Asiatic Herpetological Research

1997

Vol. 7, pp. 6-18

A Catalogue of Non-Metrical Variations in Skull Bones of Vipera lebetina(Reptilia, Viperidae)

Yuri A. Chikin
Institute of Zoology, Uzbek Academy of Sciences, A. Niyazov Street, I,

Tashkent 700095, Uzbekistan.

Abstract.- This paper presents a description, codes, and figures of 16 variations of non-metrical characters of 122
skull bones of Vipera (Macrovipera) lebetina inhabiting Central Asia and the Caucasus. Statistical characteristicsof the number of teeth on the pterigoideum and dental bones for several populations of the three subspecies of thelevantine viper are provided.

Kev words:

Reptilia, Ophidia, Viperidae. Vipera lebetina obtitsa,
Turkmenistan, Caucasus, osteology, skull bones.

VV

turanica,I.

I.

cernovi, Uzbekistan,Introduction

magnifier. The characters are arranged in groups to beCurrently, the use of non-metrical variation in cranialpresented in the tables: the characters of the bony eleas foramenal characters in Table 2.
ments in Table
1

structures for the identification of differences and thedetermination of whether an individual belongs tosome populational group, subspecies or larger taxon,widely usedin zoological studies (Berri. 1975;
Shubinand Sedokova, 1982; Larina
1978:
Yaletsky,and Eremina. 1988: Zerova and Chikin. 1992). Whileis

working on a catalogue of non-metrical variations of
thecranialtraitsof Vipera

lebetinaL..

attempted to identify the range of variability,

1758,

I

system-Table 3 provides statistical characteristics, calculatedusing standard techniques (Lakin. 1990). of the numbers of the teeth in the pterigoideum and dental bones
separately for males and females, and their total: thetotalnumber of teeth is given in Table 4. The popula-tions of V. (M.) I. turanicatau (G)

atize the data, and unify techniques for distinguishing,—are characterized in Tables 3 and 4.

described by Gurtovoy(An osteological collection in the Institute of Zoologyof the Uzbek Academy of Sciences as well asal.

(1978),

Groombridge1984). The names of the characters are coded by theletter symbols (initial letters of their Latin names).

somefrom Moscow State University were the basis

Results

A total of 268 skulls of the Levantine

viper of different ages and sexes from various localities in both Central Asia and Azerbaistan were used.

These populations are attributed by modern taxonomists to the following subspecies:

The following non-metrical variations are describedin this catalogue:(Table1)most cranial bones, (Tablevariability in the shape of2)

foramenal characters,(Table 3) the number of teeth (and whether a subspecies shows variations in this or that trait), and (Table

Vipera (Macrovipera) lebetina obtusa Dwigubsky.
1832, 35 specimens;
1.

2. V.

(M.) I. turanica Chernov. 1940. 175 specimens;

3. V.

(M.) I. cernovi Chikin and Szczerbak, 1992. 58

specimens.

The ordinal numbers of the subspecies correspondand 2 of thisto those in the fourth column of Tables
catalogue and in the first column of Tables 3 and 4.
1

Variations of the characters were revealed andillustrated while comparing the bones using the binoc-ularet(1980), Mbrkevitch and Tatarko (1983). and Szyndlar

Material and Methods

for this work.

the eastern andNames of bones and bony elements are given as

describing, and coding traits and their variations.

skulls—one fromwestern parts of the Nuratau Ridges (EN, WN) andTurkestan (T), the Malguzar Ridge (M), and Gobdun-

MBI-9 at different magnifications and with a4) some schematic drawings (Figs. 1-16), illustrating
the variations described.

Some of the variations may be considered as insignificant

anomalies (rare phenes). which characterize

only individual populations of a subspecies. However,i.e.. found in each of thethe variations are common,subspecies, and are "normal" for a species, thougheach populationalgroup. Naturally-occurring heterogeneity of bilateral
theiroccurrence

isdifferentinstructures is not reflected in the catalogue except thecharacters Mcp and Fpa on the parietal bone. These

1997

Asiatic Herpetological Researchcombinations, assessed by means of the values offluctuating asymmetry and observed even on a single(nonpaired) bone, are independent characteristics ofstable development and its description is not supposed

here.

Vol. 7.

DiscussionThispublication of the Catalogue is a slightly
extended variant of my previous work (Chikin, 1993),

which, due to its limited distribution, is inaccessible
to most scientists.

Itisnoteworthy

that

separate variations (Mpa.

Mcp). expanding the range of variability, can befact considered as the age characteristics and itissive and should be treated as a scheme, which, when

Acknowledgments
The author is indebted to Galina A. Zerova for discussions resulting in this work.

Museum of Moscow Statecal

the following:

specimensfromKhodzhaev

fordescription of variability of the other bones;(2) incorporation of new traits and variations; and(3) identification ofindividuals

known traits and variations tovarious populational and taxonomicingroupings, both living and fossil.

AppendixIFigures 1-16J^tpbFbv

My thanks are extended

to Valentina F. Orlova and Evgeny Dunayev (Zoologi-

used by researchers, will enable them to amplify it in)

my readers forin

This catalogue cannot be considered comprehen-1

will be grateful tothiscase that necessitates a study on transformations of
the characters in ontogeny.

(Itheir remarks and glad to have followers.

theirtranslation

Kraig Adler for reviewing it.

University) for thecollection.

of theDzhavkhar

manuscriptand

Asiatic Hevpetological ResearchVol. 7, p. 8

1997FedFed

MepmFigure 2. Supraoccipital bone (dorsal view). Location of the characters and their variations.

Figure 3. Parietal bone. A,B. and C are the variationsof the form of the parietal

crest and posterior margin. D, E, and F are theforamina and notches orthe anterior edge of thedescending part of thebone.

Figure 4. Postfrontal
bone, showing the surface
that joins with the parietalbone.

1997

Asiatic Herpetological ResearchVol. 7. p. 9

Figure 5. Basiparasphenoid bone.
Location of the characters: A - inferior;

D-superior views; B, C. E, Fvariations.

•iHLIzJi

7]

MtppiiMppn^-Vol. 7. p. 10

Figure 9. Maxillary bone. Adorsal views.
lateral, CMoml

1997

Asiatic Herpetological Research-anterior, B

1

Figure 10. Articular bone, a Proximal partfrom the external lateral view.

Meoa 1

Asiatic Herpetological Research

1997

F:2HS

Vol. 7, p.

1

1

Figure 15. Palatal bone. Variations of the foramina at
the dorsal part of the Internal lateral surface.

Figure 16. Squamosum bone. A variant with the fora-men.

Appendix

IITables 1-4

TABLE1.

Form of the skull bones in the Levantine Viper.

Character

Description of variations

CodeSubspecies

The main occipital bone

— basioccip-itale (Fig. 1)

Margo terminationis processus hasio-

Coral -shaped

Mtpb

Saw-shaped

Mtpb 2

1ccipitalis (Mtpb)

V-shaped
Projection in the middle

Truncated

Roundish

Margo prominentiae condyli occitocondilaris, pars dorsalis (Mpcod)

Margin of prominence sharply extending
above the bone
Margin of prominence gently extending
above the bone

Margo sinus prominentiae condili
on iti h ondilaris Mspco)

Groove absent(Groove chink shaped
Groove oval
Groove irregular
Supraoccipital bone—exoccipitale(Fig. 2)

Fades exoccipitalis, pars dorsalis

Surface is smooth or slightly wavy

Fed

Surface with hollows

Fed 21(Fed)Asiatic Herpetological ResearchVol. 7. p. 12

Character

Description of variations

1997

CodeSubspecies

Surface with deep grooves

Mai go exoccipitalis posterior, pars
media/is(Mepm

Margin salient)

Margin concave
Margin straight—Parietal boneparietale (Fig. 3)

Margo parietal is anterior (Mpa)

Margin straightMargin slightly concave
Crest absent

Margo crista parietalis (Mcp)

Crest reaches the posterior margin of thebone
Crest does not reach the posterior margin ofthe bone

Left and right parts of the crest join at acuteangles
Parts of the crest join at right angles

Parts of the crest do not join

Margo parietalis posterior Mpp()

Margin roundish

Margin right-angled
Margin obtuse-angled—

Postfrontal bone

postfrontale(Fig. 4)

Sunn a articuli ossis postfrontale

Suture starts at the projection in midpart(Suap)— basiparas-

Basiparasphenoid bonephenoid (Fig. 5)

Margo cristae basiparasphenoidis

Crest long with rounded top(Mcb)

Margo processus suborbitalis et areae
lateralis (Mpsal)

Suborbital appendix is big. in the form of asemicircle projecting beyond the line andthe margin of the lateral surface passes near
the line in the form of a concave arc

Suborbital appendix is small, projecting
beyond the line in the form of a small sectorof a circle or obtuse angle, and the marginof the lateral line parallel with the line or onit

Suborbital appendix small, parasphenoidand suborbital appendices project beyond
the line, as well as most of the lateral line

Fed 3

[997Character

Asiatic Herpetological Research

Description of variationsVol. 7. p. 13

CodeAsiatic Herpetological ResearchVol. 7. p. 14

Character

Description of variations

1997

CodeSubspecies

Sinus el crista transversus dorm salts,

1:2:3

Opening rectangular; crest well developed

Scti

The medial process is significantly larger;
the lateral process is small and smoothGpml

1I; 2;

Margin even (no projection observed)

Meoa

12; 3

pars interior (Scti)
Articular bone—articulare(Fig. 10)

Gracilis evolutionis processes media-ns et processus lateralis (Gpml—angulare

Angular bone)(Fig. 11)

Margo exiguus ossis angulares(Meoa)

Margin with process below—

Splenial bonespleniale (Fig. 12)

Margo emarginaturae ossis spleniale

Notch not restricted by the projections(Meos)
Notch posteriorly restricted by the projection

Notch anteriorly restricted by the projections

Notch both anteriorly and posteriorlyrestricted by the projections
Dental bone—dentate(Fig. 13)

Magiutudo relativa emarginaturae

Margin of notch reaches 7th toothossis articulare. pars lateralis exterior(Mrel)

Margin reaches 8th toothMargin reaches 9th toothMargin reaches 10th toothInitialio apertionis fissurae Mekkelii

Margin reaches

1

1th toothMargin reaches

1

2th tooth

Fissure opens under 7th tooth(IafM)
Fissure opens under 8th tooth
Fissure opens under 9th tooth

Mrel

13]

Asiatic Herpetological Research

997

Vol. 7. p. 15

TABLE 2. Foramenal traits of the skull bones in Levantine Viper.
Character

Description of \ariations

CodeSubspecies

The main occipital bonebasioccipitale (Fig. 1)

Foramen ossis basioccipitale, part

Foramen not present

Fbv 1

2; 3

One foramen

Fbv 2

1

Two foramina

Fbv 3

1:2

Three foramina

Fbv 4

Four foramina

Fbv 5One foramen

Fpl

Two foramina

Fpl 2ventrale (Fbv)

Prooticum bone-prooticum (Fig. 14)

Foramen ossis prooticum, pars lateralis1(Fpl)

Three foramina
Four foramina
Parietal bone-parietale Fig. 3)(Foramens et emarginaturas ossis pari-Two completely closed foramina

Fpa

One of the foramina is not closed

Fpa 2

Two open foramina, rupture in the wall

Fpa 3

1etale, pars anterior (Fpa)smaller than the diameter of the foramen

Two open foramina, rupture in the wall larger

Fpa 4than or equal to the diameter of the foramen

Of equal size on both sides

Fpa 5

Both foramen and sulcus missing

Fspt 1

Foramen present

Fspt 2

Sulcus present

Fspt 3

Both a groove and sulcus present

Fspt 4

One fossa

Facv 1

External edge of foramen not closed (partition
has sulcus, canal)

Facv 2

Basiparasphenoid bone-basiparasphenoid (Fig. 5)

Foramen et sulculus processes trabecularis (Fspt)

Foramen anterior canal is vidianis(Facv)

Premaxillary bone-premavilla (Fig. 6)

1

:2; 3

Vol. 7, p. 16

Character

Asiatic Herpetological Research

Description of variations

1997

CodeSubspecies

Foramen ossispremaxilla. pars ventralis

Foramen not present

Fpv

One foramen

Fpv 22; 3

Two foramina

Fpv 32: 3

Three foramina

Fpv 4

2: 3

Four foramina

Fpv 5Foramen not present

Fpli

One foramen

Fpli 21(Fpv)

Palatine bone-palatinum (Fig. 15)

Foramen ossis palatinum, pars lateralis

1interior (Fpli)

Two foramina
Three foramina

Squamose bone squamosum (Fig. 16)
Foramen squamosum (Fsq)

No foramen
One foramen

Angular bone-angulare (Fig. 11)

Foramen annulare (Fa)

No foramen
One foramen

Two foramina
Splenial bone spleniale (Fig. 12)

Foramen spleniale (Fsp)

No foramen
One foramen

Dental bone-dentale (F'ig.13)

Foramen mentale (date, fornix Fm)

One foramen, oval
One half-moon-shaped foramen

Two oval foramina
Situs foramen mentale (Sifm)

One foramen under 5th tooth
One foramen under 6th tooth
One foramen under 7th tooth
One foramen under 8th toothTwo foramina under 4th and 7th teeth
Two under 5th and 6th teeth
Two under 5th and 7th teeth

I;

2

1997

Character

Asiatic Herpetological Research

Description of variations

Vol. 7, p. 17

CodeSubspecies

TABLE 3.

Two under 6th and 7th teeth

Sifm 82

Two under 6th and 8th teeth

Sifm 92

Two under 7th and Xth teeth

Sifm 10

2:1

Asiatic Herpetological ResearchVol. 7, p. 18

TABLE 4. Total tooth number on the skull in the Levantine Viper.
BoneSubspecies:

1997