I

Asiatic Herpetological Research

December 1993

Vol. 5, pp. 3 1-44

J

Holocene anurans from Caucasus

ZBYNEK rocek
Department of Paleontology, Geological Institute, Czechoslovak Academy of Sciences, Rozvojova 135, CS-165
00 Prague, Czech Republic

NW

Abstract. -Holocene deposits of the Kudaro I Cave from the vicinity of Ertso Lake (South Ossetia,
Caucasus) yielded, among others, rather numerous disarticulated anuran bones. Examination of this sample

revealed that majority of this material belongs to the genus Bufo and to the family Ranidae. This generally
corresponds to the composition of the contemporary anuran fauna of that region.

Key Words: Anura, Holocene, Caucasus, osteology.

Introduction

7B). Probably also ZIL/EL/1 40 (Fig. 7 A).

The material described in the present
paper was recovered from the deposits ofthe cave Kudaro I. The cave is on the slope

Description: The margin of the cavitas
glenoidalis is elevated. Although both thepars acromialis and proc. glenoidalis arebroken off it is obvious that there was a
deep incision between them. There is amoderately prominent and rather pointed
outgrowth on the margo anterior. Bothscapulae are comparatively big elements
corresponding by their size to the below

of Mt. Chasavalskaya (1600 m altitude),valley of Dzhordzhori River, in the vicinityof Ertso Lake, about 90 km NE from thetown Kutaisi, South Ossetia, near

NW

Caucasus
Rachinsky ridge in the
also
42°
see
43°
W;
N,
Lyubin(approx.
1980a). The deposits are of the Holocene

described humeri and ilia.

age (Lyubin 1980b).
Material:
First description of the anuran and reptile

Humeri ZIL/EL/21 (Fig. 2B),

ZIL/EL/25, ZIL/EL/55,

material from this cave was published by(1980).

DarevskyHis

taxonomic

assignments generally agree with those in
the present paper.

The material consists of

ZIL/EL/79, ZIL/EL/1 12 (Fig. 2A),
ZIL/EL/132 (Fig. 2C), ZIL/EL/139,
ZIL/EL/153, ZIL/EL/173.

isolated bones; anuran bones described inthis paper are deposited in the Zoological

These specimens (except for ZIL/EL/153

ofthat includes also the most distal section of

Sciences, St. Petersburg, under collection

the crista ventralis) are preserved only as
distal parts of the humerus. All of them are

InstituteoftheUSSR Academy

numbers ZIL/EL/1 - ZIL/EL/185.

by conspicuous degree ofofthe epicondylus medialis,development
the distal margin of which extends almost
to the level of the distal margin of the caput
humeri. Hence, there is a distinct notch
between the both structures that cancontinue onto the dorsal surface of the
distal section of the bone. On the ventral
surface of the medial epicondylus one candiscern an indistinct ridge running onto its

characteristic

Anatomical terminology mostly follows
that of Bolkay (1919) and Gaupp (1896).
It should be noted that most of elements inthe sample are postcranial bones and only
few cranial bones are present. Fragmentarymaterial bearing no diagnostic characters
was excluded from the account below.

Systematic Paleontology

distal surface.

Bufo sp.
Material:

Left scapula, ZIL/EL/7 (Fig.

1993 by Asiatic Herpetological Research

The lateral epicondylus isdeveloped in lesser degree, extending
laterally from the outline marked by thecrista lateralis in some specimens (see Fig.

Asiatic Herpetological ResearchVol. 5, p. 32

2A, B). The lateral surface of the caput
humeri is worn away in larger specimens,so the ball is not complete. The wholedistal end of the bone is bent ventrally; thisis correspondingly reflected on its dorsal
surfacewhich

islongitudinal axis.

convexalong

itsSome variation may beobserved concerning the extent of themedial and lateral cristae which might be,
however, assigned to secondary sex
differences. This might be suggested also
by small specimen ZIL/EL/55 which may
represent an immature individual, and inwhich both cristae are lacking. On theother hand, all specimens have their crista
medialis directed laterally, so its ventral
surface meets the medial surface of theIndiaphysis almost perpendicularly.
ZIL/EL/25 and ZIL/EL/132 the margins ofboth cristae are rather undulated andthickened.

December 1993

be abradedspecimens (e.g., ZIL/EL/62).

olecranon

may

in

large

On the inner surface of the bone, close to
the point where the margins of thearticulation cavity of the olecranon andcapitulum meet with one another, is a small
but deep pit. Similar pit is lacking or not so
deep in ranids, but regularly present in
Bufo. It serves as a muscle insertion area
and in some specimens may be doubled.
The posterior margin of the bone (i.e., ofitsulnar part) bears an indistinct crista insome specimens.
Material:

ZIL/EL/75,
ZIL/EL/97,

Ilia

ZIL/EL/41, ZIL/EL/66,

ZIL/EL/94
ZIL/EL/99

(Fig.
(Fig.

4A),
4B),

ZIL/EL/1 24, ZIL/EL/1 29.
Description:

The

ala

ossis

ilii,

if

compared with the posterior part of the
These humeri (preserved

In large with thedetermination of size-corresponding ilia.

bone, are stout (ZIL/EL/94) or rather
slender (ZIL/EL/99). Their dorsal margin
is rounded, only in the mid-part it becomesan edge distinctly bent medially along its

However, some (esp. smaller) humeri may

whole extent.

the variation range of the ranids,
but the latter assignment lacks reliable

compressed dorsoventrally, ellipsoid

foundation if only distal part of the bone isala, approximately at the level of the highestbepoint of their arch, there is an orifice of thenarrow horizontal canal coming onto theDiscussion:

only as distal sections) are morphologicallyclosest to those of Bufo.

fall intoat the disposal.

Discoglossids may
excluded because their medial and lateralcristae are confluent with the diaphysis,
with no distinct border. The only fossil
anuran that is of similar size as ZIL/EL/1 12

Latonia seyfredi v. Mayer (=Discoglossus giganteus WettsteinWesterheimb). However, its morphology
isand stratigraphic range are different (seee.g.

Mlynarski, 1976, pi. 1/4).

Material:
Radioulnae ZIL/EL/43,
ZIL/EL/62 (Fig. 8 A), ZIL/EL/1 06 (Fig.
8B), ZIL/EL/1 16, ZIL/EL/120 (Fig. 8C),

cross-section.

In its anterior part, the ala isin

On the medial surface of thebone surface from the posterior. There iscertain variation in the location andmorphology of this canal - it may continue
as a groove for a short distanceanteriorwards, and the orifice may belocated on the bottom of a horizontal
depression developed below the above-

mentioned edge.

The depression may

terminate anteriorly on the dorsal surface of
the ala or, in some specimens (esp. smallerones), the orifice is located posterior to the

The margin of the
Description:
olecranon rimming the articular cavity is

The torus superior isdepression.
extensive, with two to three tubercles
continuing (except the most anterior one) asa short and low ridges laterally.
The
anterior-most tubercle continues as a shortformed either by calcified cartilage orossified tissue lacking periostal bone. The

medial surface of the bone.

ZIL/EL/1 62.

border between the smooth periostal boneand rough surface rimming (and also
covering) the articular cavity is distinctive.
It seems that this mostproximal part of therounded ridge anteroventrally, onto theZIL/EL/1 24(and some other ones) is much smaller but
except for size its morphology corresponds
in all principal features to that describedabove.

Asiatic Herpetological Research

December 1993

Vol. 5, p. 33

«&&

CM

EM

A- Rana sp. (ZIL/EL/148). Left humerus in ventral, dorsal and lateral views (from the left to
B-fianasp. (ZIL/EL/151). Left humenis in ventral, dorsal and lateral view (from the left to right).
C- Rana sp. (ZIL/EL/74). Ventral (left) and dorsal (right) view of the proximal section of therighthumerus (drawing reversed for comparison). Bar equals 1mm. Abbreviations: C. A. - crista adventiva;
C.L. - crista lateralis; C. M. - crista medialis; C. V. - crista ventralis; E. M. - epicondylus medialis; F. D.
FIG.1.
right).

fossula dividens; P. L. -processus lingualis.

The shape of the ilium

orifice may be doubled, and the differences

corresponds to that in contemporary Bufoin this respect may be found also between

bufo and B. viridis. The only difference isthat in both latter forms the longitudinal
depression on the medial surface of the ala

theDiscussion:much lesser degree (due toleft and right ilium of a single
individual). The same seems to hold forSize differences may befossil material.

noted that the orifice of the mentioned canalon the medial surface of the bone displays

ascribed either to interspecific variation or
to differences between both sexes (the lattermay reach quite a high degree in somecontemporary representatives of the genus

certain variation in contemporary forms (theBufo.

is developed inlesser extent of the edge).

It should

also beAsiatic Herpetological ResearchVol. 5, p. 34

December 1993

D

B

FIG. 2. A- Bufo sp. Right humerus (ZIL/EL/1 12). B- Bufo sp. Right humerus (ZIL/EL/21). C- Bufo
sp. Left humerus, drawing reversed for comparison (ZIL/EL/1 32). D- Ranidae indet. Right humerus(ZIL/EL/150). E- Ranidae indet. Right humerus (ZIL/EL/1 10). Bar is 1 mm.

Rana sp.
Humeri ZIL/EL/56, ZIL/EL/74
1C), ZIL/EL/83, ZIL/EL/128,(Fig.
ZIL/EL/1 48 (Fig. 1A), ZIL/EL/1 51 (Fig.
Material:

IB), ZIL/EL/1 79.

Description:

The crista ventralis andcrista ad ventiva delimit a wide, shallow andrather longitudinal depression for muscle
insertion.
The latter crista may be

developed to various degree, whereas thecrista ventralis is well developed in nearlylongitudinal groove on the dorsal surface of
The proc. lingularis and thethe bone.
outgrowth produced by the crista adventiva

may form together a roof over the fossula
dividens; this nearly results in a canal. The
caput humeri is well prominent ventrally(clearly seen in lateral aspect). Although
the distal part of the bone is straight, thecrista medialis and lateralis make itseemingly "S" shaped. Lateral epicondylus
Other features may be
is entirely absent.
seen in Fig. IB.

individuals, with a distinctive lingual

Some variation may be observed, mainly

process (only in ZIL/EL/128 this process ispoorly developed, and the crista ventralis
continues distally as a gradually lowering
ridge). The crista ventralis has a hollow

in the degree of development of the lingual

allinside its free margin; consequently,

italong its attachment to thediaphysis. This is manifested externally bygrooves along the insertion of the crista, on:ither side.

The crista lateralis and medialis

directed

dorsally,

adventiva.

isthinnerareprocess and in the extent of the medial andlateral cristae, as well as of the crista

forming thus a

Anatomical notes: As may be deduced
from the condition in Rana esculenta(Gaupp, 1896) the depression between thecrista ventralis and adventiva could serve as
an area of insertion for three heads of theAsiatic Herpetological Research

December 1993

Vol. 5, p. 35

FIG. 3. A- Ranidae indet. Right ilium in lateral view (ZIL/EL/87). B- Ranidae indet. Left ilium in lateralview (ZIL/EL/1). C- Ranidae indet. Lett ilium in lateral view (ZIL/EL/14). D- Anura indet. Left ilium inreversed for comparison. Bar equals 1mm. Abbreviations: A. lateral view (ZIL/EL/3). B, C, andD

acetabulum; A. O.

I.

-

ala ossis

ilii;

C. O.

I.

-

crista ossis

ilii;

P. A.

-

pars ascendens; P. C.

-

parscylindriformis; P. D. pars descendens; T. S.- tuber superius.

m.

pectoralis (portio epicoracoidea,
and abdominalis), whereas theproximal part of the crista itself (its edge)
could serve for attachment of two heads ofthe m. deltoideus (pars clavicularis and
The third head of thescapularis).
deltoideus muscle (pars episternalis) isfixed to the ventral ridge of the medial
The fossula dividens
epicondylus.
for tendon of the m.
servedprobably
coracoradialis. The groove between boththe crista medialis and lateralis served no
doubt for insertion of the caput profundumof the m. anconeus. The medial crista
serves in anurans for attachment of the m.

itssternalissex character.

flexor carpi radialis and the lateral crista for(Italy). In other individuals, the cristae andthe lingual process were developed tovarious degree, similar to the condition in
the described fossil material. In all cases

the m. extensor carpi radialis.

has

its

specialThe former

function in amplexus.

Consequently, the crista medialis is usually
better developed in males, and the degree ofdevelopment is considered secondary

Taxonomic notes: Humeri recallingthose described above may be found insome individuals of contemporary "brown"
frogs, i.e. of Rana temporaria R. arvalis,dalmatina, R. latastei, and R
macrocnemis. I found morphology closely
resembling that in ZIL/EL/1 51 (Fig. IB) inR.

.

contemporary Rana arvalis (DP FNSP
5830) from S Bohemia (Czechoslovakia),
in R. arvalis wolterstorfii (DPFNSP 6264)

from Soroksar (Hungary), andlatastei (DP FNSP 6419) fromthese humeri belonged to males.

in

R.

Como

Hence, itAsiatic Herpetological ResearchVol. 5, p. 36

December 1993

FIG. 4. A- Bufo sp. Right ilium in lateral view (ZIL/EL/94). B-Bufo sp. Left ilium in lateral view
(ZIL/EL/99). C- Ranidae indet. Left ilium in lateral view (ZIL/EL/68). D- Ranidae indet. Right ilium inlateral view (ZIL/EL/1 11). B and C reversed for comparison. Bar equals 1 mm.

may be concluded that the describedcharacters on the humerus may be ascribedto sexual

dimorphism, but they are notpresent in all males. In any case, relations
to contemporary "brown" ranids seems tobe beyond any doubt. It is quite possiblethat the above described humeri assigned toRana sp. and humeri (and other elements)
described below as Ranidae indet. might
belong to a single form.

surrounded by marginal part for the
cartilago paraglenoidalis that connects thisbone with the proc. glenoidalis scapulae.

The pars epicoracoidealis is nearly regularly
arch-like, exceeding by its antero-posterior
diameter the lateral part of the bone.

Material: Humeri ZIL/EL/1 10 (Fig. 2E),
ZIL/EL/1 50 (Fig. 2D), ZIL/EL/1 68.

Description:

Ranidae indet.

similar)Material:

Right coracoid, ZIL/EL/54(Fig. 7E).

Description:The

intumescentia

glenoidalis is circular, with distinct but
small fovea acetabuli where the ligament of
the humerus is inserted.
The fovea is

The crista ventralis humeri

in ZIL/EL/1 10 (and in ZIL/EL/1 68 that is
isprominent, gradually lowering

Part of its margin is tongue-likebent medially. Only within the proximal
section of the crista there is a groove along
the medial surface of its basis. The crista
medialis is well developed, but only in the
distally.

The lateral crista isdegree. The medial

distal third of the bone.

developed

in lesser

December 1993

Asiatic Herpetological ResearchVol. 5, p. 37

Ranidae indet. Vertebrae in ventral view. A- Sacral vertebra (ZIL/EL/10). B- V6 (ZIL/EL/70).
C- V2 (ZIL/EL/8). Bar equals 1 mm. Fig.6 Ranidae indet. Praesacral vertebrae in dorsal (above) andventral (below) views. A- V5 (ZIL/EL/17). B- V2 (ZIL/EL/125). Bar equals 1 mm.
FIG. 5.

Asiatic Herpetological ResearchVol. 5, p. 38

epicondylus is well developed, the lateral
one is rudimentary. On the medial surfaceof the diaphysis there is low but discernible
crista.
Specimen ZIL/EL/150 has its
ventral crista well developed, with rounded
and almost straight margin. It is thin along
its attachment to the diaphysis.
This,
together with the fact that the diaphysis isoval in cross-section, caused that there is agroove along the basis of the crista,
especially well developed on its medialsurface. Neither medial nor lateral crista
are developed in this specimen, and theDecember 1993

separating thus the tuber from the crista.
The acetabulum is with acute and prominent

margins, considerably lifted above the parsdescendens, but rather slanting down
towards the pars ascendens. However,even here the margin of the acetabulum isrepresented by a distinct ridge. There is aconsiderable notch between the tuber
superius and the dorsal margin of the pars
ascendens. The dorsal margin of the pars
ascendens continues anteroventrally onto
the medial surface of the bone as a roundedridge that disappears after a short distance.

ZIL/EL/29, ZIL/EL/49, ZIL/EL/68 (Fig.

ZIL/EL/1 (Fig. 3B) is essentially the samebut the crista is much lower than is the
dorsoventral diameter of the pars
cylindriformis, in whole its extent anterior
to the tuber superius. Anteriorwards it iseven getting lower, so its transition into thedorsal margin of the pars cylindriformis isindistinct.
The tuber is prominent and4C), ZIL/EL/87 (Fig. 3 A), ZIL/EL/1 11

extensive anteroposteriorly.

ZIL/EL/135, ZIL/EL/164,(Fig.
ZIL/EL/177.
Specimens ZIL/EL/2,
ZIL/EL/19, ZIL/EL/29, ZIL/EL/135 andthe crista ossis ilii is an insertion area for

ZIL/EL/177 are similar to ZIL/EL/1 1 1 (Fig.

thelateral epicondylus is absent, too.

In spiteof these differences, both humeri may be
assigned to the Ranidae.
Material:

ZIL/EL/1

Ilia

(Fig.

3B),

ZIL/EL/2, ZIL/EL/14 (Fig. 3C),
ZIL/EL/18, ZIL/EL/19, ZIL/EL/28,

4D),4D) in that the tuber superius is continuouswith the crista ilii.

Anatomical notes: The lateral surface ofm. iliacus externus, the other end ofwhich is fixed by a tendon to the proximal
part of the femur (Gaupp, 1896, figs 104,
105).

The inner surface of the crista is

The crista ossis ilii
Description:(vexillum of some authors) and the pars
cylindriformis can be well distinguished inmedial view, whereas only with somedifficulties in lateral view. The crista does
not reach up to the anterior end of the ala.

occupied by the m. coccygeo-iliacus thatruns to the urostyle. The iliacus externus
muscle is, together with the iliacus
internus, one of the most robust pelvic
muscles in ranids and perhaps it plays an

It is directedfact that its

to

the

dorsally in its section adjacenttuber

dorsomediallysuperius,
in

its

and

bent

anterior portion.

Consequently, it forms wide groove on the
medial surface of the ala, roofed dorsallyby the edge of the crista, and ventrally
delimited by the pars cylindriformis. The
tuber superius is prominent above the levelof the crista (but not in ZIL/EL/1 11),

however,its

margin corresponds by its

thickness to the edge of the latter. Only
posteriorly the tuber has a rounded margin,
declined rather laterally.
In specimen
ZIL/EL/68 the tuber is prominent but notextensive; it has conspicuous ridge running

down obliquely from its top. There is adistinct depression

between the tuber and

The depression
pars cylindriformis.
extends to the dorsal margin of the ala,

important role in jumping, despite of thetendon is fixed close to theproximal end of the femur. The tuber
superius serves for attachment of the m.
glutaeus magnus, m. ilio-fibularis, and m.
ilio-femoralis. The first is the most robust
muscle of the dorsal side of the thigh, andtogether with other heads of the m. triceps
femoris it stretches the knee joint. All
muscles that are inserted on the tuber
superius are important for locomotion.

Discussion:

Contemporary European

ranids mostly have the torus superior ilii

continuous with the crista ilii, regardless ifthey belong to the esculenta or temporaria

However, certain variation
exists in this respect, so one can findindividuals with prominent torus also incomplexes.

those forms in which it is continuous with

Asiatic Herpetological Research

December 1993

Vol. 5, p. 39

B
FIG. 6.

Ranidae indet. Prae sacral vertebrae in dorsal (above) and ventral (below) views.

A- V5(ZIL/EL/17). B- V2(ZIL/EL125). Bar equals 1 mm.

the crista in most individuals.

This is why

more precise assignment is difficult.

ZIL/EL/8) produce distinct proc. spinosus
which is, as usually in V2 of ranids,
directed posteriorly.

Material:

Praesacral vertebrae

V2 -ZIL/EL/8 (Fig. 5C) and ZIL/EL/125 (Fig.
6B); V3 - ZIL/EL/9; V5 ZIL/EL/17 (Fig.
- ZIL/EL/70
V6
6A);(Fig. 5B).
Description: The centra are procoelous,
In V2, thedorsoventrally compressed.
aredistinctly inclined
diapophyses
andanteriorly
slightly also ventrally; they
are oval in cross-section due to moderate
dorsoventral compression. ZIL/EL/125 issimilar in its preserved characters buthaving the postzygapophyses
more robust, and the posterior convexity of
the centrum more prominent (see Fig. 6B).
Besides that, the anterior-posterior distancediffers in

between the prae- and postzygapophyses isgreater than in ZIL/EL/8 because the former
processes extend anteriorly beyond the

The neural arches ofarenot preserved in(theylevel of the centrum.

ZIL/EL/125

Its

dorsal surface isflat, only anteriorly there is a narrow andlow ridge paralleled by a depression on
V3 is represented by ZIL/EL/9
which is preserved only as fragment

either side.

lacking the centrum, but its diapophysis
with some rugosity in the middle of its
length, as well as an extent of its neural
canal and shape of its praezygapophysis,suggest its relations to the ranids.
ZIL/EL/17 is V5; it has its proc. spinosus
directed dorsally (again, as usual in ranids).

Perhaps it might be associated with
ZIL/EL/70 (see below), judging by theshape of the centrum in ventral view (alsoin this specimen the posterior convexity is

divided by a slot, though visible only inposterior aspect). Peculiar feature of thisspecimen is the ventral edge of its anterior
concavity which runs out anteriorly in the
mid-line (see Fig. 6A). It is difficult to say
whether this is of some taxonomic

Vol.

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40

Asiatic Herpetological Research

December 1993

FIG. 7. A- cf. Bufo sp., left scapula in lateral view (ZIL/EL/140). B- Bufo sp., left scapula in lateral view
(ZIL/EL/7). C- Anura indet., left praearticular in dorsal view (ZIL/EL/12). D- Anura indet., parasphenoid(ZIL/EL/185). E-Ranidae indet., right coracoid in ventral view (ZIL/EL/54). F- Anura indet., rightpterygoid (ZIL/EL/141). Abbreviationsx. gl. cavitas glenoidalis; i. gl. intumescentiaglenoidalis; m.a.

margo anterior; p. a. pars acromialis; p. gl. - proc. glenoidalis. Bar equals 1 mm.

V6 is represented byimportance.
ZIL/EL/70. Its diapophyses are rounded incross-section, and are of the same diameterboth proximally and distally. They aredeveloped than in ZIL/EL/17. Signs ofsuch slots may be observed incorresponding vertebrae of someindividuals of the contemporary Ranidae

Thehorizontal, not inclined dorsally.
posterior convexity is clearly divided
vertically by a slot which is better

Rana esculenta). All the describedpraesacral vertebrae have in common adistinct indentation along the posterior edge
(e.g.,

Asiatic Herpetological Research

December 1993

Vol. 5, p. 41of the neural arches; this is interrupted only

The sulcus pro cart. Meckeli is, especiallyin the mid-line where a distinct ridge runsin the posterior moiety

down from the proc. spinosus.
All the above features suggest thatvertebrae of the Ranidae should beconcerned. Bufonids are excluded mainly
because of the morphology of their proc.
spinosus and because their neural canal isless spacious.

Material:

Sacral vertebra ZIL/EL/10(Fig. 5A).

Description:Thecentrumis

dorsoventrally compressed, bicondylar
posteriorly, both condyli being separated
The
by a comparatively wide notch.
anterior side of the centrum is not preserved
but comparison with contemporary ranids
suggests that there could be a mineralizedintervertebral disc.

The diapophyses are

and are distinctlyThe
compressed dorsoventrally.
of
thearticulation
surfaceinis,correspondence
praezygapophyses
inclineddorsally,

with the inclination of the diapophyses,
facing dorsomedially.

of the bone, only
moderately developed. These characters do
not allow precise assignment.
Material:

Right pterygoid, ZIL/EL/141(Fig. 7F).

Description: The inner surface (margo
orbitalis) of the ramus maxillaris does notbear any crista and the ramus itself isalmost straight. The distal section of theramus posterior is broken off, so the real
proportions of the bone are difficult toreconstruct.

Material: Ilium, ZIL/EL/3 (Fig. 3D).

Description:
Although this ilium ispreserved only as a small section, importantdiagnostic characters are preserved. The
crista ossis ilii is well developed, and may
be distinguished both in medial and lateral
In contrast to ranids, the torus
view.
superior is not developed, and the anterior
margin of the pars descendens is directed
posteroventrally instead of ventrally or evenanteroventrally (see Bohme 1977, fig. 9).

Anura indet.

Praesacral vertebra (mostMaterial:

probably V5 or V6), ZIL/EL/1 1.
Material:

Parasphenoid, ZIL/EL/185(Fig. 7D).

Description: Only the centrum and basesof the left transverse process incl. adjacent

Description: The shape and proportionsof the bone may be seen from Fig. 7D.
Among the characters that should bearepreserved.
praezygapophysis
However, one can conclude after thementioned are the proc. posterior which iswell developed, narrow proximal parts

declination of the proximal part of thetransverse process that V5 or V6 should beconcerned. The centrum is dorsoventrally(compared with the distal sections) of thecompressed and procoelous, thoughits

lateral processes and of the pars medialis,posterior side is also slightly concave.

Asand distinct indentations on the transition
between the pars medialis and lateralprocesses caused by a low ridge on eitherside; similar ridge continues on both sides
from the lateral edge of the proc. posterior
onto the surface of the proc. lateralis whereits

it disappears.

surface does not display spongious

structure (indicating a crack) it can beadmitted that there could be a freeintervertebral disc that in living animal
adhered the posterior end of the centrum.

The ventral surface of the centrum is almost
at the same level as the proximal section of
the transverse processes, and the centrum

Material: Left praearticular, ZIL/EL/12

itself isshort antero-posteriorly.

The(Fig. 7C).

praezygapophysis is comparatively small
and located far laterally (its distance from

Description: The proc. coronoideus iswell developed, nearly vertical in position.

the lateral edge of the proximal concavity of

the centrum

isabout half the diameter of

Vol.

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December 1993

-OL

D

Radioulnae in medial view. A- Bufo sp. (ZIL/EL/62). B-Bufo sp. (ZIL/EL/106). C- Bufo sp.
(ZIL/EL/120). D- Anura indet. (ZIL/EL/36). E- Anura indet. (ZIL/EL/46). F- Anura indet. (ZIL/EL/72).
D-F reversed for comparison. Arrows indecate border between periost and that part of the bone without
crista on the ulnar
capitulumradioulnae; OL. olecranon; cr.
periostal surface. Abbreviations: CAP.

Fig. 8.

margin. Line equals 1 mm.

this concavity).
The neural arches are
and the neural canalthin,comparatively

was obviously quite spacious.
Material: Urostyle, ZIL/EL/88.

This element fits
Description:
morphologically into the variation range ofcontemporary Ranidae. Both in Ranidae
and Bufonidae this range is rather broad
which precludes precise assignment of thecompressed in their columnar section. This
results in that their anterior and posterior
margins run out in a distinct ridge. The
smallest specimen (ZIL/EL/72), however,has its margins rounded. These radioulnae
might be ascribed to the Ranidae, however,lack of diagnostic characters of theseelements casts some doubts on thisassignment.

Accompanying Vertebrate Fauna in the
Samplespecimen.
Material:

character is that most of them are laterally

Radioulnae ZIL/EL/36 (Fig.

medium to small sized (compared with

From Kudaro I Cave, Tsepkin (1980)
gave a list of accompanying fishes,
Darevsky (1980) mentioned one lizard
genus (Lacerta sp.), Burchak-Abramovich(1980) gave a list of birds, Gadzhiev

those identified as Bufo).

(1980) bats, Gromov

8D), ZIL/EL/46 (Fig. 8E), ZIL/EL/72 (Fig.
8F), ZIL/EL/73, ZIL/EL/1 14, ZIL/EL/152.

Description:

These radioulnae are

A conspicuous& Fokanov (1980)

Asiatic Herpetological Research

December 1993&

References

rodents, and Vereshchagin
Baryshnikov(1980) large mammals. In the sample thatwas placed at my disposal

there were,
besides frogs, also two different forms of
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Ivan
Horacek,(determination by

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ofZoology,CharlesBANNIKOV, A. G.,

I.

S.

DAREVSKY, V. G.

ISHCHENKO, A. K. RUSTAMOV, AND N. N.
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BOHME, G.

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BOLKAY, S.

Determination of the material revealed

Vol. 5, p. 43

1977. Zur Bestimmung quarterer
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Bosni i Hercegovini, (1919):277-357.

that its substantial part belongs to the genus

Bufo and to the family Ranidae. Minor part
(represented by fragmentary or lessnumerous elements) could be determined
only as Anura indet. Precise determination
was impossible because of supposed
individual and developmental variation.
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generic composition of the
Holocene anuran fauna in this region was

show

that

basically the same as contemporary one.
The genus Bufo in the corresponding
altitudes of Caucasus is nowadays

represented by. Bufo verrucosissimus, andB. viridis, genus Rana by R.

macrocnemis,
R.
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et
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am grateful to Professor I. S. Darevsky(Zoological Institute, St. Petersberg) for theloan of the fossil material for study, and to
Dr. I. Horacek (Department of Zoology,
Charles University, Prague) for theofdeterminationaccompanying
micromammalian fauna. Thanks are due
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