification of the various species. Recently,
Karl et al. (2006) catalogued Bolboceratine
scarabs of the Palaearctic region. The present
study includes taxonomic information for four
species of Bolbohamatum and one species of
Bolbogonium from the Madhya Pradesh and
Chhattisgarh states in India and also
incorporates new distributional records of these
beetles. A checklist of both genera from India
is included.
Results and Discussion
Four species of the genus Bolbohamatum; B.
calanus (Westwood 1848), B. phallosum
Krikken 1980, B. marginale Krikken 1980 and
B. laterale (Westwood 1848) and one species
of genus Bolbogonium; B. insidiosum Krikken
1977 were studied from the states Madhya
Pradesh and Chhattisgarh. Bolbohamatum
calanus, B. laterale and B. phallosum are
recorded for the first time from Madhya
Pradesh, while Bolbohamatum marginale and
B. calanus constitute new reports for
Chhattisgarh. The identification of these
species is based on the structure of external
male genitalia, pronotal ornamentation and
clypeal dentations, which are shown in figures
1 to 9. Bolbogonium insidiosum shows
variations in the structure of clypeofrons (Fig.
9). The checklist for 19 Indian species of both
Bolbohamatum (11 species) and Bolbogonium
(8 species), along with their distribution within
and outside of India, are provided in Table 1.
Materials and methods
Specimens for the study were collected using
light trap from various protected areas by
scientific teams of ZSI based in Jabalpur,
Madhya Pradesh. Pinned specimens were
identified with the help of available taxonomic
revisions of the studied genera (Krikken,
1977b, 1980). Specimens were examined under
a binocular microscope (Leica M205 A) and
photographs were taken with the help of an
attached digital camera. Male specimens were
dissected, with the abdomen separated from the
body and the aedeagus extracted from the
abdomen. The genitalia were then cleaned and
softened in a dish of hot water and further
cleaned in a hot water solution of 10% KOH.
All parts of the aedeagus were washed in 95%
ethanol and photographed. After examination,
the genitalia were stored in a glass vial
containing 70% ethanol.
Systematic Account
Family: Geotrupidae Latreille, 1802
Subfamily: Bolboceratinae Mulsant, 1842
Tribe: Eubolbitini Nikolajev, 1970
Genus Bolbohamatum Krikken, 1980
Bolbohamatum Krikken, 1980: 5 (Type species:
Scarabaeus cyclops Olivier, 1789: 60)
The genus includes the species, presenting one
of the largest Bolboceratine scarabs which are
distributed in both the Palaearctic and Oriental
geographic regions. It likely evolved on the
Indian subcontinent and spread at a relatively
late stage through Myanmar into Sundaland and
China (Krikken, 1980).
Generic diagnosis: Metasternum anterroiorly
always with a small spiniform protrusion and
with anterior lobe narrowly separating middle
coxae. Head of males with a pair of tubercles
on clypeus. Pronotum in case of male possess
median and lateral protrusions with the surface
between them usually concave. Fore tibia with
7-10 external denticles.
The details of specimens examined, registration
number of specimens, distribution inside and
outside India, main diagnostic characters,
description, illustration of external male
genitalia, and identification key to the species
level within the genus Bolbohamatum are
provided. The classification adopted in the
article is after Smith (2006). Identified
specimens were deposited in ZSI, Jabalpur,
Madhya Pradesh (India).
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Identification key to the species of Bolbohamatum Krikken, 1980 from Central India:
1. Lateral tubercles of pronotum well developed but not marginally situated. Apex of parameres
not with reflexed paramerites …………………...………………………...………………….. 2
Lateral tubercles of pronotum well developed or completely reduced or absent if present then
marginally situated. Apex of parameres dorsally with short reflexed paramerites
…………………………………………………..……………………….……………………. 3
2. Dorsally the parameres moderately sclerotized, relatively narrow and with poorly developed
paramerite. Ventral side of parameres devoid of distinct paramerites. Basal capsule relatively
narrow ………………………………………...……..……………….. Bolbohamatum calanus
Dorsally the parameres foliate and ventrally with a pair of more or less glider-like
paramerites. Basal capsule in lateral view distally strongly emarginated ……………...………
…………………………………………….……………………….. Bolbohamatum phallosum
3. Paramedian tubercles of pronotum closely approximated and separated by less than to interocular distance while lateral tubercles well developed and marginally situated
……………………………………………………..…………….…. Bolbohamatum marginale
Paramedian tubercles of pronotum not closely approximated and separated by more than
inter-ocular distance while lateral tubercles absent ………………….. Bolbohamatum laterale
throughout. Paramedian tubercles separated by
less than inter-ocular distance. Lateral
impression of pronotum shallow.
Bolbohamatum calanus (Westwood, 1848)
Bolboceras calanus Westwood, 1848a: 385,
(description, distribution).
Bolboceras tumidulus Westwood, 1852: 22,
(description, distribution, illustration).
Bolbohamatum calanus Krikken, 1980: 20,
(description, keyed, distribution, illustration, comb.
nov.).
Specimens
examined:
Chhattisgarh:
ZSI/CZRC-A/16601; male (Length: 15.0 mm;
width: 10.0 mm); Barnavapara camp,
Barnavapara Wildlife Sanctuary, Raipur (21o
24.00’ N, 82o 25.314’ E; alt. 303.8 m); K.
Chandra & party, 01 July 2011, light trap.
Madhya Pradesh: ZSI/CZRC-A/16602; male
(Length: 16.0 mm; width: 9.0 mm); Forest Rest
House, Bandhavgarh National Park, Umaria, K.
Chandra, 10 August 2005, light trap;
ZSI/CZRC-A/16603; male (Length: 17.0 mm;
width: 9.0 mm), Karmajhiri, Pench Tiger
Reserve, Seoni; K. Chandra, 13 June 2001.
Diagnosis: (Fig. 1). Brown, shiny and pilosity
yellow brown. Cephalic tubercles more or less
dentiform, isolated and placed simply on
clypeal disc. Dorsal outline of left mandible
sinuate lobate. Pronotum with a pair of feebly
developed, slightly transverse median tubercles
with lateral callosities. Pronotum abundantly
punctate, but never densely punctate
Figure 1: B. calanus (scale: 5 mm), ZSI/CZRCA/16601, Barnawapara Wildlife Sanctuary, 2011.
External male genitalia: (Fig. 2) Dorsally, the
parameres moderately sclerotized, relatively
narrow and with poorly developed paramerite
while the ventral side of parameres devoid of
distinct paramerites. Basal capsule relatively
narrow.
Geographical distribution: India: Assam,
Bihar, Chhattisgarh, Karnataka, Madhya
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view, the basal capsule distally strongly
emarginated.
Geographical distribution: India: Madhya
Pradesh, Maharashtra and East India.
New state and district record: Madhya Pradesh
(Seoni and Mandla).
Remarks: B. calanus (Westwood, 1848) and B.
phallosum Krikken, 1980 show close
resemblance in their morphological characters
and cannot be separated on the basis of external
characters only, but the phalli of both the
species are very different and only the
characters of the phallus distinguish both the
species.
Pradesh, Maharashtra, Tamil Nadu, West
Bengal and Uttarakhand.
Elsewhere:
Bangladesh and Java.
New state and district record: Chhattisgarh
(Raipur) and Madhya Pradesh (Umaria and
Seoni).
Figure 2: Dorsal & ventral view of external male
genitalia of B. calanus (scale: 2 mm), ZSI/CZRCA/16601.
Bolbohamatum phallosum Krikken, 1980
Bolbohamatum phallosum Krikken, 1980: 21,
(description, keyed, distribution, illustration).
Figure 3: B. phallosum (scale: 5 mm), ZSI/CZRCA/16604, Pench Tiger Reserve, 2001.
Specimens examined: Madhya Pradesh:
ZSI/CZRC-A/16604; male (Length: 15.0 mm;
width: 9.0 mm); Turiya, Pench Tiger Reserve,
Seoni; K. Chandra, 23 June 2001; light trap;
ZSI/CZRC-A/16779; male (Length: 17.0 mm;
width: 11.0 mm); Kisli Rest House, Kanha
National Park, Mandla; M. Limje & party, 13
September 2003; light trap; ZSI/CZRCA/16780; male (Length: 16.0 mm; width: 10.0
mm); Kisli Rest House, Kanha National Park,
Mandla; M. Limje & party; 09 September
2003.
Diagnosis: (Fig, 3). Brown, shiny and pilosity
yellow brown. Clypeus with a pair of dentiform
tubercles. Pronotum with closely approximated
paramedian tubercles and lateral protrusion not
shifted to antero-lateral corner. Juxtasutural
punctures of elytra sub obsolete and discal
striae shallowly impressed and finely punctate.
Elytral inter-striae very slightly convex and
minutely and sparsely punctured.
External male genitalia: (Fig. 4) Dorsally,
parameres foliate and ventrally with a pair of
more or less glider-like paramerites. In lateral
Figure 4: Dorsal & ventral view of external male
genitalia of B. phallosum (scale: 2 mm), ZSI/CZRCA/16604.
Bolbohamatum marginale Krikken, 1980
Bolbohamatum marginale Krikken, 1980: 30,
(description, keyed, distribution, illustration).
Specimens
examined:
Chhattisgarh:
ZSI/CZRC-A/16599; male (Length: 16.0 mm;
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Remarks: The species can be easily
distinguished from its close relatives in having
pronotal lateral tubercles situated almost
marginally and very closely approximated
paramedian tubercle.
Geographical distribution: India: Chhattisgarh,
Madhya Pradesh, Tamil Nadu, Karnataka and
Uttarakhand. Elsewhere: West Pakistan.
New state and district record: Chhattisgarh
(Bilaspur) and Madhya Pradesh (Mandla).
width: 9.0 mm); Ataria Forest House,
Amarkantak Biosphere Reserve, Bilaspur; A.
Singh & party; 18 April 2004; light trap.
Madhya Pradesh: ZSI/CZRC-A/16600; male
(Length: 15.0 mm; width: 9.0 mm); Kisli,
Kanha National Park, Mandla; M. Limje &
party, 13 September 2003; light trap.
Diagnosis: (Fig. 5). Dorsal outline of left
mandible lobate. Clypeus with a pair of
dentiform tubercles each placed against lateral
margin. Pronotum with strongly approximated
paramedian tubercles and a pair of lateral
tubercles situated almost marginally. Median
longitudinal zone and lateral declivities of
pronotum densely and coarsely punctured while
impression between paramedian and lateral
tubercles virtually devoid of punctures and
opaque. Fore tibia with seven external
denticles.
Bolbohamatum laterale (Westwood, 1848)
Bolboceras lateralis Westwood, 1848: 385
(description, distribution).
Bolbohamatum laterale, Krikken, 1980: 33,
(description, keyed, distribution, illustration, comb.
nov.)
Specimens examined: Madhya Pradesh:
ZSI/CZRC-A16730; male (Length: 19.0 mm;
width: 12.0 mm); Sitapar, Singhori Wildlife
Sanctuary, Raisen; D. K. Harshay, 16
September 2009; day collection.
Diagnosis: (Fig. 7) Cephalic tubercles
dentiform and placed on lateral margins.
Paramedian tubercles of pronotum well
separated by more than inter ocular distance.
Lateral tubercles almost absent. Median cavity
of pronotal disc absent. Punctation on pronotal
disc abundant. Front tibia with eight denticles.
Figure 5: B. marginale (scale: 5 mm), ZSI/CZRCA/16599, Amarkantak Biosphere Reserve, 2004.
External male genitalia: (Fig. 6) Parameres
reduced and basal capsule enlarged. Basal
capsule of the phallus very robust in
comparison to B. laterale (Westwood, 1848)
and B. kuijteni Krikken, 1980.
Figure 7: B. laterale (scale: 5 mm), ZSI/CZRCA/16730, Singhori Wildlife Sanctuary, 2009.
External male genitalia: (Fig. 8) Apex of
parameres dorsally with short reflexed
paramerites. Basal capsule of phallus is robust
but not too much extant as of B. marginale
Krikken, 1980.
Geographical distribution:
India: Assam,
Maharashtra, Madhya Pradesh, Jammu &
Kashmir and Karnataka.
New state and district record: Madhya Pradesh
(Raisen).
Figure 6: Dorsal & ventral view of external male
genitalia of B. marginale (scale: 2 mm), ZSI/CZRCA/16599.
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U shaped, sparsely punctate impression.
Pronotum with anterior declivity impressed and
punctation generally sparse. Scutellum finely
punctate. Elytral striae deeply impressed well
defined and with large punctures. Elytral stria
two extending further caudad. Front tibia with
8-9 denticles.
Geographical distribution: India: Madhya
Pradesh, Maharashtra, Tamil Nadu and Uttar
Pradesh.
New district record: Madhya Pradesh (Raisen).
Remarks: The species shows variation in the
shape of clypeus, ornamentation of frons and
vertex. (Fig. 9 a, b).
Remarks: The species can be easily
distinguished from its close members, B.
marginale Krikken, 1980 and B. kuijteni
Krikken, 1980 having only one pair of lateral
pronotal protrusions and abundantly punctate
pronotum.
Figure 8: Dorsal & ventral view of external male
genitalia of B. laterale (scale: 2 mm), ZSI/CZRCA/16730.
Tribe: Bolbelasmini Nikolajev, 1996
Genus Bolbogonium Boucomont, 1911
Bolbogonium Boucomont, 1911: 340 (as subgenus
of Bolboceras Kirby; Type species: Bolboceras
triangulum Westwood, 1852: 342).
Bolbogonium Krikken, 1977: 79 (stat. nov.).
Generic diagnosis: Middle coxae widely
separated by an anterior lobe of pyriform
metasternal disc. First antennal club segment
on proximal side shiny and glabrous distinctly
separated from surrounding pubescent surface.
Seven elytral striae between elytral suture and
humeral umbone and all virtually reaching
base.
Bolbogonium insidiosum Krikken, 1977
Bolbogonium insidiosum Krikken, 1977:
(description, keyed, distribution, illustration).
95,
Specimens examined: Madhya Pradesh:
ZSI/CZRC-A/16605; male (Length: 9.0mm &
width: 5.5mm); Kartoli, Singhori Wildlife
Sanctuary, Raisen (23o 11.200’ N, 78o 12.085’
E); S. S. Talmale; 13 December 2010;
ZSI/CZRC-A/16778; male (Length: 8.0mm &
width: 5.0mm); Bhamori rest house, Singhori
Wildlife Sanctuary, Raisen; S. Sambath &
Party; 17 September 2011; day collection.
Diagnosis: (Fig. 9 a, b) Yellowish brown, shiny
and pilosity yellowish. Clypeal surface regulate
punctate. Frons with three small isolated
tubercles between eye-canthi. Vertex with large
Figure 9: Variation of B. insidiosum (scale: 2 mm),
ZSI/CZRC-A/16605 & 16778, Singhori Wildlife
Sanctuary, 2010 & 2011.
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Table 1: Checklist of genera Bolbohamatum and Bolbogonium from India
Distribution
Name of the species
India (states)
B. cyclops (Olivier, 1789)
B. calanus (Westwood, 1848)
Elsewhere
Genus Bolbohamatum Krikken, 1980
Bihar, Himachal Pradesh, Madhya Pradesh, New
Delhi, Uttarakhand, Uttar Pradesh and West Bengal
Assam, Bihar, Chhattisgarh, Karnataka, Madhya
Pradesh, Maharashtra, Tamil Nadu, West Bengal
and Uttarakhand
B. phallosum Krikken, 1980
Assam and Himachal Pradesh
B. robustum Krikken, 1980
B. laevicolle (Westwood, 1848)
B. pyramidifer Krikken, 1980
B. meridionale Krikken, 1980
B. marginale Krikken, 1980
Himalayan Region
Assam, Maharashtra, Orissa and Himalayan Region
Orissa
Puducherry
Chhattisgarh, Madhya Pradesh, Tamil Nadu,
Karnataka and Uttarakhand
B. kuijteni Krikken, 1980
Maharashtra
B. laterale (Westwood, 1848)
Maharashtra, Madhya Pradesh, Sikkim, Jammu &
Kashmir, Karnataka and West Bengal
Bangladesh
Pakistan
Genus Bolbogonium Boucomont, 1911
West Bengal
B. howdeni Krikken, 1977
Bihar
B. impressum (Wiedemann, 1823)
Himachal Pradesh and Uttarakhand
Karnataka, Madhya Pradesh, Maharashtra, Tamil
Nadu and Uttar Pradesh
B. insidiosum Krikken, 1977
B. davatchii (Baraud, 1973)
B. punctatissimum (Westwood, 1852)
B. scurra Krikken, 1977
B. triangulum (Westwood, 1852)
Bangladesh
and Java
Chhattisgarh, Maharashtra and Madhya Pradesh
B. pseudogrande Krikken, 1980
B. bicornutum Krikken, 1977
Nepal
Jammu & Kashmir, Himachal Pradesh, Uttar
Pradesh and Uttarakhand
Uttar Pradesh
Tamil Nadu
Andhra Pradesh, Bihar, Madhya Pradesh, Himachal
Pradesh, Uttar Pradesh, Uttarakhand and West
Bengal
Pakistan
Afghanistan
Bangladesh
Iran
(Nikolajev,
2008)
Myanmar,
Bangladesh
Pakistan
Acknowledgements
The authors are thankful to K. Venkataraman
(Director, ZSI) for providing necessary
facilities and encouragement.
Kral, D., I. Löbl and G. V. Nikolajev, 2006.
Bolboceratidae. In: Catalogue of the Palaearctic
Coleoptera - 3. Löbl, I. and A. Smetana (eds.).
Apollo Books, Stentrup, Denmark: 82-84.
Literature cited
Boucomont, A., 1911. Contribution à la
classification des Geotrupidae (Coleoptera),
Annales de la Société entomologique de France,
79: 335-350.
Krikken, J., 1977a. The genus Bolbelasmus
Boucomont in Asia, with notes on species
occurring in other regions (Coleoptera:
Geotrupidae). Zoologische Mededelingen, 51
(17): 278-292.
Carpaneto, G. M., R. Mignani and E. Piatella,
1993. A revision of the Afro-Indian genus
Bolboceratops
(Coleoptera,
Scarabaeoidea,
Geotrupidae), Journal of African Zoology, 107:
329-353.
Krikken, J., 1977b. Asian bolboceratine scarabs
of the genus Bolbogonium Boucomont
(Coleoptera-Geotrupidae),
Tijdschrift
voor
Entomologie, 120 (3): 77-108.
XX
75
TAPROBANICA VOL. 04: NO. 02
BOLBOCERATINE SCARABS OF CENTRAL INDIA
Krikken,
J.,
1978a.
The
Afro-Asian
Bolboceroides validus group (Coleoptera;
Geotrupidae), Zoologische Mededelingen, 52
(26): 301-311.
Nikolajev, G. V., 2003. The taxonomic
composition of the subfamily Bolboceratinae
from the Palaearctic faunistic region. Tethys
Entomological Research, 8: 187-206.
Krikken, J., 1978b: Taxonomic notes on
Bolboceras indicum Westwood and its relatives
(Coleoptera:
Geotrupidae).
Entomologische
Berichten Amsterdam, 38: 72-77.
Nikolajev, G. V., 2008. On the systematic
position of Iranian species Bolboceras davatchii
Baraud, 1973 (Coleoptera: Scarabaeoidea),
Caucasian Entomological Bulletin, 4 (2): 199201.
Krikken, J., 1979. The Genus Bolbocerosoma
Schaeffer in Asia (Coleoptera: Geotrupidae),
Zoologische Mededelingen, 54 (3): 35-51.
Ochi T. and M. Kawahara, 2002. Description of
the female of Bolbelasmus shibatai Masumoto,
1984. Kogane, 3: 31-33.
Krikken, J., 1980. Bolboceratine scarabs of the
Oriental genus Bolbohamatum nov. (Coleoptera,
Geotrupidae), Tijdscrift voor Entomologie, 123:
1-38.
Ochi T. and K. Masumoto, 2005. Systematic
Position of Bolbelasmus ishigakiensis Masumoto
Elytra, 33 (2): 244
Krikken, J., 1984. A generic reclassification of
the Afrotropical Bolboceratini (Coleoptera:
Geotrupidae). Zoologische Mededelingen, 58: 2345.
Ochi T., K. Masahiro and K. Masakazu, 2010. A
new species of Bolbochromus from the
Philippines (Coleoptera, Scarabaeidae). Kogane,
11: 97-99.
Li, C. L., C. C. Wang, K. Masumoto, T. Ochi and
P. S. Yang, 2008. Review of the Tribe
Bolboceratini s.l. from Taiwan (Coleoptera:
Scarabaeoidea: Geotrupidae) with a Key to the
Eurasian Genera. Annales of the Entomological
Society of America, 100 (3): 474-490.
Ochi T., M. Kon and M. Kawahara, 2011.
Four new taxa of Scarabaeoidea from Southeast
Asia. Special Publication of the Japanese Society
of Scarabaeoidology, Tokyo 1: 153-162.
Scholtz, C. H. and D. J Browne, 1996. Polyphyly
in the Geotrupidae (Coleoptera: Scarabaeoidea): a
case for a new family. Journal of Natural History,
30: 597-614.
Masumoto, K., 1984. New coprophagous
Lamellicornia from Japan and Formosa. The
Entomological Review of Japan, 39 (1): 73-83.
Smith, A. B. T., 2006. A Review of the FamilyGroup Names for the Superfamily Scarabaeoidea
(Coleoptera) with Corrections to Nomenclature
and a Current Classification, Coleopterists
Society. Monograph Number, 5: 144-204.
Nikolajev, G. V., 1970. Taxonomic status of the
groups belonging to the subfamily Geotrupinae.
Results of the Second Scientific Conference of
Young Specialists and Postgraduates, celebrated
100 years from the date of birth of V. I. Lenin and
50 years from the date of origin of Kazakhstan:
31-34.
Westwood, J. O., 1848. Description of some new
or imperfectly known species of Bolboceras,
Proceedings of the Linnaean Society London, 1:
384-387.
Nikolajev, G. V., 1979a. Eine neue BolbelasmusArt aus Asien (Coleoptera, Scarabaeidae,
Bolboceratinae.
Reichenbachia,
Staatliciies
Museum fur Tierkunde in Dresden, 17 Nr. 27,
225-227.
Westwood, J. O., 1852. Descriptions of some new
or imperfectly known species of Bolboceras
Kirby, Transactions of the Linnean Society
London, 21: 19-30.
Nikolajev, G. V., 1979b. Neue Gattungen und
Untergattungen der Blatthornkäfer (Coleoptera,
Scarabaeidae).
Reichenbachia,
Staatliciies
Museum fur Tierkunde in Dresden, 17 Nr. 23,
189-191.
70
76
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