Cour. Forsch.-Inst. Scnckcnberg I 245 I 281-313 I
7 Figs, 4 Pls
I FrankfUJ1 a. M., 30. 12.2003
-----------------------------------------------------------------
Conodonts across the Permian-Triassic boundary in the Southern Alps
With 7 figs, 4 pls
Maria Cristina PFRRI & Enzo FARABEGOLI
Abstract
The Tesero and Hulla sections (Dolomites, Italy) currently considered 01' pivotal importance l'or thc l'ermian-Triassic (P-T) boundary in the western Tethys were characterised hy a medium to high rate of shallow-marine sedimentation (5-]() cm k.y.-').
The thin, uppermost carbonate facies ofthe Permian Bellerophon Fmn (here named the "Bulla Member") is bounded regionally by unconformable surfaces. The upper unconforrnity coincides with the base of
the Werfen Fmn historically identificd with the P-T boundary. In the Bulla scction, it slightIy antedates (by
1.3 m) the P-T boundary-defined hy conodonts, specifically the firs! appearance of Hìndeodus parvus. The
Bulla Mbr yielded Hì. typicalis and Hi. praeparvus, extending their ranges above the P-T boundary into the
Wcrfcn Fmn The conodont associations l'rom the lower part ofthe Werfen Fmn consist mainly ofhindcodids
and isarciccllids showing graduai morphological change up-scction. The thickness and swcll1l1g ofthe basai
cavity anù of th.e attachment area of the denticlcs-giving the conodont a swollen appearance-has been
considercd laxonomically significant for ùefining rorms to be grouped in Isareìee/la. Four ncw specics
are describcd: Ifindeodus pìsai. Isareicella lobala. Is. m(lala and Is. peculiaris. Seven conodont biozoncs
have been discriminated, three are defined for the first time: Lower and Upper praeparvlIs. parvlIs. loba!a.
slaeseheì, ìsareica and aeqllabì/is zones. A possible lincagc between the genera Hindeodus and [mreicclla
is suggested. JJi. praeparvus has been considered the anccstor of Hi. parvus, the last represcntativc offlindcodus in the Southern Alps. Ili. praeparvIIs is also ancestral to Isarcicella prìsea tÌ'om which the subsequent
Isarcìcefla lineage evolved, namely Is. prìsca to Is. turgida, Is. lobata, lI'. s!aeschei and Is. isarciea.
The carbonate-terrigenous deposits around the P-T boundary oceur in thin (dm-m thick) depositional
cycles, recording superimposed high frequency, fourth- (l 00 k.y. period) and fifth- (\ 0-25 k.y.) order cycles driven by short-terrn sea-Ievel fluctuations. The main mass-extinction event occurred dunng the rapid
sea-Ievel fall followed by sea-Ievel rise at the contact of the Belferophon and Werfen Fmns. The lowstand
exhumed the top of the Bellerophon Fmn shallow-watcr carbonates resulting in moderate weathering and
erosion ofbcdrock. Many remanié Perrnian fossils occur in the basai Werfen Fmn transgrcssive trae!. The
high conodont diversity in the lower Tesero \llbr accords with rapid adaptative radiation following c1imatic
change. On the basis ofour resu\ts, we propose thallhe Bulla section---(;haracterised by relatively abundant
conodonts prccisely located in readily discriminahle high- depositional cycles-is the mosl rcliahle parastratotype scction for the P-T boundary in the western Palaeotethys.
K e y w o r d s: conodonts, Permian-Triassic boundary, Southern Alps, Italy.
Introduction
Since the nineteenth century, the Perrnian-Triassic boundary (P- T boundary) in the Southem Alps (Italy, Austria)
has been equated with the lithostratigraphic boundary
between the Bellerophol1 Fmn (below) and Werfen Fmn
(above) (RICHTHOFEN 1860, LEPSIUS 1878, MOJSISOVICS
1879, MERL;\ 1930, LEOI\\RI)I I935, PIA) 937, BOSELLINI
1964, ASSERFTO et al. 1973), two units cropping out extensively (fig. l) over an area of about 40,000 sq. km. VENZO
(1957) and subsequently PASINI (1985) found that the oldest
layers ofthe Werfen Fmn produced foraminifers, fusulinids
and calcareous algae ofPerrnian aspect. Attribution ofthe
lowermost Werfen Fmn to the Pcrrnian, supported by additional palaeontologic data (NERI & P\SI'i1 1985, BROGLIO
LORIGA et al. 1986, POSENATO 1988), became genera\1y accepted during the last decade when more than 20 detailed
contributions to litho- and hiostratigraphy, sedimentology
and depositional architecturc around the P T boundary
have been proposed for correlation between the Southem
Authors' addresses: Dipartimento di Scienze della Terra e Geologico-Ambientali, University of Bologna, Italy. Fax: 0039 051 2094522.
>perri@;geomi.n.unibo.it<, >fàra@geomin.unibo.it<
PERRI & FARAIlECiOI.l: Conodonts across the Permian - Triassic boundary in the Southem Alps
D
D
\dallH:lllì 11111 ｌ｜ｾｉ＠
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Fig. 1: Distribution ofuppermost Permian-Lower Triassic dcposits in the Southem Alps, and location of stratigraphic sections. AF:
Agordo-Frasscné, AS: Ansiei, BT: Butterlock, BU: Bulla, CH.: Col di Rioda, FA: Faedo, MC: Masi Saracini, MN: Monte Naro, MO:
Montagna, MR: Monte Rosà, MS: Maestrin, PR: Pressano, SP: San Pellegrino Pass, ST: Strigno, TS: Tesero. Black dots indicate
sections bearing conodonts.
Alps and well-studied coeval sections in China, Pakistan,
India and Iran.
Important stratigraphic questions remain unanswered.
Specific aims ofthis paper are: (i) precise location ofconodont taxa across the P-T boundary in two c\assic sections
(Bulla and Tesero sections) in the Dolomites, with particular regard to the species Hindeodus parvus, the entry
of which identifìes the base of the Triassic; (ii) revision
of conodont taxonomy aeross the P-T boundary in the
Southem Alps; (iii) reeognition of actual boundary SUffaces (conformable, uneonfonnable, or paraconformable
sensu DUNBAR & RODGERS 1957) and architecture of the
depositional sequences (cycles) near the P-T boundary - in
order to propose palaeoenvironmental-palaeogeographic
reconstructions consistent with field data.
Prob1cms regarding the P-T boundary in the
Southem Alps
Lithostratigraphy, depositional sequences and cyclcs
Nearly ali modem researchers identified an unconformity
(s3, in fig. 2) located a metre or less below the Bellerophon Fmn-Werfen Fmn boundary (BROGLIO LORICiA et al.
1986, 1988, FARABEGOLI et al. 1986, MAGARITZ et al. 1988,
WIGNALL & HALLAM 1992, MASSARI et al. 1994, WIGNALL
et al. 1996, CIRILLI et al. 1998, F ARAI3EGOLI & PERRI 1998,
NERI 1999, NERI & POSENATO I 999a), but most research282
ers deny existenee ofthe "classic" unconfonnity between
the Bellerophon and Werfen Fmns detected in the field by
BOSELLlNl (1964), ASSERETO et al., (1973), F ARABEGOLI &
VIEL (1982), FARABEGOLI & PERRI (1998) (rTl in fig. 2).
NERI & POSENATO (1999a), for instance, defined it as a
"paradigmatic" unconfonnity. Four international congresses held in the Southem Alps during the last twenty years
("Riecardo Assereto and Giulio Pisa Field Symposium on
Triassic Stratigraphy in Southem Alps". Bergamo lune
1979; "Pennian and Pem1ian- Triassic Boundary in the
South-Alpine Segment ofthe Western Tethys" from Italian
IGCP 203 Group 1986; European Conodont SymposiumECOS VII 1998; Intemational Field Conference on "The
Continental Permian of the Southem Alps and Sardinia
(ltaly), Regional Reports and GeneraI Corre\ations" 1999),
produced either an increase in stratigraphic information
and, paradoxieally, augmented confusion about the precise location and the nature ofthe P-T boundary. Current
opinions about the latter can be grouped as follows:
l. The contact between the Bellerophol1 and Werfen
Fmns is gradational and the P-T boundary is placed at a
thin oolitic unit (VENZO 1955, ULCIGRAI 1966), named the
Tesero Horizon by BOSELLIl\f (1964) and later the Tesero
Mbr by FARAREGOLI & VIEL (1982); it is loeated at or near
the very base of the Werfen Fmn (NERI & PASINI 1985,
BROGLIO LORIGA ct al. 1986, 1988, NOE' 1986, 1987,
SCHDNLAUB 1991, WIGNALL & HALLAM 1992, CIRILLI et al.
1998, NERI 1999).
Cour. Forsch.-Inst. Senckenberg, 245, 2003
w
Lombardy
Lake Como
Dolomites
Camonica Valley
o
Adige Valley
50km
Bellerophon Fm.:
Badiota faciescarbonates
ｾ＠
Werfen (Servino) F1/1.
III
ｾ＠
BasaI oolitic llOrizon
c::J sulphate evaporites
E
Cortina
Gardena fm.:
ｾ＠
conglomerate,
sanstone, pelite
. '....
Fiemma::.::.a facies-
..
Basement
Werfen Fm.: mari/le
ｾｧｩＺｮ･＠
Bellerophon Fm.:
marine limestone
polimictic-breccia
peritidal dolomite
f'-4--"L4..4
gypsmn-dolomite
clastic gypsum
ｾ＠
Gardena Fm.:
continental &
.,
... marine
, i ••'. conglomerates
and sandstones
(J)- nautiloid
ｆｾ］Ｚ｝＠
ｾ＠
ｾ＠
Athesian
volcanics
breccia
m
metamorphic
Farabegoli 2000
Fig. 2: Above: the «classic» sketch of the Middle Perrnian-Lower Triassic sequence and the P-T boundary in the Southem Alps.
A regional unconforrnity separates the Upper Perrnian Val Gardena and Bellerophon Fmns from similar westwards transgressive
oolitic beds (Tesero Horizon) ofthe Werfen Fmn (after ASSERETO et al. 1973).
Below: Upper Perrnian-Lower Triassic lithostratigraphy, facies architecture and depositional sequences in the eastem sector ofthe
Southem Alps. Val Gardena Fmn: l - Ora Member, 2 - Butterloch Mbr, 3 - Rio Bavaro Mbr; Bellerophon Fmn: 4 - Lavardet Mbr, 5
- Lozzo Mbr, 6 Rioda Mbr, 7 - Rio Barbide Mbr, 8 - Casera Razzo Mbr, 9 - Bulla Mbr; Werfen Fmn: lO - Tesero Mbr, Il - Mazzin
Mbr, 12 - Andraz Horizon, 13 - Siusi Mbr (after FARABEGOLI et al. 1986, slightly modified). Ali modem workers emphasized the
unconforrnable surface s3. Not many Authors pointed out the «classic» ravinement surface (rTl) connected with the last eustatic
event - predating the P-T boundary (defined by conodonts) by a few thousand years - located in the Werfen Fmn The upperrnost
trangressive sequence (tsP) corresponds to the Bulla Mbr, here defined.
283
PERRI & FARABEGou: Conodonts across the Permian - Triassic boundary in the Southem Alps
2. The contact between the Bellerophon and Werfen
Fmns is gradational, but the P-T boundary, defined by
conodonts, is located in the Mazzin Mbr of the Werfen
Fmn about 8 m abovc the top of Tesero Horizon and 15
m from the base ofthe Werfen Fmn (WIGNALL et al. 1996,
KOZUR 1998b).
3. A sharp, not clear, subaerial exposure surface separates the Bellerophon Fmn from the Werfen Fmn, whereas
the P-T boundary lies near the very bottom of the Tesero
Mbr (FARABEGOLI et al. 1986).
4. An unconformity (BOSEtLINI 1964, AssERETo et al.
1973, BRANDNER & MosTLER 1982, FARABEGOLI & VIEL
1982, MOSTLER 1982) divides the 8ellerophon and Werfen
Fmns. The P-T boundary corresponds to the base ofthe
Tesero Horizon (i. e. the base of Werfen Fmn), whereas
the hiatus equates with the uppermost part ofthe Permian
(ASSERITO et al. 1973).
5. An unconformity separates the Bellerophon and
Werfen Fmns whereas the P-T boundary, corresponding
to the FAO of Hi. parvlI.\. is placed 1.3 m above the base
of the Tesero Mbr (F-\.RAflFCiOLI & PERRI 1998); the corresponding hiatus has a very short duration, not apparent
from conodont data through time (this paper).
Other opinions, connecti ng palaeontological events
with depositional sequence architecture have been suggested producing a very complicated suite of proposals
(for a synthesis, see NERI 1999: tab. l).
Moreover, some subdivisions ofthe interval from the
upper Bellerophon Fmn to Mazzin Mbr - into sequences
or high frequency depositional cycles - has been proposed
(FARA BEGOLI et al. 1986, MAGARITZ et al. 1988, WIGNALL
& HALLAM 1992, MASSARI et al. 1994, WIGNALL et al.
1996, MASSARI & NERI 1997, CIRILLI et al. 1998, NERI
1999, NERI & POSENATO 1999a) (fig. 2). Because of lack
of reliab1e geochronological dating (according to recent
time-scales the duration ofthe whole Scythian may range
from 4 to about IO m.y., cf. 1[ARLAND et al. 1982, Om" &
LETOLLE 1982, HAQ et al. 1987, BRAcK et al. 1996, YI'\; et
al. 1997, GLlNISTER et al. 1999), identification of Upper
Permian-Lower Triassic sequences tends to he specula-
ｾｔ＠ ,
,
,
MO
FA
,
tive with numerous interpretations differing in terms of
hierarchy or cause (F ARABEGOLI et al. 1986, NOE' 1986,
1987, MAGARITZ et al. 1988, POSENATO 1988, KOZUR 1998a,
1998b, NERI & POSENA TO 1999a).
[n the following, unless otherwise specified, we will
generally use sequence terminology without reference to
hierarchical order (timing, intensity, geographical extent),
or possible cause (GOLDHAM\1ER et al. 1990).
Biostratigraphy and chronostratigraphy
In the Southern Alps the uppermost Permian-Lowcr Triassic shallow water sequence, lacking ammono id faunas.
has been investigated for brachiopods, calcareous algae,
foraminifers, bivalves, gastropods, pollen, fungi (PASINI
1985, NERI & PASINI 1985, BROGLIO LORlGA et al. 1986,
1988, POSENATO 1988, WTGNALL et al. 1996, CIRILLI et al.
1998. KOZUR 1998a, b, NERI & POSENATO 1999a) and,
mainly, by means of conodonts for biostratigraphic and
chronostratigraphic alignments.
HUCKRIEDE (1958) first deseribed conodonts from the
Southem Alps defining the new Triassic species Spathognathodus isarcicus. STAESCHE (1964) carri cd out important
research on Scythian conodonts from the South Tyrol; this
became the milestone for Lower Triassic conodont workers. SWEET (in ASSERETO et al. 1973) first extracted a few
conodonts from the uppermost part ofthe Bellerophon Fmn
and some from the overlying lower portion ofthe Werfen
Fmn KOZUR & MOSTLER (in MOSTLER 1982) proposed the
first local conodont zonation, discriminating eight biozones in the Werfen Fmn SCHONLAUB (1991) published a
very rich conodont fauna from the lower part ofthe Werfen
Fmn collected from the Gartnerkofel-l core in the Carnic
Alps. F ARABEGOLI et al. (1986), PERRI (1986, 1991, 1998),
PERRI & ANDRAGHETTI (1987), F ARABIGOLI & PERRI (1998),
PERRI & FARABEGOLI (1998) and NICOR.\ & PERRI (1999)
contributed new data on the conodont fauna - mainly in
terms of multielement taxonomy - of thc uppermost Bellerophon Fmn and the entire Werfen Fmn with particular
,
PR
,
MC
MR
,
ST
ｾＮＭ
I l mudstone,
L
I siJtstone
ｾ＠
ｾ＠
oolitic storm
laycrs and shoals
ｾ＠
oolitic hars
[ •ｾ＠ • Jsand
shallu\\ manne
amI hrecclU
,
Farahegoh 2{)()(J
MN
ti
+
't:
Tesero è\th.
'"
ellerophon Fm +
. al Gardena l'm.
_ _ tidal cvaporites
and slltstones
c===J
Fig. 3: Lithostratigraphic and facies scheme ofthe lower Werfen Fmn in the Westem Oolomites. Section locations in Fig. 1.
284
"
Cour. Forsch.-Inst. Senckenberg, 245, 2003
regard to the P-T boundary. The chronostratigraphic
significance of conodonts ahout the P-T boundary of
this region has been discussed by SCHONLAUB (1991),
KOZUR (1996, 1998a, b), ORCHARD (1996), WIGNALL et al.
(1996), FARABEGOLI & PERRI (1998), ORCHARD & KRYSTYN
(1998), YIN & TONG (1998) and NICORA & PERRI (1999).
The very different opinions presented derive either from
the Iithostratigraphic nomenclature used (i.e. Tesero vs.
Mazzin Mbr) or from criteri a used to define the P-T
boundary, but really these derived from the scarcity of
palaeontological data.
Bulla Section
The Bulla section (BU) is well exposed along thc road
joining Ortisei to Bulla (fig. 1), about 0.5 km NW ofBulla
(Pufels). It consists of marine environmcnts extending
from upper Permian to upper Scythian, overlain with an
unconformable contact by the Richthofen Conglomerates
(upper Anisian) (MOSTLER 1982, PERRI 1991, FARABEGOLl
& PERRI 1998).
The complete lithic column can be summarized as
follows:
- Bellerophon Formation;
- Unconformity;
Werfen Fmn, consisting or in asccnding order:
Tesero Member: oolitic grainstone-packstone a few m
thick, interfingering with the over1ying Mazzin Mbr;
Mazzin Member: marine clays alternating with mudstones, passing upwards to alternating siltstones, thin
mudstone and subordinate wackestone layers and subordinate thin calcarenitic storm layers. It passes gradually
upwards to:
Andraz Mem ber: siltstones and very fine reddish marly
sandstones alternating with yellowish supratidal (evaporitic) dolomitic laycrs.
The sequence ncar thc P T boundary will be described
in detail below and compared with the corresponding interval in the Tesero section.
Lithology and sedimentology near the P-T boundary
in the Bulla Section (fig. 4)
Bellerophon Formation
Dolomite facies: yellow-grey dolomite and sandy
dolomite beds, 10-20 cm thick, containing ostracods,
bivalves, foraminifers (Geinit::ina sp.) and ?radiolarians
deposited in an open shallow lagoon above wave base;
dolomitization was prohahly post-depositional. This facies
corresponds to the basaI part of the uppermost transgressi ve sequence in F ARABEGOLl et al. (1986), to Cycle VI p.p.
in MASSARI et al. (1994), and to thc Casera Razzo Mbr (8)
in fig. 2 (below).
- Unconformity.
- Bulla Member (1.30 m thick): predominantly dark
fossiliferous packstones alternating with bioturbated
wackestones and thin bedded dark clayey siltstones. Ihe
lowermost calcareous beds consist of mm-size lithoclasts
(pebbles) mixed with an undoubtedly reworked microfauna offoraminifers, fusulinids, ostracods, crinoids, and
red and green algae. The siltstones contain a few crinoid
fragments. The middle and upper layers contain macrofossils - undoubtedly not reworked - with geopetal fillings
(e.g. Bellerophon sp.) as well as other fossils of doubtful
origino Hummocky and wave ripples indicate a shallow (6
m or less) shore environment where high-energy events
(storm layers) alternated with low energy periods. A regional unCOnfOlll1ity separates this unit from the overlying
Tesero Mbr ofthe Werfen Fmn
Thc Bulla Mbr is a transgressive-rcgrcssive faciescouple extending from Carnia to the Dolomites (BROGLIO
LORIGA et al. 1986, FARABIGOLl et al. 1986, NERI et al.
1986, POSENATO 1988, CASSINIS et al. 1993, WIGNALL &
HALLAM 1996, KOZUR 1998b, NERI 1999). Ihis interval has
been referred to by various informai names, e.g.: Uppermost Bellerophon Fmn, Member 2, interval IV (BROGLIO
LORIGA et al. 1986); upper unit of the Bellerophon Fmn
(NERI et al. 1986); Event l or Comelicania beds (POSE'JA 10
1988); uppe11110st beds of the Black Limestone Unit of
the Be/lerophon Fmn (NoE' 1987, MAGARITZ et al. 1988);
Comelicana-Nankinella Assemblage (BROGLIO LORIGA et
al. 1988); uppellllost part of parasequence I at the base
of third order sequence 6 (MASSARI et al. 1994, NERI &
POSENA TO 1999); diverse benthos packstone of the Uppermost Bellerophon Fmn (HALLAM & WIGNALL 1999);
Comelicania beds ("horizon") and Unit 3 (NERI 1999).
To stabilize lithostratigraphic nomenclature, FARABEGOLI
(herein) proposes to name it the "Bulla Member of the
Bellerophon Fmn"; the Bulla scction is proposed as the
reference section.
- Unconformity, readily identified at microscopic scale
(F ARA.BEGOLI & PERRI 1998: 295, fig. 4.3.4.). Ihis surface
corresponds to the c1assic erosional surface first depicted
by ROSELLTNI et al. (1973: fig. 6).
Werfen F01111ation
Tesero Member: lower lithosome (2.30 m, samples
BU9-BUI2C). The lowermost "fossiliferous" dark packstone-grainstone beds (20 cm) contain many small, subrounded, "terrigenous" lithoc1asts mixed with a reworked
fauna of foraminifers, fusulinids, ostracods, crinoids, red
and green algae, and with rare oolites. Ihey pass upwards
to five gray oolitic grainstones and packestones, containing
both reworked and unreworked fossils (e.g. Bellerophon cf
vaceki) alternating with c1ayey siltstone and wackestonc.
Except l'or the lower 20 cm, the lower Tesero Mbr produccd
a well preserved conodont fauna (samplcs BU9-BUI2C
scc F ARABEGOLl & PERRI 1998 and this paper). Grainstones
and packstones feature hummocky bedding, passing upwards to wave ripples. They can be interpreted as st01111
events within a transgressive shallow (3-6 m) shoreface.
The depositional environment was very similar to that of
the underlying Bulla Member of the Bellerophon Fmn
285
PERRI & FARABEGOU: Conodonts across the Permian - Triassic boundary in the Southem Alps
Bulla Section
Tesero Section
samples (8l')
samples
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Fig. 4: Upper Permian-Lower Triassic deposits in the Bulla and Tesero sections.
286
｟ｾ］＠
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t
TES71 ' - '
TES69'-'
ｾ＠
Ｎｾ＠
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TES62'-'
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current direction Q9 ·.>radiolaria
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11
:---:=::::::
= --
/
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.
--,..I__---,
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(6'!--"'1-l1
l'acies
/
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2
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w
ｾＧＭｴｌＬ＠
Marly
Dolomite
ｾＭＺｳ［］＠
TSIO'-'
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TS9 .-,
13
Jì
-=r
---=
-'li..
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(T)
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r- 2.
)
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)*
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-13
•
ev
z . ｾ＠ ｾ＠
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•
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-12e'-'
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-
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I
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L)
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f
Ostracod
root traces
bioturbation
____ stilolith
COUf. Forsch.-Inst. Senckenberg, 245, 2003
except for the great reduetion in autochthonous skeletal
biomass in the oolitie grainstones.
The sueeeeding faeies consists of l m thick alternating
mudstone and pelletoid wackestone with oolitie fossiliferous paekstone containing rare reworked fossils. This tàcies
produeed a rieh conodont fauna. It is regarded as calm
marine in-offshore (below wave base) interrupted by rare
distaI storm layers. It corresponds to Units 4 and the 5 a,
band c portions ofthe Tesero section (NERI 1999). It may
be a high-stand trae t of the sequence, modified by two
short-term Milankovieh eyeles.
This transgressive traet ofthe lowermost Tesero Mbr
extended over a wide area from Carnia to Lombardy,
eovering both the Upper ealeareous "members" of the
Bellerophon Fmn and the uppermost Val Gardena Sandstones (BOSELLlNI et al. 1973, BR()C;1I0 LORIGA et al. 1986,
FARABEGOLl et al. 1986, NERI et al. 1986, NERI 1999).
Only in the Trento area (centraI western Dolomitcs) did
eontinental eonditions persist over some tectonically rejuvenated topographic highs (F ARABEGOLl & VIEL 1982,
this paper: fig. 3).
with pelletoid waekestone, referred to as probably algal
stromatolites (WIGNALL et al. 1996), deposited on a calm
marine bottom below wave base. Only the lowermost part
of this member produced conodonts. The middle of this
facies may be the transgressive, HST traet ofthe sequence,
modified by short term Milankovieh cycles.
Tesero Member: upper lithosome (1.35 m thick). It
consists ofa positive (thinning fining upwards) sequence,
composed offive oolitic packstone and grainstones layers,
altemating with mudstone-wackestones. This progradational-retrogradational facies may be interpretcd either
in terms of inereasing coastal dynamics, eausing parti al
destruetion of oolitic bars and shoals developed on the
Trento area struetural highs, and contemporaneous offshore deposition or, more probably, as horizontal shifting of eoastal fàeies caused by high frequeney sea level
changes (i.e. regressive-transgressive system tracts related
to Milankovich cycles).
Mazzin Member: upper lithosome. Thin bedded,
coupled pelletoid and micritic mudstone, with lamination of probably algal stromatolitic origin, alternating
with siltstones and subordinate packstones, deposited on
a marine botto m Iying below wave base. If we eonsider
this composite faeies as due to high frequency sea levcl
changes, the siltstone intervals eould eorrespond to more
humid intervals related to a late transgressive traet.
The lower lithosome of the Tesero Mbr passes gradual1y
upwards to:
Mazzin Member: lower lithosome (2.9 m thick, samples BU13A-BUI3B): thin bcds ofmudstone alternating
ti-
ｾ＠
§
...
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..D
E
U
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r:/:J
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'"
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o
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BU26
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BU23
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28.47
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-
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2.82
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-8
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ｾ＠
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:1:
ｾ＠
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ｾ＠ ｾＺＱ＠
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...;
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L
U
Perrl lODO
Fig. 5: Cono don t range-chart o[ the Bulla (BU) section across the P-T boundary.
From left to right: chronostratigraphy: systems; lithostratigraphy: formation and members; biostratigraphy: inferred biozones,
samples numbers, sample distances in m [rom the base ofthe Werfen Fmn Ha.: Hadrodontina, Hi.: Hindeodus, Is.: Isarcicella, St.:
Stepanovites.
287
PERRI & FARABEGOLl: Conodonts across the Perrnian - Triassic boundary in thc Southem Alps
Ｍｾ
Conodonts from the Bulla Section
The conodont associations found in the lower part of the
Bulla section, even though presented in PERRI (1991) and
F ARAIlEGOLI & PERRI (1998), are here tabulated in fig. 5 in
conncction with reporting new data obtained from taxonomic revision; only productive samplcs, the subject of
this revision, are reported.
Samplcs from the Bellerophon Fmn and the first 0.20
m ofthe Werfen rmn produced no conodonts; neither did
the sequence ofabout 25 m in the Mazzin Mbr, from 3.50
m (above BUI3B) up to 28.50 m (BU23).At 1.30 m, in the
lower Tesero Mbr, Hi. parvus (KOZUR & PJATAKOVA 1976)
makes its appearance in sample BU12B in association
with Hi. praeparVlIS, Hi. pisai. Is. prisca and Is. turgida
(KOZUR et al. 1975). Thc entry ofthis species defines the
base of the Triassic.
Tesero Section
The Tesero section (TS) crops out in the neighbouring of
the Tesero village, about 35 km S ofBulla, and is one of
the most studied P-T boundary sections (LEONARDI 1929,
1935, BosELLINI 1964, ASSERETO et al. 1973, PASINI 1985,
NERI & PASINI 1985, BROGLIO LORIGA et al. 1986, 1988,
BRANDNER et al. 1986, Noi 1987. MAGARITZ et al. 1988,
PERRI 1991, WIGN\LL & H AI.LAM 1996, WIGNALL et al.
1996, KOZlJR 1998b). Many authors regard this section
as thc western Palaeotethys referencc section for the P-T
boundary in global correlations (WIGNALI. et al. 1993, NERI
& POSFNA TO 1999b).
Lithology and sedimentology in the Tesero Seetion
(fig. 4)
- Bellerophon Formation
F oss i l i fero u s Do 10m i te fac i es: fine-grained dolomitic mudstone and wakestone with algae, foraminitèrs,
molluscs and ostracods. This facies corresponds to the
lowcr part of the Dolomite Unit of MAGARITZ et al. (1988)
and to Unit I ofNERI (\999).
Marly Dolomite facies_(morethan 2.5 m thick): Iightgrey marly dolostone characterized by frequent subaerial
exposurc surfaces aIternating with thin pelitic beds. This
facies corresponds to the upper part ofthe Dolomite Unit
ofMAGARITZ et al. (1988), to the top ofIIIrd-order depositiona! sequence n. 5 of MASSARI et al. (1994), and to Unit
2 ofNERI (1999).
- Unconformity.
Bulla Member (about 0.50 m thick): blackish pelites
grading upwards into marls and mixed terrigenous-fossiliferous calcarenites. The coarse terrigenous components are represented mostly by reworked foraminifers
and algae of Permian-type. Recently, PIRINI RADRlZZANI
(1999) considered this to be an autochthonous fauna. This
288
unit passes upwards into the Tesero Mbr. Hummocky and
wave ripples point to a shallow shore environmcnt. No
conodonts (P ERRI 1991, NlcoRA & PERRI 1999), macrofossi!s (POSENATO 1999) or po1\en (CIRILLI 1999) have been
reported from this unit in the Tesero section. It corresponds
to Unit 2 p.p. ofMAGARITZ et al. (1988), to Unit 3 ofNERI
(1999) and the lowermost pari of the Comelicania beds
in nearby areas (NERI & POSENATO 1999 b).
- Unconformity.
- Werfen Forn1ation
Tesero Member: (7.60 m thick: TES62 TS10A). Thc
lowermost dark calcareous beds (Unit 4a of NERI 1999)
contain numcrous small, angular to sub-roundcd, terrigenous Iithoclasts mixed with a clearly reworkcd Permiantypc fauna of foraminifers, fusulinids, ostracods, crinoids,
and red and grccn algae. The autochthonous components
consist of rare oolites and the Ombonia-Orthotetina assemblage (POSENATO 1999). The succeeding 0.90 m of
thick oo!itic grainstones (Unit 4b ofNERI 1999) contains
Ombonia, Bellerophon vaceki. conodonts and sparse, very
probably reworked, Permian-type algae (Gymnocodium).
They pass upwards to five grey oolitic grainstones and
packstones (Unit 5a ofNERI 1999) containing either reworked (fusulinids, algae) and unreworked fossils (Rellerophon cl vaceki). Grainstones and packstones feature
hummocky bedding, passing upwards to wavc ripples;
they can be interpreted as storrn events within a shallow
shoreface.
The succeeding tracts (Unit 5b of NERI 1999 - with
stromatolite carbonate mounds alternating with thin storm
layers and the Crurithyris fauna - and Unit 5c) could correspond to the transitional beds from lower Tesero Mbr
to lower Mazzin Mbr in the Bulla section. Ihe topmost
oolitic grainstone (Unit 6 ofNERI 1999) corresponds to the
upper Tesero Mbr at Bulla. The Tesero Mbr has produced
a conodont fauna (PERRI 1991, NICORA & PERRI 1999).
M azzin M em ber: This (TS 16-TS28) consists ofbioturbatcd to laminatcd marly lime mudstone alternating with
thin fossiliferous biocaIcarenite storm layers, rcadi Iy COfrelated with the upper Mazzin Mbr in the Bulla section.
Thc depositional environment was offshore or shelfbelow
the wave base.
Conodonts from the Tesero Section
The conodont fauna from the Tesero section was previously studied by SWEET (in ASSERETO et al. 1973), PERRI
(1991) and subsequently by NICORA & PERRI (1999).
The lower portion ofthe section, consisting ofthc uppermost part of the Bellerophon Fmn and the first 35 cm
of the Werfèn Fmn, failed to produce conodonts. as did
the interval ofabout 8.70 m from 2.40 (abovc TSI0A) to
Il m (IS 16). The lowermost conodont fauna occurs in the
Tesero Mbr in sample TES62 (0.35--0.40 m: fig. 4). The
conodont ranges are reported in fig. 6. Noteworthy is the
finding of Is. changxingensis. According to WANG (1995)
COUI. Forsch.-Insl. Scnckenberg, 245, 2003
--------------------------------------------':.:i-<
<\l
TS28
28.00
S
TS26
26.00
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"ti
e -S!""
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(\)
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s::
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et:
r
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Q
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:::'"
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o
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ｾ＠
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15.30
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:::
-
TS10A
2.20
TSlO
2.00
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1.70
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1.30
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Z
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et:
[.LJ
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e
(\)
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r
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:::5
-
r-
TES69
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....;
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":
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!
!:
Sl
Sl
e
14.30
11.00
ｾ＠
;.
ｾ＠
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:lj
ｾ＠ <;.;;
ｾ＠
Ｎｾ＠
ｾ＠
ｾ＠ .:;
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c:;
--'Ｎｾ＠ -
<;.;;
I....
'=!
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'=!
ｾ＠
Q
ｾ＠
Ｎｾ＠
24.00
22.50
TS 16
r
24.40
TS23
TS19
ｾ＠
'=!
-Q
Q
-ｾ＠
'ｾ＠
.;::::
.::-j
'=!
ｾ＠
ｾ＠
ｾ＠
ｾ＠
<;.;;
ｾ＠
'::J
"l
ｾ＠
>-...;
;:s
.::-j
.::-j
ｾ＠
c:;
<;.;;
.::2
....
J::l..
,.;
>-...;
TS62
0.30
Pem 2000
Fig. 6: Conodont range-chart ofthe Tesero (TS and TES) section.
From left to right: chronostratigraphy: systems; Iithostratigraphy: lànnation and members; biostratigraphy: infcrrcd biozones, samples
numbers, sample distances in m from the base ofthe Werfen Fmn Ha.: Hadrodul1tina, Hi.: Hindeodus, Is.: Isarcicella.
and KOZUR (1996), this species has a very short range in
China overlapping with the entry of Hi. parvus.
Conodont fauna
For the present contribution the entire conodont fauna of
the uppermost beds of the Bellerophon Fmn and of the
lower Werfen Fmn (Tesero and Mazzin Mbrs) collccted
from sections along the Southem Alps (Carnic Alps to
Dolomites) has been revised (FARABEGOLI et al. 1986, PERRI
& ANDRA(;] IETII 1987, PERRI 1991 ) (fig. I). Conodonts were
found in the Col di Rioda and Ansiei sections cio se to
Auronzo (Belluno, Eastem Dolomites) in the uppermost
beds ofthe Bellerophon Fmn (PERRI & AN DRAGHETII 1987).
The more interesting and abundant data come from the
Western Dolomites, and particularly from the previously
described Bulla section (Gardena Valley, Bolzano) (PFRRI
1991, FARABFGOLI & PERRI 1998, PERRI & FARAIlHiOLl
1998) and thc Tesero section (Fiemme Valley, Trento)
(PERRI 1991, NICORA & PERRI 1999). The high deversity
of forms found in the studied materia! has stimu!ated the
present research directed towards c1arifYing morphological
trends among Hindeodus and Isarcicella.
Befare describing the sequence of conodont associations, it is appropriate to describe some ofthe broad evo-
lutionary trends noted in the hindeodids compared with
isarcicellids. The changes readily recognizable are:
• reduction in conodont length compared with height;
• the wide and laterally compressed dentic1es ofthe biade
tending to be reduced to higher, narrow and rounded
dentic1es;
• the cusp being nearly the same height as the denticles
gradually becomes more than twice the height of the
denticles;
• the basai cavity tending to inerease, beeoming eupshaped;
• the width and asymmetry of the basai cavity tending
to increasc until starting to become cup-shaped with
lateral 10bes;
• owing to reduction in element length, the pit shifts from
a proximal position to a centrai situation.
Ali speeies recognized i n the praeparvus-prisca-isarcica evolutionary trend display cither forms with the
posterior tip ending abruptly or spccimens with denticles
reaching the posterior end - as already mentioned by
PERRI (1991, 1998). These morphological differences al10w discrimination oftwo morphotypes in Hi. praeparvus
as well as the subspecies parvus and erectus within Hi.
parvus (KOZUR 1995, 1996, ORCHARD & KRYSTYN 1998).
A salient characteristic of this evolutionary trend is that
the thickness ofthe basai cavity, mainly in the inner more
289
PlKKI
& FARABEGOU: Conodonts across thc Pcrmian
Triassic boundary in the Southem Alps
expanded side, and ofthe attachment area ofthe denticles,
gives the conodont a swollen appearance. This peculiarity
was firstly noted by KOZUR (1975) when he describcd Is.
turgida and by KOZLR (1995) when hc dcfìncd 1.1'. prisca
by discriminating it from Hi. praeparvus solely on the
basis ofthis thickening. Though KOIl;R (1995, 1996) suggested that the thickening and swcll ing of thc cup could be
eeo\ogically control\ed, conodont associations from the U.
praeparvlIs Zone up to the isarcica Zone display incrcasing
devclopmcnt ofthis peculiarity. Swollen elcments are more
abundant in the younger beds, contrasting with decrease in
number of un-swollen elements of llindeodlls. The highcst
percentage of swollen and thickened forms oeeurs in thc
lobata, staeschei and isarcica zoncs where reprcsentatives
of isareieella are predominant. Un-swollen elements are
already very rare in levels where Is. lobata is present.
Graduai increase in asymmetry, width, swelling and
thickening of the basai cavity in the studied material
cou Id be evidence of transition between Ilindeodus and
Isarcieella. The medium-high sedimentation rate of the
sequenee about the P-T boundary in the Southern Alps
has al\owed reeognition ofthe above transition. The same
conodont fauna, found scattered through 20-40 metres in
the Alps, occurs in only a fcw mctrcs of section in China,
Kashmir and Pakistan.
Conodont Zonation
For the Southern Alps, the conodont zonation ofthe uppennost Pennian and lowennost Triassic has becn based
on hindeodids and isarcicellids. The uppcrmost beds of
the Bellerophon Fmn and the lower portion ofthe Werfen
Fmn, comprising the Tesero Mbr and part of the Mazzin
Mbr, have bcen subdivided into scvcn biozoncs by means
of conodonts (fig. 7).
l. Lower praeparvus Zonc
Lower limit: defined by the fìrst entry of Hindeodus
praeparvus KOZUR 1996.
Upper limit: defincd by thc first appcaranee of Isarcicella
prisca KOZUR 1995.
Associated fauna: Hi. typicalis.
In the studied successions, the lower Iimit of the biozone has not been discriminated beeausc Hì. praeparvus
oecurs in the first productive samples of the uppernlOst
levels ofthe Bellerophon Fmn in the western Carnic Alps
and extends into the Werfen Fmn Thc biozonc is part of
the flpicalis Zone of PERRI (\99\).
2. Upper praeparvus Zone
Lower limit: defined by the first appearance of lsarcicella
prisca KOZLR 1995.
Upper Iimit: defined by the first entry of Hindeodlls parvus
(KOZUR & PJATAKOVA 1976).
Associated fauna: Hi. tvpicalis is present. Ili. pìsai, isarcìcella changxingensis, and Is. pecliliaris first occur within
the biozone. The biozone, hcrc proposed, eorresponds to
part of the t.picalis Zone of PIRRI (\99\ ) and to the lati290
Ｍｾ
dentatus Zonc of FARA BEGOLI & PERRI (1998).
3. parvlIs Zone
\.ower I imit: defined by the first entry of Hindeodus parvlls
(KOZLR & PJATAKOVA 1976).
Lpper limit: defined by the first appcarancc of Isorcicella
lobota n. sp.
Associated fauna: Hi. typìcolis is present.is. turgida seems
to enter from the base, associated with Hi. pan'us. Hi. pisai
ends at the base ofthe biozone whereas Hi. praeparvus and
is. prisca end in the lower part of the biozonc. Thc biozone
corresponds to part of thc typiealis Zone or PERR! (1991)
and of the parvus Zone of F ARABEGOLI & PERRI (1998).
4. lobata Zone
Lower lim it: defined by the first entry ofisarcicella [ohata
n. sp.
Upper limit: defined by the first appearance of lsarcicella
staesehei DAI & ZHANG 1989.
Associated fauna: Hì. typicalis, Hi. parvlIs and Is. turgIda arc prcscnt. is. inflata is prescnt from the base or the
biozone.
This biozone, herein proposcd, corresponds to the
lower portion of thc isarciea Zonc of PERRI ( \991 ), to the
upper part ofthe parvus Zone (sample BU23) of FARABECiOLl & PERRI (1998) and to part of the staeschei Zone
(sample TS23) ofNlcoRA & PERRI (1999).
5. staesehei Zone
Lower limit: dcfined by the first cntry of Isarcicella staeschei 0.\1 & ZHA'JG 1989.
Upper limit: defined by thc first appearance of isarcicella
isarcica (HLCKRIEDE 1958).
Associatcd fauna: Hi. parvus cnds within thc biozone. Aiso
present are is. turgida, is. in/lata and Is. [obala.
Samples BU25A-BL26A in the Bulla section and
samples TS24-TS25 in thc Tescro section have been assigned to the biozone. Il con'esponds to part or the isarcica
Zone OfPERRI (1991), the lower part ofthe isarcica Zone
of FARABEGOLI & PERRI (1998) and to the upper part ofthe
staeschei Zone ofNICORA & PIRRI (1999).
6. isarcica Zone
Lower limit: defined by the first appearance of Isareicella
isareiea (HUCKRIEDE 1958).
Upper limit: defined by the disappearance of isareicella isareiea (HUCKR1EDE 1958) and by the presence of
Hadrodontina aequabilis STAES('llE 1964, marker species
ofthe following local biozonc rccognised in the Southem
Alps.
Associatcd fauna: Is. injla/a, [l'. [o ba/a, Is. s/aeschei and Is.
isarcica end their ranges near or at the top of the biozone.
Ha. aequahilis occurs in the bim-:one.
The biozone corresponds to the upper part ofthe isareiea Zone (samples BU27-BU27 A and samplc TS26) of
PERRI (1991) and ofFARAFlrt,OL! & PI:RRI (1998) and to the
isarcica Zone (sample TS26) ofNlCORA & PERRI (1999).
Cour. Forsch.-Inst. Scnckenhcrg, 245, 2003
ｾ
ｾ＠
.!:;.
Ｍ
,<t.....
ｾ＠
ｾ＠
.§
s:-
'-'
-.]1
1 -
......
!\ .....' '
ｾｓｬ＠
ＢＬｾＧ＠
ｾ＠ ..• '
Ｎｾ＠ｾ＠
ｾ＠
-
ｾ＠
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i,
Il
•
Ｍｾ＠
ｾＮＧ＠ Ｍ｜ｾｖ＠
..
ｾ＠
' '§
'"
ｾ＠
Ｎｾ＠
ＭｾＬ＠
c
ｾ＠
ｾ＠
- - - - - - - - - - - - - - - s;: - - - - - . ,
. .i
t..
"
..
1 :r:
e
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:r:"'"
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JIssvnLl
Fig. 7: Phylogeny of Hindcodus and Isarcicella in the uppermost Permian and lowermost Triassic, infcrrcd from Southem Alps
data.
From Jeft lo right: chronostratigraphy: systems, lithostratigraphy: formations and members, biostratigraphy: biozones, conodont
ranges and phylogeny. ｐｾ＠ TB: ｐ･ｲｭｩ｡ｮｾ＠
Triassic Boundary defined by conodonts.
291
PERRI
& FARABEGOU: Conodonts across the Permian - Triassic boundary in the Southem Alps
Phylogeny or Hindeodus in the uppermost Permian
and Iowermost Triassic
Hindeodus is a biostratigraphically important genus especially for research on sequences spanning the Permian-Triassic boundary. In the SouthemAlps, representatives ofthe
genus Hindeodus are more frequent in the lower part of the
studied succession in the Upper praeparvus and parvus
zones - with the species Hi. typicalis, pisai, praeparvus
and parvus (fig. 7).
The new species Hindeodus pisai could have been derived from Hi. minutus because of its numerous narrow
denticles, the presence of some small denticles anterior to
the cusp, and be cause of the profile of the denticulation
seen in lateral view. Hi. praeparvus could also have been
derived from Hi. typicalis by reduction in number ofthe
denticles and increase in width ofthe last denticles ofthe
biade. Frequent elements of Hi. praeparvus and rare Hi.
typicalis occur in the first productive levels, their ranges
overlapping with the first occurrences of Hi. parvus. Several transitional forms between Hi. praeparvus and Hi.
parvus have been identified among the numerous specimens assigned to Hi. praeparvus in bed BUI2A, below
the first appearance of Hi. parvus. These elements, even
though already showing reduction in conodont length compared with height and displaying increase in cusp height,
stili have the last three denticles ofthe bIade wide (pl. 2,
figs 22-24). In the Bulla section, transition between these
two species is clearly visible. It is therefore possible to accept that Hi. parvus was deri ved from Hi. praeparvus. HL
parvus ranges in the studied material up to the slaeschei
Zone; it is the last species ofthe genus.
Phylogeny of Isarcicella in the uppermost Permian
and lowermost Triassic
The genus Isarcicella, typical of the base of the Lower
Triassic, could have originated from one of the latest
species of Hindeodus (fig. 7). In this paper, Isarcicella
has been enlarged to include not only forms with a wide
inflated cusp bearing lateral denticles on one or both sides
of it, but also elements without lateral denticles, though
always with a wide asymmetrical swollen cusp. Forms not
necessarily contemporaneous (and mainly stratigraphically
lower) displaying swelling and thickening of the biade,
of the attachmen t area of the denticles, and of the basai
cavity - though stili having a slightly asymmetrical reduced basai cavity - do occur. Such elements, assigned
to various species such as peculiaris, changxingensis, prisca and turgida, have been included in Isarcicella. The
stratigraphically lowest representatives showing these
peculiarities have been included in Isarcicella? prisca by
KOZUR (1995). In the Southem Alps numerous specimens
of this species occur in association with Hi. praeparvus
- which has the same denticulation - and from which it is
differentiated only by the swelling\thickening ofthe basaI
cavity. Because Is. prisca occurs in levels stratigraphi292
cally younger than those with the first occurrences of Hi.
praeparvus, the latter could be the ancestor of Isarcicella
prisca. Hi. praeparvus occurs in the uppermost beds of
the Bellerophon Fmn but without Is. prisca. Among the
first forms showing swelling of the basai cavity is Is.
changxingensis. WANG (1995) described and figured the
species, assigning it to Hindeodus. According to PERRI,
who assigned ali forms showing swellings to Isarcicella,
the species changxingensis should also be placed in this
group. The only specimen found in the SouthemAlps was
obtained from the Tesero section (sample TES71) in the
praeparvus Zone at 1.30 m from the base of the Werfen
Fmn It has a widely swollen basai cavity like the forms
figured by WANG (1995). On the basis of denticulation, Is.
changxingensis - with portion of the biade fused - could
be derived from the stratigraphically lower Permian form
Hi. julfensis. Other swollen forms, always from the base
ofthe Werfen Formation, are included into the new species
Is. peculiaris; the latter could have been derived from Is.
prisca by extreme enlargement of the denticles and rounding oftheir apices. Is. turgida was derived from Is. prisca
within the parvus Zone by reduction in conodont length
compared with height and increase in height of cusp and
denticles - such as occurred in Hi. parvus, the latter was
derived from Hi. praeparvus. Is. turgida is the ancestor
of both Is. lobata and Is. inflata. Increase in asymmetry
of the basaI cavity - tending to enlarge and swell up to
become a cup with a laterallobe - transforms Is. turgida
into Is. lobata. Extreme enlargement ofboth sides ofthe
cup is characteristic of the forms assigned to the new
species Is. infiala. This too was derived from Is. turgida
within the base of the lo baIa Zone. Appearance of one or
a series of nodes or dentici es characterises forms assigned
to Is. slaeschei. There are numerous elements very cio se
to Is. lobala and some to Is. infiala, differentiated only
by bearing a little node or denticle on one side ofthe cup;
these demonstrate that Is. staeschei was derived from Is.
lobala and, it in tum, probably developed into Is. infiata. In
the analysed material very high intraspecific variability is
displayed by Is. slaeschei. Forms bearing one or a series of
denticles on one side ofthe cup co-occur from the first bed
having this form, whereas in Spiti (India) and Kashmir the
first elements of Is. slaeschei have a single lateral denticle.
Is. slaeschei enters at levels stratigraphically younger than
the first occurrence of Is. lobala. Is. isarcica was derived
from Is. slaeschei by appearance of one or more nodes or
denticles on both sides of the cupo
In summary the studied material displays evolutionary
transition between Hindeodus and Isarcicella. Is. prisca
derived from HL praeparvus could be the ancestor of Is.
lurgida, and it the ancestor of Is. lobala and I. infiala,
whereas Is. staeschei could have derived from Is. lobala
and gave rise to Is. isarcica.
Cour. Forsch.-Inst. Senckenberg, 245, 2003
--------------------------Permian-Triassic boundary
The proposal by YIN (1993) to utili se the first appearance
of Hindeodus parvIIs to define the lower limit of the Triassic System has now been ratified by the Intemational
Stratigraphic Commission on Stratigraphy with the GlobaI Stratotype Section and Point for the Permian-Triassic
boundary on section D at Meishan, China (YTN et al. 200 l).
The choice was suggcsted bccause Hi. parvus has been
shown to be present in alI investigated sections crossing
the P-T boundary. lt is present in both shallow water and
pelagic biofacies. The stratigraphically lowesl occurrence
of Hi. parvus in the SouthemAlps was first pointed out by
MOSTLER (1982) in the Bulla section at about 2.5 m from
the base ofthe Werfen Fmn but, unfortunately, no plate figuring the conodont was published in the guide-book on the
excursion to Siusi in Siidtirol. According to FARABEGOLI &
PERRI (1998), the first appearance of Ri. parvus in the Bulla
section occurs in thc lowcr Tesero Mbr at 1.30 m from the
base ofthe Werfen Fmn In the Tesero section, the base of
the parvus Zone was indicatcd by WIGNALL et al. (1996)
to be in the middle part ofthc Mazzin Mbr. Since then the
base ofthe biozone in the Southem Alps has been reported
in that position, i.c. the base ofthe Triassic System is taken
to occur in the middle part ofthe Mazzin Mbr. NICORA &
PERRI (1999), also, found the first representatives of Hi.
parvus in the Tesero section in sample TS 16 in the Mazzin
Mbr about II m from the base ofthe Werfen Fmn At 1.30
m from the base (sample TES71) Is. changxingensis occurs
and at 2-2.20 m (samples TS lO and TSI0A) some broken
specimens have been found seeming to display transitional
features between Hi. praeparvus and Hi. parvus. NICORA
& PERRI (1999) think that in the Tesero section Hi. parvus
should be present at a lower leve\.
In the Bulla section (FARAllHiOLl & PERRI 1998), the
first metre ofthe Tesero Mbr has numerous well preserved
specimens of Hi. praeparvus passing, in the upper part
of this basai sequence - just before the first entry of Hi.
parvus - to associations with typical specimens of HL
praeparvus co-occurring with obviously transitional forms
between Ili. praeparvus and Hi. parvus. The lower limit of
the Triassic can be identified by the first appearance of Hi.
parvus in a continuous sequence with a clear\y recognisable transition between Hi. praeparvus and Hi. parvus. Thc
biostratigraphic boundary - defined by means ofcondonts
in the lower Tesero Member - is thus closer to the lithostratigraphic boundary. The latter obviously indicates an
important event associated with a discontinuity recognisable in most sections across thc P- T boundary.
Conclusions
Litho- and biostratigraphic ana\ysis ofthe uppermost part
of the Bellerophon Fmn (Permian) in the Southem Alps
has highlighted two unconformable surfaces that bound 1
m or less of shallow-marine carbonate facies (here namcd
the Bulla Mbr). The upper unconformity, for a long time
ignored after its first discovery (BOSELLINI 1964, ASSERETO
et al., 1973), corresponds to the basaI contactofthe Werfen
Fmn Tt slightly antedates the P-T boundary defìned by
means ofconodonts. One research goal was to defìne in the
Southcm Alps thc evolutionary appearance of l/indeodus
parvus, the index species for identitying the basc of thc
Triassic. fii. parvus fìrst appears at 1.30 m (BU 12B) above
the base ofthe Werfen Fmn The fìnding of Hi. parvus in
the Bulla section allows identifìcation of the base of thc
Triassic System in the lower Tesero Mbr (FARABEGOLl &
PERRI 1998), lowering the position of the P-T boundary
previously believed to be in the middle of the Mazzin
Member (WIGNALL et al. 1996).
A review of taxa across the P-T boundary, mainly
focused on hindeodids and isarcicellids, has been carri ed
out. The swelling\thickening of parts of conodonts has
been considered taxonomically signifìcant for defining
forms grouped in 1sarcicella. Such forms become dominant at stratigraphically highest levels. Ali Pa scaphate
elements presenting no swellings were let1 in the gcnus
Hindeadus.
The Bulla Mbr (Upper Permian) yielded Hi. typicalis
and Hi. praeparvus onlyin the Col di Rioda andAnsiei sections close toAuronzo (Belluno, Eastem Dolomites). These
forms persisted upwards above the P-T boundary. Four
new species from the lower Wcrfen Fmn are described:
Hi. pisai, Is. peculiaris, Is. fabata and ls. iriflata.
The biostratigraphical data allow subdivision of the
interval from the top ofthe Bellerophon Fmn to the Mazzin Mbr ofthe Werfen Fmn into sevcn conodont biozones:
Lower and Upper praeparvus, parvus, fabata, staeschei.
isarcica and aequabilis zones. The fìrst entry of 1.1'. prisca
defìnes the base ofthe Upper praeparvus Zone hercin proposed. In this biozone, Is. changxingensis, 1.1'. peculiaris
and Hi. pisai occur. High conodont diversity in the lower
Tesero Mbr can be explained by rapid adaptative radiation following climatic change. Entry ofthe new specics
II'. fabata a1\ows defìnition of a new biozone, thc loha/a
Zone, located between the parvus and staeschei zones. It
corresponds to the upper portion ofthe parvus Zone or to
part ofthe staeschei Zone ofprevious papers.
A possible Iineage between the genus Hindeodus and
Isarcicella has been suggested. GraduaI morphologic
changes can be discriminated in thc conodont associations, such as reduction in conodont length compared with
height, incrcase in width and asymmetry ofthe basaI cavity, and the lattcr becoming a cup with lateral lobcs bearing nodes. Thc laterally compressed wide denticles tend
to be reduccd to higher, narrow, rounded denticles. The
cusp doubles its hcight. The posterior end can be either
adenticulate, cnding abruptly, or denticulate with !ittlc
denticles reaching the posterior tipo
GraduaI increase in asymmetry, width, and swelling
and thickening of the basaI cavity in the studied materiai could be evidence of transition between the genera
Hindeodus and Isarcicella. The medium to high sedimen293
PERRI & FARABEGOU: Conodonts across the Permian - Triassic boundary in the Southem Alps
tation rate of the sequence across the P-T boundary in
the Southern Alps enabled recognition of the transition.
The same conodont fauna occurs through a few metres
in China, Kashmir and Pakistan, but in the Alps ranges
through 20-40 metres.
Hi. praeparvus is the ancestor of Hi. parvus, the last
representative of Hindeodus in the Southern Alps, and of
Is. prisca from which the enti re Isarcicella lineage could
have originated. The evolutionary trend appears to have
been Is. prisca, Is. turgida, Is. lobala, Is. slaeschei and
Is. isarcica.
The succession around the P-T boundary consists of
depositional sequences that can be related to short-term
(25-100 k.y.) sea-Ievel fluctuations, conceivably isochronous at global scale. The main mass-extinction event
occurred in conjunction with coupled, rapid, sea-Ievel fall
followed by sea-Ievel rise at the contact ofthe Bellerophon
and Werfen Fmns. The lowstand exhumed the Bellerophon shallow-water carbonates and resulted in moderate
weathering and erosion ofbedrock. Many remanié fossils
(fusulinids, calcareous algae) were incorporated into the
transgressive basai tract.
The Bulla section, characterized by relatively abundant
conodont discoveries related to well identified depositional
sequences, seems to be the most reliable parastratotypical
section for the P-T boundary in the western Palaeotethys.
Similar, partly condensed sequences, detectable both in
shelf (China) and nearshore (Pakistan) sequences, are
probably isochronous at global scale.
Remarks: The conodonts here assigned to the scaphate
Pa element of the multielement apparatus of Hindeodus
present a symmetrical or slightly asymmetrical lachrymiform basai cavity not bordered by flange-like brims.
Biade, attachment area of the denticles and basai cavity
show no swelling and thickening. The posterior tip can
end either abruptly or with denticles decreasing in height
and reaching the tipo
Range: Lower praeparvus-staeschei zones: uppermost
Permian-Lower Triassic, lower Scythian.
Hindeodus parvus (KOZUR & PJATAKOVA 1976)
(pl. 2, figs 4-12)
1975
1976
d-e.
1981
1987
1988
1995
1996
1996
Systematic Palaeontology
1998
Genus Hindeodus REXROAD & FURNISH 1964
Ty P e s p e c i es: Trichonodella imperfecla REXROAD
1957
(= Spathognathodus cristulus YOUNGQUlST & MILLER
1949)
294
Hindeodus parvus (KOZUR & PJATAKOVA) - MATSUDA: 91, pl.
5, fig. 2.
Anchignathodusparvus KOZUR & PJATAKOVA- YANG et al. (eds):
pl. 36, fig. 2.
Anchignathodus typicalis SWEFT - BUDUROV & GUPTA: fig. 8.
b,f
1995
1996
1996
Descriptions of species are based only on Pa elements.
Preservation ofthe Pa elements of Hindeodus REXROAD &
FURNISH 1964 and Isarcicella KOZUR 1975 is go od whereas
the ramiforms ofboth genera are often broken, not allowing complete reconstruction of the apparatuses. The Pa
elements have to be observed in various views. Ideally,
Lower Triassic conodont workers should publish plates
with numerous elements figured in upper, lower and lateral views. A lateral view is inadequate because in several
species the denticulation is so similar that it is necessary
to observe ali views and especially the upper view. Synonymies are limited to key citations and are not intended to
be comprehensive. Figured specimens are housed in the
collections ofthe Museo Capellini ofthe Dipartimento di
Scienze della Terra e Geologico-Ambientali, University
ofBologna.
Anchignathodus parvus KOZUR & PJAT AKOVA - KozuR, MOSTLER
& RAIlIMI-YAZD: 4, pl. I, figs 13-15, pl. 7, figs 7, 9.
Anchignathodus parvus KOZUR & PJATAKOVA: 123, figs 1a-b,
1998
Hindeodus parvus morphotype l (KOZUR & PJATAKOVA) - KoZUR: 69, pl. 2, fig. 6.
Hindeodus parvus morphotype 2 (KOZUR & PJATAKOVA) - KoZUR: 69, pl. 2, fig. 9.
Hindeodus parvus (KOZUR & PJATAKOVA) - llN et al.: pl. 5. 3,
figs 6, 8.
Hindeodus parvus (KOZUR & PJATAKOVA) - YIN & ZHANG: pl.
2. 8, fig. l.
Hindeodus parvus erectus KOZUR: 97, pl. 2, figs 6, 8.
Hindeodus parvus parvus (KOZUR & PJAT AKOV A) - KOZUR: 96,
pl. 2, figs 7.
Hindeodus parvus morphotype I (KOZUR & PJATAKOVA)
- FARAREGOLI & PERRI: pl. 4.3.1, figs IO-Il.
Hindeodus parvus (KOZUR & PJATAKOVA) - KOZUR: pl. I, figs
4,7,12.
1998
1999
1999
Hindeodus parvus (KOZUR & PJATAKOVA) - ORCHARD & KRYSTYN: 351, pl. 6, figs 9, 16, 17,20.
Hindeodus parvus erectus KOZUR - NICORA & PERRI: pl. 3, figs
8-9, Il.
Hindeodus parvus parvus (KOZUR & PJATAKOVA) - NlcoRA &
PERRI: pl. 3, tigs 7, 12.
Description: The Pa element is small with a big and
rather slender cusp conspicuously higher than the succeeding denticles.
Remarks: The posterior end can be either adenticulate
abruptly ending, or with denticles decreasing in height
towards the tipo Based on this peculiarity, KOZUR (1995)
defined two morphotypes, respectively morphotype l for
forms with denticles small, nearly of the same size and
with abrupt posterior end, and morphotype 2, to which
the holotype belongs, with denticulate end. Subsequent1y
KOZUR (1996) elevated the two morphotypes to subspecies,
Hi. parvus erectus and Hi. parvus parvus respectively. In
the studied material both subspecies morphologies have
been discriminated, having about the same stratigraphic
range. Differing morphologies of the posterior end are
Com. Forsch.-Inst. Senckenberg, 245, 2003
recognized in several species ofHindeodus andisarcicella;
such differences are here construed as characteristics of
morphotypes and not used as a basis for discriminating
subspecies.
In the present paper, Hi. parvus is restricted to forms
lacking swelling ofthe biade and Iacking thickening and
swelling of the attachment area of thc denticles and of
the basaI cavity, peculiarities not prcsent in the holotype.
Specimens assigned to Hi. parvus, likc that figured by
MATSUDA (1981: pl. 5, fig. 3), showing an asymmetrical
swollen basaI cavity, arc hcrc included in the Isarcicella
lobata. Several transitional forms between Hi. praeparvus
andHi. parvus have been collected. They display reduction
oflength and increase in height ofthe cusp, but still have
wide dentic1es, especially the last three.
Occurrence: Werfen Fmn, Tesero and Mazzin Mbrs,
Bulla section, samples BUI2B-BUI3B. Tesero section,
samples TS 16-TS25.
Range: parvus-staeschei zones: Lower Triassic, lower
Scythian.
ｈｩｮ､･ｯｵＬｾ＠
1998
pisai n. sp.
(pl.l,figs 1-12)
Hindeodlls n. sp. A FARABEUOL! & PERRI: pl. 4.3.1, fig. 9.
Derivatio nominis: In honour of our late friend and
colleague GIULIO PISA of the University of Bologna, specialist on Triassic.
H o lotype: IC 1749-989176, figured in pl. l, figs 7-9, and
in FARABEGOLI & PERRI 1998: pl. 4.3.1, fig. 9.
P aratypes: IC 1771-989178, IC 1773-989177 figured in
pl. l, figs 1-3, 10-12.
the dentic1es dccrease in height very gently up to 2/3 of
the unit but abruptly in the posterior third.
Description: Fragile elements tending to be elongate,
bearing 13-15 narrow, high, needle-shaped denticles,
mainly rounded in section, fused at their bases and free at
the apices. The 9-10 denticles behind the cusp are about
the same height, decrcasing very gently along two-thirds
ofthe unit, whereas thc last 5-7 denticles constituting the
posterior third decrease abruptly. Cusp narrow and higher
than denticlcs. One to three little dentic1es can he prcscnt
anterior to the cusp. Denticles and cusp show longitudinal
striae. Thc lacrhymiform basai cavity, extending along the
posteri or two-thirds of the unit, is narrow and shows the
pit in its anterior third. The uni t lacks swelling.
Remarks: The only specimen with features recalling Hindeodus pisai is one assigned to HL minutus by
SCHONLAUB (1991 : pI. I, fig. lO). The element is elongate
with about 15 denti cl es which, even though in a large part
broken, seem to have the same lateral profile as the new
species. lt was found about 40 cm from the base of the
Wcrfcn Fmn in the Reppwand section on the north slope
01' Gartnerkofel in the Carnic Alps (Austria). The outline
of the numerous little denti cles seen in lateral view and
the presence or denticles anterior to the cusp suggest Ili.
min1ltus could be ancestral to Hi. pisai.
The new spccies occurs in the Bulla section in samplcs
BU12Aand BUI2B at 1.20-1.40 m from the base ofthe
Werfen Fmn In the Strigno section it is present in sample
ST1B about 2.30 m from the base ofthe Tesero Mbr.
Occurrence: Werfen Fmn, Tesero Mbr. Bulla section, samples BUI2A-BU12B; Strigno section, sample
STIB.
Rangc: Uppcr praeparvus-parvus zones: uppermost
Permian Lower Triassic, lowermost Scythian.
L o c u s ty P i c u s : Bulla section at about 0.5 km along the
road NW of Bulla village, Province of Bolzano, Western
Dolomites.
S tratum typicum: Levei ofsample BUl2B ofthe Bulla
section, 1.30 m from the base of the Werfen Fmn in the
Tesero Mbr.
Repository: Museum G. Capellini, Dipartimento di
Scienze della Terra e Geologico-Ambientali, University
ofBologna.
Material: 16 specimens.
Hindeodus praeparvus KOZUR 1996
(pU, figs 13-16; pl. 2, figs 22-36)
1981
1987
1991
1991
1991
1991
1995
1995
Diagnosis: The Pa element is characterised by a biade
consisting of 13-15 very narrow necdle-shaped dentici es.
Thc narrow cusp is higher than the succeeding dentic1es.
One to three little denticles can be present anterior to the
cusp. Basai cavity is narrow, tear-shaped. In lateral view
1996
1996
1998
Hindedlls minulus (EUISON) - MATSUOA: 78, pl. I, fig. 9.
Hindeodus typicalis (SWEET) - PERRI & ANIJRAGHElTl: 308, pl.
32, figs 3-4.
Hindeodus /ypicalis (SWEET) - PERRI: 40, pl. 3, figs 2, 5, 6.
Hindeodus cf. ialidcntalus (KozuR, MOSTI.ER & RAHIMI-YAZD)
- SCH6NLAUB: pl. I, fig. 9.
Hindeodus parvus (KOZUR & PJATAKOVA) - SCIl6NL\1In: pl. l,
fìgs 8, 18.
Hindeodlls cf typicalis (SWEET) - SCH6NLAUB: pl. I, fìgs 15-17.
Hindmdus lalidenlatus (KOZUR, MOSTLER & RAilIMI- Y\!f))
- KO/.I!1<: 67, l'l. I, figs 5-8, pl. 2, figs 2, 3.
l1indl'Odlls lalidenlalus (KOZUR, MOSTLER & RAHIMI- Y,\/f))
- WAVi: pl. 2, figs 4-5.
Hindeodus lalidentalus praeparvus KOZUR: 93, pl. 2, figs 1-4.
Hindeodus lalidenlalus (KOZUR, MOSTLER & RAHIMI-YAZO)
- WIGNALL, KOZUR & HALLAM: fig. la.
Hindeodus latidenlalus morpholype 1 (KOZUR, MOSTLER &
RAHIMI-YAZD) - FARABEGOLl & PERRI: pl. 4.3.1, figs 1-3.
295
PERRI & FARABEGOU: CO!1odonts across the Permian - Triassic houndary in the Southem Alps
ｾＭ
1998
1998
1998
1999
1999
Hindeodl/s lalidenlalus morphotype 2 (KOZUR, M()SI LER &
RAIIIMI-YAZD) - F\RAIlI(;OIl & PERRI: pl. 4.3.1, figs 4-5.
Hindl'Odus laridenlall/s l'raeparvus KOZUR - KOZUR: pl. I, tigs
5-6, 9, II.
Hindeoduspraeparvus KOZUR - ORCHARD & KRYQYN: 352, pl.
6, figs 22-23.
Hindeodlls praeparvus morphotype I KO/<J!< - NlCORA & PERRI:
pl. 3, figs 1,3, 5-6.
Hindeodus praeparvus morphotype 2 KOZUR N'CORA & PERRI:
pl. 3, fig. 2.
Descri pti O n: The Pa element is characterised by a small
to moderately large cusp, with several denticles of the
posterior biade distinctly larger than those ofthe anterior
bIade.
Remarks: According to KOZUR (1996) Hi. latidentatus
may be subdivided into two subspecics Hi. latidentatus
latidentatus and Hi. latidentatus praeparvus. The forms
with U-shaped widely-spaced denticles in the posteri or
part ofthe bIade were assigned to Hi. latidentatus latidenfatus whereas ali elements with V-shaped, spaced, broad
denticles were assigned to Hi. latidentatus praeparvus.
ORCHARD (in ORCHARD & KRYSTYN 1998) elevated the two
subspecies to the rank of species. On the basis ofthis revision, specimens includcd hy PERRI (in FARABEGOLI & PERRI
1998) in Hi. latidentatlls are assigned to Hi. praeparvus.
In Hi. praeparvlIs two morphotypes were distinguished
on the basis of an abrupt (morphotype l) or denticulate
(morphotypc 2) posterior end. Both have been tòund in
the examined materia!. ORCHARD (in ORCHARD & KRYSTYN
1998) suggested to separate morphotypc l as a discrete
specics, and described and figured a specimen (p. 354, pl.
6, fig. 24) not close to morphotype l of Hi. praeparvus
but that could represent of a new species.
Occurrence: Uppermost part ofthe Bellerophon Frnn
Col di Rioda section, sample CR3; Ansiei section, sample
AS2. Werfen Fmn, Tesero Mbr. Bulla section, samples
BU9-BUI2B; Tesero seetion, samples TES62-TS9A.
Werfen Fmn, Mazzin Mbr. Tesero seetion, sample TS 16;
Maestrin section, sample MS3.
Range: praeparvlls-parvus zones: uppermost PermianLower Triassic, lowermost Scythian.
Hindeodus typicalis (SWEET 1970)
1970a
1970b
1977
1987
1987
1988
1993
296
Anchignathodlls Il'pim/is SWEET: p.7, pl. l, figs 13,22.
Anchignathodus IJjJica!is SWEET - SWEET: 222, pl. 1. tigs 13,
20.
Hindcodus tvpicalis (S\\FI T) - SWEET: p.223, Hindeodlls-pl. 2.
figs 1-6.
Hind<!Odl/s ftpicu!is (S\\I.I.1) - PERRI & ANDRAGHETTI: 308, pl.
32, figs 1-2.
Anchlgnathodl/s mil1l/lus (ELUSON) - YANG et al. (eds): pl. 36,
figs 1,6.
Anchignathodlls typicalis SWEET - BUDUROV & ｇｕｐｔｾＺ＠
fig. 8.
c, i. k.
Hindeodlls (vpicalis (SWEET) - TIAN: pl. I, figs 13-21.
1995
199(,
199X
Hindeodus typicalis (SWEET) - KO/L'R: 65, pl. l, figs 1,3,4.
Hindeodlls tvpicalis (S\\FFT) - YI'\ ci al.: pl. 2. 8. figs 5, 13.
Hindeodus typicalis (SWEEr) - OkCII\RD & KRYSTYN: 357, pl.
6, figs 18-19,25-26.
Description: The Pa element is characterised by a low
cusp and denticles ofthe bIade decreasing slight1y in height
towards the postcrior end. Denticles present are free only
at their apices. The pit is located in the first third of the
lachrymiform narro w basaI cavity.
R e m a r k s: In the studied material, Hindeodlls typicalis
is rare even though present from the upperrnost levels of
the Bellerophon Frnn up to the lower part of the Werfen
Fmn--the Tesero and Mazzin Mbrs. Forrns c10ser to the
holotype have been found aeross thc lithostratigraphic
boundary ofthe two tòrmations.
Occurrence - Uppermost levels of the Bellerophon Fmn
Col di Rioda section, sample CR3; Casera Federata section, sample CFI5. Werfen Fmn, Tesero Mbr. Bulla section, sample BUIO; Strigno section, sarnple STIB. Mazzin ｾ｢ｲＮ＠
Casera Federata section, sample CFI6; Tesero
section, sample TS 19.
R a n g e: Lower praeparvlIs-parvlIs zones: uppermost
Permian-Lower Triassic, lower Scythian.
Genus Isarcicella KOZUR 1975
Type species: Spathognathodus isarcicus HUCKRIEDE
1958
Revised diagnosis: Multielernent apparatus with
scaphate element in Pa position characterized by swelling/
thickening of the basai cavity bordered by flange-like
brims. Nodes or denticles may or may not be present on
one or both sides ofthe cup.
Remarks: According to PERRI'S opinion the genus lsarcicella defined by KOZUR (1975) for Pa clernents showing
a very wide and inftated cup bearing lateral nodes or denticles has to be extended also to elements with weakly to
highly asymmetrical and swollen cup but without lateral
nodes. Forms occurring stratigraphically lower displaying weak asymmetry and swelling of the basaI cavity
and without lateral nodes have been included. The species changxingensis, peculiaris. prisca and tllrgida will,
accordingly, be assigned to Isarcicella. Species such as
/ohata and inflata showing an asymmetrical. swollen and
thickened cup and appearing very c10se to 1.1'. staeschei and
Is. isarcica, but without lateral nodes or denticles, are here
placed in lsarcicella. Taxonomically signifìcant characters
offònns referred to are: 1) asymmetrical weakly to highly
inflated cup bordered by flange-like brirns; 2) pit tending
to be centraI; 3) cusp wide, higher than the other dentici es
- the latter tend to become rounded in section and discrete
Cour. Forsch.-Inst. Senckenberg, 245, 2003
in species stratigraphically higher; 4) posterior end mainly
abrupt but sometimes denticulate.
ｬｾ｡ｲ｣ｩ･＠
should have a multielement apparatus cIose
to that of Hindeodus even though different. In samples
where many Pa elements of Isarcicella are present there are
few ramiforms which could be referred to the Isarcicella
apparatus. Unfortunately these are broken and numerically
much fewer compared with Pa elements so it was impossible to be sure ofthe constitution ofthe apparatus.
Range: Upperpraeparvus Zone: uppermost Permian.
Isarcicella inflata n. sp.
(pl. 4, figs 10-11, 15-19)
1981
1991
1991
Range: Upper praeparvus-isarcica zones: uppermost
Permian-Lower Triassic, lower Scythian.
Isarcicella changxingensis (WANG 1995)
(pl. I, figs 17-19)
1995
1995
1999
Hindeodus changxingensis WANG: 149, pl. 2, figs 14-18
Hindeodusjulfensis (SWEET) - WANG: pl. 23 fig. 1
Hindeodus changxingensis WANG - NICORA & PERRI: pl. 3, fig. 4.
Description: The Paelementis characterised for having
part ofthe biade straight, adenticuiate, smooth.
Remarks: In the hoIotype described by WANG (1995: pl.
2, figs 16-18) only two denticIes are present posterior to
the high and wide cusp in front ofthe smooth biade. WANG
figured (1995: pl. 3, fig. 1) another specimen assigned
to Hindeodus julfensis. According to PERRI that form is
closer to the holotype of changxingensis - from which is
differentiated solely by having several denticIes behind
the cusp - rather than to Hindeodus julfensis. This last
species is characterised by a peculiar "hump" lacking in
WANG'S specimen; it therefore could be assigned to Is.
changxingensis.
In the Alps, NICORA & PERRI (1999: pl. 3, fig. 4) found
a specimen at 1.30 m from the base ofthe Werfen Fmn in
the Tesero section; it was referred to Hi. changxingensis.
Behind the unfortunateIy broken cusp it dispIays five
narrow denticIes followed by portio n of an adenticulate,
smooth biade. The posterior third ofthe unit shows three
little denticIes decreasing in height. The smooth part of
the biade is shorter than in the hoIotype of the species
- indicated by numerous free denticIes behind the cusp.
The specimen found in the Alps has the same peculiarity - visible in WANG'S previously cited specimen (pl. 3,
fig. l) - showing 3-4 denticI es folIowed by a piece of
non-denticulate bIade. AlI the forms, either from China
or Tta\y, have an asymmetrical very wide and inflated cupo
In this paper, PERRI considers this feature as a taxonomic
character for distinguishing Hindeodus from Isarcicella;
representatives of the species changxingensis should be
compared with primitive forms showing swelling and
thickening of the basaI cavity and which have been included herein in Isarcicella.
Occurrence: Werfen Fmn, Tesero Mbr. Tesero section,
sample TES71.
Hindeodus parvus (KOZUR & PJATAKOVA) - MATSUDA: 91, pl.
5, fig. I.
Isarcicella isarcica morphotype l (HUCKRIEDE) - PERRI: 42, pl.
4, fig. IO, pl. 5, fig. 5.
Isarcicella isarcica morphotype 2 (HUCKRIEDE) - PERRI: 42, pl.
5, fig. 4.
Derivatio nominis: Pa element characterised by an
enormously swollen and lobed cupo
HoIotype: ICI465-8923, figured in pl. 4, figs 15-17 and
in PERRI (1991) in pl. 4, figs IO a-b.
Locus typicus: Bulla section about 0.5 km along the
road NW ofBulla village, Province ofBolzano, Western
Dolomites.
Stratum typicum: Level ofsample BU27 ofthe Bulla
section; Mazzin Mbr 45 m from the base of the Werfen
Fmn
Repository: Museum G. Capellini, Dipartimento di
Scienze della Terra e Geologico-Ambientali, University
ofBologna.
Material: 12 specimens.
Diagnosis: The Pa element is characterised by a very
wide, asymmetrical, lobed, swollen and thickened cup
lacking Iaterai nodes or denticIes. BIade composed of
6-8 denticIes. Cusp high.
Description: The cup is extremely expanded and can
be lobed on both sides and bordered by flange-like brims.
The 6-8 denticles of the biade can be either distinct or
quite completeIy fused with apices free. In the holotype
the apices of the denticIes are partly abraded. The cusp is
straight or slightly bent posteriorwards and higher than
the denticIes.
Remarks: Representatives of the species, often broken
with regard to the Iarge dimension of the cup, occur in
association with Isarcicella lobata, Is. staeschei and Is.
isarcica. Forms with fused denticIes seem to be more
frequent in stratigraphically higher levels. Is. inflata is
differentiated from Is. lobata by the very much wider and
inflated cup, and by the outer side ofthe cup being always
great1y expanded. The species couId be derived from Is.
lobata by extreme dilation ofthe cup and be the ancestor of
Is. staeschei. MATSUDA (1981 : pl. 5, fig. l) figured a specimen assigned to Hindeodus parvus which can be incIuded
in Is. inflata because of its very wide and inflated cupo
297
PERRI & FARABEGOLl: Conodonts across the Pennian - Triassic boundary in the Southem Alps
Occurrence: Werfen Fmn, Mazzin Mbr. Bulla section,
sample BU27. Tesero section, samples TS23-TS25.
R a n g e: lobata-isarcica zones: Lower Triassic, lower
Scythian.
Isarcicella isarcica (HUCKRIEDE 1958)
(pl. 4, figs 1-6)
1958
1964
1970
1991
1993
1997
1999
Spathognathodus isarcica HUCKRlEDE: 162, pl. IO, fig. 7
Spathognathodus isarcicus HUCKRlEDE - STAESCHE: 288, fig. 64.
Anchignathodus isarcicus (HUCKRIEDE) - SWEET: pl. I, figs 18-19.
lsarcice/la isarcica morphotype 3 (HUCKRIEDE) - PERRI: 42, pl.
4, fig. l, p1.6, figs 1-3.
lsarcice/la isarcica (Huckriede) - WANG & CAO: 254, pl. 55,
figs 8-9.
lsarcice/la isarcica (Huckriede) - WANG & WANG: 165, pl. 2,
fig. I.
lsarcice/la isarcica isarcica (HUCKRIEDE) - NICORA & PERRI: pl.
3, fig. 16.
Description: The Pa element is characterised by a wide
and inflated cup bearing on both sides one or a series of
nodes or denticles.
Derivatio nominis: In the Pa element the high asymmetrical inner side of the cup develops so as to form a
laterallobe.
Holotype: IC 1787-200961, figured in pl. 3, figs 21-23.
Paratypes: ICI777-200965 figured in pl. 2, figs 13; ICI785-200967, ICI786-200963, ICI788-200966,
IC1789-200962 figured in pl. 3, figs 15-20,24-29; IC176299175 figured in pl. 4, figs 12-14.
Locus typicus: Tesero section close to Tesero Village,
Province ofTrento, western Dolomites.
Stratum typicum: Level ofsample TS23 in the Tesero
section, 22.5 m from the base of the Werfen Fmn in the
Mazzin Mbr.
Repository: Museum G. Capellini, Dipartimento di
Scienze della Terra e Geologico-Ambientali, University
ofBologna.
Material: 61 specimens.
Remarks: Like Isarcicella lobata and Is. staeschei, this
species shows very wide intraspecific variability (PERRI
1991). It occurs in the Southern Alps after the entry of
Is. staescheì - in the Bulla section sample BU27 at 45 m
from the base ofthe Werfen Fmn, and in the Tesero section
sample TS26 at 26 m.
Occurrence: Werfen Fmn, Mazzin Mbr. Bulla section,
sample BU27-BU27 A. Tesero section, sample TS26; San
Pellegrino Pass section, sample SP2.
Range: isarcica Zone: Lower Triassic, lower Scythian.
Isarcicella lobata n.sp.
(pl. 2, figs 1-3; pl. 3, figs 15-29; pl. 4, figs 12-14)
1964
1981
1991
1991
1993
1995
1995
1996
1998
1999
1999
298
Spathognathodus isarcicus HUCKRIEDE - STAESCHE: text-fig. 61.
Hindeodus parvus (KOZUR & PJATAKOVA) - MATSUDA: 91, pl.
5, fig. 3.
Hindeodus parvus (KOZUR & PJATAKOVA) - SCHONLAUB: pl. l,
fig. 3.
lsarcice/la isarcica morphotype I (HUCKRIEDE) - PERRI: pl. 4,
fig. 8.
lsarcice/la turgida (KozuR, MOSTLER & RAHIMI-YAZD) - GULLO
& KOZUR: fig. 2/2-3.
lsarcice/la? turgida (KozUR, MOSTLER& RAHrMr-YAZD) - KOZUR:
pl. 2, fig. 8.
Hindeodus parvus (KOZUR & PJATAKOV A) -lsarcice/la Transition ? - METCALFE: pl. I, figs 12- 13.
lsarcicella? turgida (KozuR, MOSTLER & RAHIMI-YAZD) - KOZUR:
pl. 5, figs 7-8.
Hindeodus parvus morphotype 2 (KOZUR & PJATAKOVA) trans.
form lo lsarcice/la isarcica staeschei DAI & ZHANG - FARABEGOL! & PERRI: pl. 4.3.1, fig. 13.
Hindeodus parvus erectus KOZUR - NICORA & PERRI: pl. 3, fig. 7.
Hindeodus parvus parvus (KOZUR & PJATAKOVA) - NlcORA &
PERRI: pl. 3, fig. 14.
D i a g n o s i s : The Pa element is characterised by an asymmetrical to highly asymmetrical swollen and thickened
cup with a lateral bulge forming a lobe. The cup has no
lateral nodes or denticI es. The cusp is higher than the succeeding denticles.
Description: The blade bears 4-7 denticles rounded
in section and discrete, and shows the attachment area
ofthe denticles to be swollen. The wide cusp is twice as
high as the denticles. The posterior end ofthe unit can be
either adenticulate, abruptly ending, or denticulate with
Iittle denticIes decreasing in height towards the posterior
tipo The cup is always asymmetrical and swollen; it has
a Iaterai buige forming an inflated Iobe on the inner side.
The Iobe can be variousIy expressed because of high
variability in the width of the cup. The outer side of the
cup is usually more reduced than the inner side and may
show undulations. Stratigraphically higher specimens can
develop a secondary lobe on the outer side. The cup is
always without laterai nodes or denticles. The pit tends
to be centrally located.
Remarks: Two morphotypes can be distinguished, morphotype 1 (pl. 3, figs 21-23) and morphotype 2 (pl. 3, figs
15-17), on whether or not the posterior end is abrupt or
denticuiate. In laterai view the denticuiation is very close to
that of other species such as Hìndeodus parvus, Isarcicella
turgida and some elements ofIs. staeschei and Is. Isarcica.
In specimens with the cup highIy expanded, the appearance is very cIo se to that of Is. staeschei and Is. Isarcica
from which they are distinguished only by lack oflaterai
nodes or denti cles. Is. lobata includes forms regarded as
morphotype 1 ofthe apparatus of Is. isarcica by STAESCHE
Com. Forsch.-Inst. Senckenberg, 245, 2003
(1964), SWEEl (1977) and PFRRI (1991 ). Thc three morphotypes are discriminated by bearing on the cup l) no nodes
or denticles, 2) one or a series of nodes or dentic1es on
one side, 3) one or a series of nodes or denti cles on both
sides. They were considered to be elements of a unique
apparatus because of occurring consistently in association. Recent research such as on the Bulla and Tesero
sections, brought out that the three forms enter at different
stratigraphical levels. This implies branching into three
discrete species. Specimens previously assigned to Ili.
parl'lIS and 1.1'. tllrgida are here included in Is. lobata.
As discussed under the remarks on those species, forms
like that figured by GULLO & KOZUR (1993: fig. 2.3-3)
and KOZUR (1995: pl. 2, fig. 8 and 1996, pl. 5, figs 7-8)
- which have a much too asymmetrical and swollen cup
to be referred to Is. tllrgida - have been assigned to Is.
lobata. In PERRI'S opinion the GULLO & KOZtJR specimcn
shows features not fitting with the originai diagnosis and
characteristics of Is. turgida. Similarly, forms like that
figured by MATSUrlA (1981: pl. 5, fig. 3) as Hi. parvus
have to be included in Is. lobata. They were placed in
Hi. parl'US because of the denticulation and cusp being
c10se to Hi. parvus. though the basaI cavity was too widc
and swollen to be Hi. parvus. Both morphotypes of Hi.
parvus display an approximately symmetrical basaI
cavity and no swelling. According to PERRI, forms with
a swollen and asymmetrical cup recalling 1.1'. staeschei
and 1.1'. isarcica ne ed to bc recognized as discretc species. The entry of Is. lobata moreover is stratigraphically
higher than the entry of Is. turgida. Is. lobata oecurs in
the Bulla section from sample BU23 at 30 m from thc
base of the Werfcn Fmn up to 8U27 A at 46 m, and in
the Tesero section from TS23 at 22.5 m to TS26 at 26
m. In both sections Is. lohata enters some metres below
the first appearance of 1s. staeschei and co-occurs with
Is. staeschei and the higher 1s. isarcica. The first oceurrence of Is. lohata has biostratigraphic significance and
is uscd to define the base of a new biozone betwecn the
parvus and staeschei zones.
Paratype: TCI775-989162 figurcd in p!. 1, figs 26-28.
Loeus typicus: Bulla section at about 0.5 km along
the road NW of the Bulla village, Province of Bolzano,
Western Dolomites.
Stratum typicum: Level ofsample BUIO ofthe Bulla
section at 20 cm from the base ofthe Werfen Fmn in the
Tesero Mbr.
Repository: Museum G. Capellini, Dipartimento di
Scienze della Terra e Geologico-Ambientali, University
ofRologna.
Material: 2 spccimens.
Diagnosis: The Pa element is characterised by 5-6
very wide and discrete dcnticles with the profile of the
apices extremely rounded. The wide and squat cusp, corresponding to about 1/3 ofthe unit length, is higher than
the dentic1es. Some Iittle denticles or "germinai denticles"
ean be present anterior to the cusp. The wide spoon-shaped
basai cavity is slightly swollen.
Description: Very robust, large formo Tn latera l view
the profile of the denticles is straight with abruptly ending posterior tip in the juvenilc speeimen, whereas it is
rounded prescnting dentielcs deereasing in height up to the
posterior end in the adult element, the holotype. The very
wide and straight eusp is twiee the height ofthe denticles.
Cusp and dcnticles show striae. Thc widc spoon-shaped
basai cavity is slightly asymmetrical, displaying a wcak
swelling\thickening. The pit is located in the anteriorthird
of the basai cavity.
Isarcicella peculiaris n. sp.
(p!. l, figs 26-31)
Remarks: Even though only two specimens have been
[ound, these have been assigned to a new species because
ofthe extremcly pcculiar shape. The swelling ofthe basai
cavity is close to that of Isarcicella prisca and Is. turgida.
Is. peculiaris is dift'erentiated from Is. prisca by having a
wider and squatter cusp occupying Lｾ＠ of the length (p!. l,
figs 26, 29) whereas in Is. prisco the cusp occupies only
V. or the length. In the new species therc is no evidence
ofthe largest denticles being located in the second half of
the bIade; this is characteristie of Is. prisca.
No conodont cJose to these forms has becn previously
described and figured. The element assigned to Hindeodus
cf. parvus parvus, figured and described by K07UR (1996:
pl. 3, fig. Il), is a specimen with unusually broad, distally
rounded denticles; it was obtained from a fioat bloek in
Sicily (sample KS3). Tt resembles the new species only
with respect to its denticJes.
Derivatio nominis: Referring to the extremelypeculiar
shapc of thc Pa elemcnt of the species.
Occurrence: Werfen Fmn, Tesero Mbr. Bulla section,
sample BUIO.
Ho I otype: ICI776-989157, figured in pl. 1, figs 29-31.
Range: Upper praeparvus Zone: upperrnost Permian.
Occurrence: Werfen Fmn, Mazzin Mbr. Bulla section, samples 8U23-BU27 A; Tesero section, samples
TS23-TS26; San Pellegrino Pass section, sample SP2;
Agordo-Frassené section, sample AF II.
Range: lobata-isarcica zones: Lower Triassic, lower
Scythian.
299
PERRI & FARABEGOLl: Conodonts across the Pennian - Triassic boundary in the Southem Alps
Isarcicella prisca KOZUR 1995
(pl. l, figs 20-25; p1. 2, figs 16-21)
1991
1995
1996
1998
1998
Hindeodus typicalis (SWEET) - PERRI: 40, pl. 3, figs l, 3, 4.
Isarcicella? prisca KOZUR: 73, pl. 2, fig. l.
Isarcicella? prisca KOZUR - KOZUR: lO l, pl. 4, fig. 9.
Isarcicella? prisca morphotype l KOZUR - FARAREGOLI & PERRI:
pl. 4.3.1, fig. 6.
Isarcicella? prisca morphotype 2 KOZUR - FARABEGOLI & PERRI:
pl. 4.3.1, fig. 7.
D e s c r i p t i o n: The Pa element has largest dentici es in the
second half of the biade, a broad cusp, the inner side of
the basai cavity distinctly thickened, and the rather broad
outer si de less thickened.
Remarks: KOZUR (1995) chose the specimen figured by
PERRI (1991: pl. 3, fig. l, sample BUl O) as holotype ofthe
species. Denticulation ofthe species is extremely close to
that of HL praeparvus from which it can be distinguished
by the sweIling and thickening ofthe basaI cavity, mainly
on its inner side. Several specimens have been found in the
studied material.1s. prisca has been utilised here to divide
the praeparvus Zone into Lower and Upper praeparvus
zones. The first occurrence ofIs. prisca is stratigraphicaIly
higher than that of HL praeparvus and defines the base
ofthe U. praeparvus Zone. In the Bulla section both Hi.
praeparvus and 1s. prisca are present in sample BU9, the
first productive sample, about 20 cm above the base ofthe
Werfen Fmn In the Tesero section Is. prisca first occurs
in sample TS9 whereas Hi. praeparvus is found lower in
the sequence at sample TES62, 30 cm from the base of
the Werfen Fmn (Fig. 4). In sections from the western
Carnic Alps, beds ofthe Bellerophon Fmn producing Ri.
praeparvus yielded no Is. prisca.
The species can be divided into two morphotypes.
Morphotype l includes all forms with the posterior tip
ending abruptly, and morphotype 2 with tiny denticles
occurring to the posteri or end.
Occurrence: Werfen Fmn, Tesero Mbr. Bulla section,
samples BU9-BUI2C; Tesero section, samples TS9,
TESIOI.
R a n g e: U pper praeparvus-parvus zones: uppermost
Permian-Lower Triassic, lowermost Scythian.
Isarcicella staeschei DAI & ZHANG 1989
(pl. 3, figs 1-14; pl. 4, figs 7-9)
1964
1975
1981
1989
1991
300
Spathognathodus isarcicus HUCKRIEDE - STAESCIIE: 288, figs
62-63.
lsarcicella isarcicus (HUCKRIEDE) - KozuR, MOSTLER & RAHIMIYAZD: 6, figs 4-5,8.
lsarcicella isarcica (HUCKRIEDE) - MATSUDA: 93, pl. 5, figs 47.
lsarcicella staeschei DAI & ZHANG: 430, pl. 45, figs 16-17, pl.
46, figs4-7, 11-13, 18-19, pl. 53, figs 13-14.
Isarcicella isarcica morphotype 2 (HUCKRIEDE) - PERRI: 42, pl.
1993
1995
1997
1998
1998
1999
4, figs 2-6, pl. 5, figs 1-3, pl. 6, figs 4-5.
Isarcicella isarcica (HUCKRIEOE) - TIAN: pl. I, figs 22-24.
Isarcicella isarcica staeschei DAI & ZHANG - KOZUR: pl.6, fig.
18.
Isarcicella staeschei DAI & ZHANG WANG & WANG: pl. 2, figs
2-5.
Isarcicella isarcica staeschei DAI & ZHANG - FARABEGOLI &
PERRI: pl. 4.3.1, figs 14-15.
Isarcicella staeschei DAI & ZHANG - ORCHARO & KRYSTYN: pl.
6, figs 4-8, 10-12.
lsarcicella isarcica staeschei DAI & ZHANG - NIl'ORA & PERRI:
pl. 3, fig. 15.
Description: The Pa element is characterised by a very
asymmetrical wide and inflated cup with a lateral lobe
bearing one or a series of nodes or denticles.
Remarks: The species shows high morphologic intraspecific variability (PERRI, 1991). In the Southem Alps
it occurs first after the entry of 1sarcicella lobata at 43.60
m from the base of the Werfen Fmn in the Bulla section
(sample BU25A) and at 24 m in the Tesero section (sample
TS24). One element with a rudimentary latera1 denticle,
found in the Tesero section (sample TS19) 14.30 m below
the entry in abundance of Is. staeschei (sampleTS24), was
previously assigned to 1s. isarcica morphotype 2 (= Is.
staeschei) in PERRI (1991). Here it has been considered a
transitional form towards Is. staeschei but not yet a true
representative of that species because it displays no evidence of swelling of the basaI cavity. It is stressed that at
the leve! of first occurrence of Is. staeschei in both the
Bulla (sample BU25A) and Tesero (sample TS24) sections,
forms with one denticle on one side ofthe cup have been
found in association with e1ements showing a lateral series
of denticles different from the first representatives of Is.
staeschei - having a single lateral denticle - found in Spiti
(India) and Kashmir (KOZUR 1996; WANG 1996; ORCHARD
& KRYSTYN 1998). Moreover, in the Alps, Pa e1ements of
Is. staeschei bearing more than one denticle on one side
ofthe cup occur stratigraphically lower than Pa elements
of Is. isarcica. Therefore the data from the Alps disagree
with Kozur's (1996) opinion that the forms of Is. staschei
bearing two or more denti cles exclusively co-occur with
Is. isarcica and that only specimens with one denticle on
one side ofthe cup have a different (lower) range probably
representing a different taxon.
Occurrence: Werfen Fmn, Mazzin Mbr. Bulla section,
samples BU25A-27A; Tesero section, samples TS24-26;
San Pellegrino Pass section, sample PSP2; Agordo-Frassené section, sample AF Il.
Range: staeschei-isarcica zones: Lower Triassic, 10wer
Scythian.
Coue Forsch.-Inst.
Isarcicella turgida
(KozuR, MOSTLER & RAHIMI-YAZD 1975)
(pl. 2, figs 13-15)
1975
1993
1995
1996
Anchigna/hodus lurgidus KOZUR, MOSTLER & RAHIMI-YAZD: 5.
pL 7, figs 11-12.
Isarcicella turgida (KOZUR, M05TLER & RAHIMI-YAlD)- GULLO
& KOZUR: fig. 217-9.
lsarcicella? turgida (KozuR, MOSTLER & RAHIJl.ll- YAlD)- KOZUR:
72, pl. 2, fig. 5.
Isarcicella? turgida (KOZUR, MOSTLER & RAHIMI- YAlD) - KOlUR:
p.IOO. pl. 4, fig. 8.
Descri ption: The Pa element has a thickened bIade and
slightly asymmetrical basai cavity. The upper part of the
swollen basai cavity is completely smooth,
Rem arks: KOZUR (1975) defined Isarcicella turgida as a
conodont with a swollen biade and a slightly asymmetrical
basaI cavity. The description fits with the holotype (KOZUR
1975: pl. 7, fig. 12) in which the bIade is weakly inflated
and the asymmetry ofthe basai cavity is unrecognisable.
Together with the holotype, KOZUR (1975: pl. 7, fig, 11)
figured another specimen with biade more swollen, but
with the basai cavity always slightly asymmetrical. GULLO
& KOZUR (1993: fig. 2/2-3) figured a specimen from the
Sosio Valley ofSicily with a very wide asymmetrical and
swollen cup, It was assigned to Is. furgida. KOZUR (1995,
1996) interpreted the species very broadly, enlarging the
originaI concept to include forms like that found in Sicily
which, subsequently, he took to be representative of Is.
turgida. PERRI considers that form very different from the
holotype of Is. turgida. In the studied material, specimens
like that found in Sicily appear 13.60 m in the Bulla section
and 1.5 m in the Tesero section below the first occurrence
of Is. staeschei and co-occur with both Is. staeschei and Is.
isarcica. According to PERRI'S point ofview, these forms
- included in Is. lobata correspond to the adenticulate
elements of morphotype I of the apparatus of Is. ìsarcica
OfSTAESCHE (1964), SWEET (1977) and PERRI (1991) as
discussed under the remarks on Is. lobala. They seem to
derive from representatives of the species Is. turgida by
increase in the width and asymmetry of the basaI cavity
(which becomes a cup bearing a lobe). Here Is. turgida is
restricted to forms dose to the originai diagnosis; more
asymmetrical and inflated forms appearing stratigraphically higher have been assigned to Is. lobata.
KOZUR (1995) indicated that the distinctive character
for discriminating between Hi. parvus and Is. turgida is
only the swelling of the biade; the denticulation is the
same. Forms corresponding to the originai diagnosis of
Is. turgida have been found in the Bulla section in the
same level (BU13B) where Ili. parvus first occurs. These
specimens in lateral view are very dose to HL parvus, but
in upper view show slight asymmetry ofthe basai cavity,
thickened mainly on the inner side.
Occurrence: Werfen Fmn, Tesero and "\1azzin Mbrs.
245,2003
Bulla section, samples BUI2B, BU23; Tesero section,
samples TSI9, TS23.
Range: parvus-lobata zones: Lower Triassic, lower
Scythian.
Acknowledgements
We are deeply grateful to KIRIL BUDUROV and MIKE ORC'HARD for thoroughly rcvicwing the manuscript and making fruitful suggcstions, and to JOHN TALENT t'Or editing
and criticai reading of the manuscript. We thank CARLO
BRAZZOROTTO who undertook the heavy liquid separations,
PAOLO FERRIERI who took SEM photos of conodonts, and
Dr. BENIAMINO COSTANTI\\[ who helped generate the computer drawings. The Agorolo-Frassené and Maestrin sections
were sampled with ALDA NICORA and the San Pellegrino
Pass section with CLAUDIO NERI. PERRI is responsible for the
conodont data and discussion; FARABEGOLI is responsible
forthe geological and sedimentological parts ofthe paper;
condusions were written in collaboration. The research
was funded by ex 60% "\1URST grant to M.C. PERRI.
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Cour. Forsch.-Inst.
245,2003
Plates
305
PERRI & F ARABEGOU: Conodonts across the Pennian - Triassic boundary in the Southem Alps
Plate 1
Ali magnifications are x 100
Figs 1-12
Hindeodus pisai n. sp.
1-3. Lateral, lower and upper views respectively oflCI77l-989l78; Bulla: BU12A.
4--6. Lateral, lower and upper views respectively oflC1772-989l8l; Bulla: BUI2B.
7-9. Holotype, lower, upper and lateral views respectively ofIC1749-989l76; Bulla: BU12A.
10-12. upper, lower and lateral views respectively ofIC1773-989l77; Bulla: BUI2A.
Figs 13-16
Hindeodus praeparvus KOZUR 1996.
13-14. Morphotype I, lateral and upper views respectively ofICI 742-989 160; Bulla: BU lO.
15-16. Morphotype 2, lateral and upper views respectively ofICI744-989173; Bulla: BUI2A.
Figs 17-19
Isarcicella changxingensis (WANG 1995).
Lateral, upper and lower views respectively ofICI759-990087; Tesero: TES71.
Figs 20-25
Isarcicella prisca KOZUR 1995.
20-22. Morphotype I, lower, upper and lateral views respectively ICI774-200971; Bulla:
BUl2B.
23-25. Morphotype 1, lower, upper and lateral views respectively ICI746-989182; Bulla:
BUI2B.
Figs 26-31
Isarcicella peculiaris n. sp.
26-28. Morphotype I, lateral, lower and upper views respectively of ICl775-989l62; Bulla:
BUIO.
29-31. Holotype, Morphotype 2, lateral, lower and upper views respectively ofICl776-989157;
Bulla: BU lO.
306
Cour. Forsch.-lnst. Senckenberg, 245 , 2003
Plate l
PERRI & FARABEGOU: Conodonts across the Permian
Triassic boundary in the Southcm A1ps
ｾＭ
Plate 2
Ali magnifications are x l 00
Figs 1-3
lsarcicella lobata n. sp.
Morphotype l, lateral, lower and upper views respectively ofIC I 777-200965; Tesero: TS24.
Figs 4-12
Hindodus parvus (KOZUR & PJATAKOVA 1976).
4-6. Morphotype 2, lateral, lower and upper views respeetively ofICI761-990086; Tesero: TS 19.
7-9. Morphotype I, lower, upper and lateral, views respectively of IC 1751-989194; Bulla:
BUl3B.
10-12. Morphotype I, lateral, lower and upper views respeetively oflCI750-989191; Bulla:
BUl2B.
Figs 13-15
lsarcicella turgida (KozuR, MOSTLER & RAHIMI- YAZD 1975).
Morphotype l, lower, upper and lateral views respeetively ofiC 1778-989184; Bulla: BUI2B.
Figs 16-21
lsarcicella prisca KOZUR 1995.
16 18. Morphotype 2, lateral, 10wer and upper views respeetively of IC 1747 -989169; Bulla:
BUI2A.
19-21. Morphotype 2, lower, upper and lateral views respeetively of IC 1779-989170; Bulla:
BUI2A.
Figs 22-36
Hindeodus praeparvus KOZUR 1996.
22-24. Transitional form to Hi. parvus, upper, lower and lateral views respectively of IC 1748989164; Bulla: BUI2A.
25-27. Morphotype 2, lateral, lower and upper views respeetively of ICI780-989163; Bulla:
BU12A.
28-30. Morphotype I, lateral, lower and upper views respeetively of ICl781-989168; Bulla:
BUl2A.
31-33. Morphotype 2, lower, upper and lateral views respectively of IC 1745-989183; Bulla:
BUl2B.
34-36. Morphotype l, lower, upper and lateral views respeetively of IC1743-989156; Bulla:
BUIO.
308
Cour. Forsch.-Inst. Senckenberg, 245 , 2003
Plate 2
PERRI & FARABEGOU: Conodonts across the Permian - Triassic
in tbe Southern
Plate 3
Ali magnifications are x I 00
Figs 1-14
Isarcicella staeschei DAI & ZHANG 1989.
l, 2. Lateral and upper views respectively of IC 1755-989196; Bulla: BU25A.
3-5. Lower, lateral and upper views respectively oflCI782-200970; Tesero: TS24.
6-8. Lower, upper and lateral views respectively oflC 1754-989197; Bulla: BU25A.
9-11. Lateral, lower and upper views respectively ofICI783-200969; Tesero: TS24.
12-14. Upper, lower and lateral views respectively ofIC 1784-200968; Tesero: TS24.
Figs 15-29
Isarcicella lobata n. sp.
15-17. Morphotype 2, lateral, lower and upper views respectively of ICI785-200967: Tesero: TS24.
18-20. Morphotype 1, lateral, lower and upper views respectively or ICI786-200963: Tesero: TS24.
21-23. Holotype, morphotype l, lateral, upper and lower views respectively of ICI787-200961; Tesero:
TS24.
24-26. Morphotype l, lower, upper and lateral views respectively ofIC 1788-200966; Tesero: TS24.
27-29. Morphotype I, lower, upper and lateral views respectively ofIC1789-200962; Tesero: TS24.
310
Cour. Forsch.-Inst. Senckenberg, 245 , 2003
Plate 3
PERRI & FARABEGOU: Conodonts across the Pennian - Triassic
hrulnrt",1"V
in the Southem
Plate 4
AlI magnifications are x 100
Figs 1-6
lsarcicella isarcica (HUCKRTEDE 1958).
1-3. Lateral, lower and upper views respectively of ICI466-9832; Bulla: BU27.
4-6. Lower, upper and lateral views respectively of IC 1451-8940; Bulla: BU27.
Figs 7-9
lsarcicella staeschei DAI & ZHANG 1989.
7-9. Lower, lateral and upper views respectively ofIC1453-8939; Tesero: TS26.
12-14
Figs 10-11, 15-19
312
lsarcicella lobata n. sp.
Morphotype 2, lateral, lower and upper views respectively of IC1762-99175; Tesero:
TS24.
lsarcicella inflata n. sp.
10-11. Morphotype 1, lateral and upper views respectively ofICI464-8924; Bulla: BU27.
15-17. Holotype, Morphotype l, lateral, lower and upper views respectively oflC 1465-8923;
Bulla: BU27.
18-19. Morphotype 2, upper and lateral views respectively of lC 1460-8922; Tesero:
TS23.
Cour. Forsch.-Inst. Senckenberg, 245, 2003
Plate 4
Enzo Farabegoli