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L Si

ILLh.ou. LIBRARY
AT URBANA-CHAMPAIGN

FIELDIANA: ZOOLOGY

A Continuation of the
ZOOLOGICAL SERIES

of
FIELD MUSEUM OF NATURAL HISTORY

VOLUME 65

NATURAL HISTORY SURVEY

DEC 6 1976

LIBRARY

FIELD MUSEUM OF NATURAL HISTORY
CHICAGO, U.S. A.

FIELDIANA
Zoology

Published by Field Museum of Natural History

Volume 65, No. 4 June 13, 1974

Preliminary Key to the Turtles, Lizards,
and Amphisbaenians of Iran

Steven C. Anderson

Callison College, University of the Pacific
Stockton. California

INTRODUCTION

Research toward a monographic treatment of the lizards, turtles, and
amphisbaenians of Iran has been completed recently. Preparation of the
final manuscript has been delayed, and a further delay in publication
seems inevitable. For this reason, it seems desirable to publish the key to
this fauna in a preliminary form as an aid to collectors and others con-
cerned with the fauna of Iran and Southwest Asia generally. It is hoped
that through use, its shortcomings will be revealed and communicated
to the author, so that an improved, illustrated version can be included
in the monograph.

To facilitate use of the key and to aid in recognition of significant
range extensions, the distribution of turtles, lizards, and amphisbaenians
is shown according to political divisions (ostans) in Table 1. The distri-
bution according to natural geographic regions has been discussed in
detail elsewhere (Anderson, 1968).

Non-herpetologists using this key are referred to Peters (1964) for
definitions of unfamiliar terms. Species preceded by an asterisk (*) have
not yet been recorded definitely from Iran. Certain difficulties attend
the use of a key not accompanied by illustrations, diagnoses, and de-
scriptions of each species, and individual specimens, especially juvenile
and damaged specimens may not be identifiable on the basis of the key
alone. This is particularly true in the case of geckos, in which loss of the

US ISSN 0015-0754

Library of Congress Catalog Card Number: 74-77215

Pubucauon „86 27 NATURAL HISTORY SURVEY

LIBRARY

FIELDIANA: ZOOLOGY, VOLUME 65

SAUDI ARABIA

Fig. I. Map of Iran showing the primary political divisions (ostans). 1. Tehran, 2.
Gilan, 3. East Azarbaijan, 4. West Azarbaijan, 5. Kordestan-Kermanshah, 6. Khuzestan-
Lorestan, 7. Esfahan, 8. Fars, 9. Kerman, 10. Baluchestan-Sistan, 11. Khorasan, 12. Maz-
andaran. Boundaries and ostan numbers after Fisher, 1968, p. 4. Redrawn).

tail is frequent; in some cases it has been necessary to rely on caudal
characters in the key. This is true in the case of certain species which I
have not examined, and have had to base the dichotomies on descrip-
tions in the literature, and in other instances where statistical criteria
are the only definitive means of separating taxa other than the char-
acters 1 have used in the key. If the locality is known for a specimen, the
distribution table (table 1) should aid in a tentative identification.

ACKNOWLEDGEMENTS

Thanks are owed a great many people, and I will enumerate my debts
to these people in the monograph. Two groups of people must be men-
tioned here, however: the collectors and the museum curators, without
whom all work in biosystematics would be impossible. Collectors of
material (from all countries in Southwest Asia) that I have examined in
preparation of this key include: Jeromie A. Anderson, William T. Blan-
ford. Erica and Richard Clark, Anthony F. De Blase, Henry Field, E. S.

ANDERSON: KEY TO REPTILE GROUPS 29

Fraser, John Gasperetti, Jerry Hassinger, Harry Hoogstraal, Walter P.
Kennedy, Douglas Lay, H. Loftier, Yusuf Lazar, C. W. McEwan, Richard
A. Martin, R. P. Miller, Sherman A. Minton, Jr., John W. Neal, Jr., Knud
Paludan, R. W. Redding, Charles A. Reed, A. R. M. Rickards, Janice K.
and William S. Street, Howart Stutz, Robert G. Tuck, Jr., Dan Womochel,
and N. A. Zarudny.

Mr. and Mrs. William S. Street deserve special mention in this list,
as their two Iranian expeditions recently have helped to assemble the
most comprehensive collections in U. S. museums. Their collections,
deposited in Field Museum of Natural History, along with my own ear-
lier collection from southwestern Iran (deposited in the California Acad-
emy of Sciences) have formed the basis of my studies of the amphibians
and reptiles of Iran. Street Expedition material from Afghanistan has
also provided comparative data.

Also deserving of special mention are the anthropological expeditions
of Henry Field. Dr. Field has maintained an active concern with promot-
ing knowledge of the fauna of Southwest Asia, and specimens collected
by his expeditions and by other collectors at his request have formed the
foundation of U. S. herpetological collections from that area over the
past 45 years. Most of this material is in Field Museum of Natural History
and at the Museum of Comparative Zoology, Harvard.

Curators who have loaned me material and /or made me welcome at
their institutions during the past 15 years include: Alan E. Leviton, Cali-
fornia Academy of Sciences; Robert Inger and Hymen Marx, Field
Museum of Natural History; James A. Peters, United States National
Museum; Richard Zweifel and Charles Myers, American Museum of
Natural History; Charles Walker, Donald Tinkle, and Arnold Kluge,
Museum of Zoology, University of Michigan; Ernest E. Williams, Muse-
um of Comparative Zoology, Harvard; Robert C. Stebbins, Museum of
Vertebrate Zoology, University of California, Berkeley; Ilya Darevsky,
Zoologicheski'i Institut, Leningrad; Josef Eiselt, Naturhistorisches Mu-
seum, Wien; Alice Grandison, J. C. Battersby, and E. N. Arnold, British
Museum (Natural History); Jean Guibe; Muse6 Nationale d'Histoire
Naturelle, Paris; F. W. Braestrup, Universitetes Zoologiske Museum,
Copenhagen.

Much of the research leading to the development of this key was
done while I was Associate Curator in the Department of Herpetology,
California Academy of Sciences.

I thank Hymen Marx, Field Museum of Natural History, for com-
ments and suggestions.

30 FIELDIANA: ZOOLOGY. VOLUME 65

The work was supported in part by a grant from The American Phil-
osophical Society (Grant No. 4959 - Penrose Fund, 1968) and by two
Faculty Research Grants (1971 and 1972) from the University of the
Pacific.

KEY TO THE TURTLES OF IRAN1

la. Carapace without homy plates; feet with 3 claws Trionyx euphraticus (Daudin 1802)

lb. Carapace with horny plates; feet with 4 or 5 claws 2

2a. Head covered with undivided smooth skin; digits fully webbed 3

2b. Head covered by shields; digits not webbed 4

3a. Plastron united to carapace by bony suture; plastron not hinged, immovable; anal
plates of plastron pointed, their median suture shorter than interabdominal suture

Mauremys caspica caspica (Gmelin 1774)
3b. Plastron united to carapace by ligamentous attachment; plastron more or less dis-
tinctly hinged, movable (in adults); anals rounded, their median suture longer than

interabdominal suture Emys orbicularis (Linnaeus 1758)

4a. Forelimb with 4 claws Testudo horsfieldii Gray 1844

4b. Forelimb with 5 claws 5

5a. Shell oval in outline, with smooth, rounded posterior margin; ground color light olive,
with large, distinct, individual dark markings . . . Testudo graeca ibera Pallas 1814
5b. Shell elongate in outline, with upturned, emarginate posterior margin; ground color
brownish olive, with very indistinct dark markings

Testudo graeca zarudnyi Nikolsky 1896

KEY TO THE LIZARDS AND AMPHISBAENIANS OF IRAN

I a. Limbs absent 2

I b. Limbs present 5

2a. Eyelids well developed and movable; osteoderms underlie scales of head and body.

ANGU1DAE 3

2b. No movable eyelids; no osteoderms underlie scales of head and body 4

3a. A deep lateral fold from head to level of vent; teeth blunt, with conical crowns

Ophisaurus apodus (Pallas 1775)

3b. No lateral fold; teeth long and sharp Anguis fragilis colchicus (Nordmann 1840)

4a. Body ringed with distinct annuli; eyes very small, beneath head shields; scales not

imbricate Diplometopon zarudnyi Nikolsky 1907

4b. Body not ringed with distinct annuli; eyes usually large, well developed, with distinct

iris and pupil, sometimes small (Typhlopidae and Leptotyphlopidae); scales imbricate

snakes (not covered in this work)
5a. Skin soft, with granules, rarely imbricate scales; no paired, symmetrically arranged

shields on top of head, which is covered by granules; neither suborbital nor fron-

tosquamosal arch present on skull; clavicles broadened, forming loop at inner end;

tongue smooth or covered by thread-like papillae; pupil of eye usually vertically

elliptical (except in Pristurus). GEKKONIDAE 6

Marine turtles of the Persian Gulf are not included.

ANDERSON: KEY TO REPTILE GROUPS 31

5b. Skin never soft, composed of scales, plates, or granules; either suborbital and/or fron-
tosquamosal arch present on skull; clavicles not broadened on inner end, or if broad-
ened, then tongue covered by imbricate, scale-like papillae or by oblique folds . 44

6a. Eyelids movable; digits not dilated; procoelous vertebrae 7

6b. Eyelids immovable (spectacle); digits dilated or not; amphicoelous vertebrae 8

7a. Subdigital lamellae smooth . . .Eublepharis angramainyu Anderson and Leviton 1966
7b. Subdigital lamallae each with several small tubercles

Eublepharis macularius (Blyth 1854)

8a. Pupil of eye round Pristurus rupestris Bland lord 1874

8b. Pupil of eye vertically elliptical 9

9a. Digits strongly dilated 10

9b. Digits not dilated 17

10a. Each digit dilated at base, with double row of lamellae beneath, forming pads; ter-
minal phalanges conpressed II

10b. Each digit dilated at apex, terminating in subtriangular expansion, claw lying in

longitudinal groove dividing apical expansion 15

I la. Tail with sharp, denticulated lateral edge; outer postmentals not in contact with

labials Hemidactylus garnotii Dum^ril and Bibron 1836

lib. Tail without sharp, denticulated lateral edge (although in H. flaviviridis there is a
ventrolateral row of small pointed tubercles); outer postmentals in contact with

labials 12

12a. No enlarged dorsal tubercles, or if tubercles present, these are rounded, feebly keeled,
not regularly arranged (none present in Iranian, Afghan, Pakistan, or northern
Indian specimens examined); males with femoral pores only

Hemidactylus flaviviridis Rilppell 1835

1 2b. Enlarged dorsal tubercles numerous, strongly keeled, arranged in more or less regular

longitudinal series; males with preanal pores only, or with both preanal and femoral

pores 13

13a. Males with 15-27 femoral and preanal pores; 6-10 lamellae under 4th toe

* Hemidactylus brookii Gray 1845
13b. Males with preanal pores only; 8-14 lamellae under 4th toe 14

14a. 8-11 lamellae and pairs of lamellae under basal expanded portion of 4th toe; 7-10
supralabials and 7-9 infralabials; males with 2-10 preanal pores.

Hemidactylus turcicus turcicus (Linnaeus 1758)
14b. 12-14 lamellae and pairs of lamellae under basal expanded portion of 4th toe; 10-12
supralabials and 8-10 infralabials; males with 9-13 preanal pores

Hemidactylus persicus Anderson 1872
15a. Apical expansion of digit with fine lamellae beneath; postanal sacs present.

Ptyodactylus hasselquistii (Donndorff 1789)
15b. Apical expansion of digit smooth beneath (low magnification); postanal sacs absent

16a. Largest dorsal turbercles more than one-half height of ear opening; tubercles extend-
ing onto occiput and temporal area, much larger than surrounding granules; whorls
of caudal tubercles separated by 3-4 transverse rows of small scales

Asaccus elisae (Werner 1895)

16b. Largest dorsal tubercles less than one-half height of ear opening; tubercles becoming

much smaller on nape, usually not extending onto head, or if so, few in number,

scarcely larger than surrounding granules; whorls of caudal tubercles separated by

5-6 transverse rows of small scales

Asaccus griseortotus Dixon and Anderson 1973

32 F1ELDIANA: ZOOLOGY. VOLUME 65

17a. Digits with well-defined lateral fringe of elongated, flexible pointed scales 18

1 7b. Digits without lateral fringe of elongate, flexible pointed scales, although scales may

be denticulate 23

18a. Dorsal scales intermixed with larger rounded tubercles

Crossobamon eversmanni (Wiegmann 1834)

18b. Dorsal scales uniform, not intermixed with tubercles 19

19a. Dorsal scales small, not cycloid; scales of tail not large, not plate-like, and not strongly

imbricate 20

19b. Dorsal scales large, cycloid; tail covered above (at least on posterior two-thirds) by

single row of large, plate-like, strongly imbricate scales 21

20a. Back with 4 dark crescentic crossbars; 10-11 supralabials; forelimb does not reach

beyond tip of snout Stenodact ylus affinis (Murray 1884)

20b. No dark crossbars on back; 12-15 supralabials; forelimb reaches beyond tip of snout

Stenodact ylus doriae (Blanford 1874)
21a. Large cycloid scales of dorsum extend forward to occiput

Teratoscincus scincus (Schlegel 1858)

21b. Large cycloid scales not extending forward beyond shoulders 22

22a. Not more than 60 scales round middle of body Teratoscincus bedriagai Nikolsky 1899
22b. About 100 scales round middle of body . . . .Teratoscincus microlepis Nikolsky 1899

23a. Dorsal scales uniform, small, homogeneous 24

23b. Dorsal scales heterogeneous 27

24a. No postmentals (chin shields) Tropiocoiotes latifi Leviton and Anderson 1972

24b. Postmentals present 25

25a. A single pair of postmentals, not in contact; dark crossbars of body absent or indis-
tinct, sometimes two dorsolateral series of spots

Tropiocoiotes helenae (Nikolsky I907)1

25b. Two pairs of postmental shields; dark crossbars of body and tail distinct 26

26a. Dark dorsal crossbars of body and tail broader than interspaces

Tropiocoiotes persicus bakhtiari Minton, Anderson, and Anderson 1970
26b. Dark dorsal crossbars less than one-half width of interspaces

Tropiocoiotes persicus persicus (Nikolsky 1903)

27a. Dorsal scales of many sizes, all scales except labials and chin shields strongly keeled

Tropiocoiotes heteropholis Minton, Anderson, and Anderson 1970

27b. Dorsal scales small, intermixed with larger tubercles; at least some scales of head and

body smooth (except Bunopus aspratilis) 28

28a. Subdigital lamellae with a single transverse series of tubercles, particularly on the free
margin, seen under magnification (sometimes worn down in later part of epidermal

cycle); distal phalanges not compressed 29

28b. Subdigital lamellae smooth; distal phalanges compressed or not 31

29a. Postmentals (chin shields) absent Bunopus tuberculatus Blanford 1874

29b. Postmental shields present 30

30a. Ventrals strongly keeled; tail with large, strongly keeled, sharply pointed tubercles,

no subcaudal plates Bunopus aspratilis Anderson 1973

30b. Ventrals smooth; tail without enlarged tubercles, posterior three-fourths with en-
larged subcaudal plates Bunopus crasstcauda Nikolsky 1907

1 Schmidtlcr and Schmidtler (1972) have described a new subspecies, Tropiocoiotes
helenae fasciatus. from Kordcstan-Kermanshah and Khu/.estan-Lorestan Provinces. The
two subspecies are distinguished as follows: T. h. helenae— 65-84 dorsal scales between
axilla and groin. 0-6 indistinct dark dorsal crossbars with white posterior margins; T h.
fasciatus 80-92 dorsal scales. 5 distinct crossbars with white posterior margins.

ANDERSON: KEY TO REPTILE GROUPS 33

31a. Postmentals (chin shields) present, and well differentiated in size and shape from
granular small scales of chin and throat 32

31b. Postmental shields absent (sometimes a short row of enlarged, subcircular scales
present behind mental) 43

32a. Subfemoral tubercles present among granules of lower surface of thigh, in short row of
2-6, often in contact with posterior row of large imbricate scales; males with con-
tinuous series of preanal and femoral pores 33

32b. No subfemoral tubercles; males with preanal pores only 34'

33a. 24-29 strongly keeled, nonmucronate trihedral or subtrihedral tubercles in paraverte-
bral row from occiput to level of vent; males with 28-41 (32-40 in Afghan specimens
examined) preanal and femoral pores (total of both sides).

Cyrtodactylus fedtschenkoi (Strauch 1887)

33b. 19-23 strongly keeled, mucronate tubercles in paravertebral row from occiput to
level of vent; males with 23-31 (24-29 in Afghan and Iranian specimens examined)
preanal and femoral pores (total of both sides)

Cyrtodactylus caspius (Eichwald 1831)

34a. Subcaudal scales one head-width behind vent small, not enlarged and plate-like . 35

34b. Subcaudal scales one head-width behind vent enlarged, plate-like, 2 serially arranged
plates, or pairs of plates covering each caudal segment 38

35a. Subcaudal plates smooth 36

35b. Subcaudal plates distinctly keeled 37

36a. Scattered small keeled tubercles among the large trihedral dorsal tubercles which
form fairly regular longitudinal rows; tubercles on tail arranged around middle of
each segment, not in terminal scale row . . . .Cyrtodactylus russowii (Strauch 1887)

36b. No scattered small tubercles among the rows of enlarged dorsal tubercles; caudal
tubercles form terminal rings of each annulus

Cyrtodactylus kachhensis (Stoliczka 1872)

37a. 23-30 abdominal scales across middle of belly (about 1 1 scales in a distance across
belly equal to length of snout)

Cyrtodactylus heterocercus heterocercus (Blanford 1874)

37b. 14-16 abdominal scales across middle of belly (less than 10 scales in a distance across
belly equal to length of snout) Cyrtodactylus saggitifer (Nikolsky 1899)

38a. Subcaudal plates in 2 median series; dorsal tubercles distinctly smaller than inter-
spaces; snout 2 to 2'/i times longer than diameter of eye

Cyrtodactylus kirmanensis (Nikolsky 1899)

38b. Subcaudal plates in a single median series; dorsal tubercles smaller or larger than
interspaces; snout length less than twice diameter of eye 39

39a. Caudal tubercles arranged around middle of each caudal segment, not forming termi-
nal ring of each segment * Cyrtodactylus kotschyi (Steindachner 1870)

39b. Caudal tubercles (or enlarged keeled scales) forming terminal ring of each segment
40

40a. Dorsal tubercles distinctly smaller than interspaces, rounded, smooth or weakly keeled
to subcorneal, but not distinctly trihedral; peritoneum and investiture of some in-
ternal organs of abdominal cavity darkly pigmented; limbs and tail thin, attenuate.41

1 DeWitte (1973) has described a new genus and species, Rhinogekko misonnei, from
the Dasht-e Lut (Kerman, Baluchistan-Sistan, and Khorasan Provinces). It would appear
at this point in the key, and is distinguished from all other species in having the nostril
situated at the apex of a prominent caruncle composed of four scales. It is closely related
(if not identical) to " Agamura" femoralis Smith 1933, which is known from Baluchistan,
Pakistan.

34 F1ELDIANA: ZOOLOGY. VOLUME 65

40b. Dorsal tubercles distinctly larger than interspaces, strongly keeled and trihedral,
peritoneum and investiture of organs of abdominal cavity without melanocytes;
limbs and tail sturdy 42

41a. 24-28 abdominal scales across middle of belly (14- 15 scales across belly in distance
equal to length of snout); snout length less than l'/$ times diameter of eye

Cyrtodactylus agamuroides (Nikolsky 1899)

41b. 10-16 abdominal scales across middle of belly (6-8 scales across belly in distance
equal to length of snout); snout length I1/: times diameter of eye

Cyrtodactylus gastropholis (Werner 1917)

42a. 12-16 dorsal tubercles in longest transverse (chevron-shaped) series across back;
width of dorsal tubercles distinctly smaller than greatest diameter of ear opening;
10-14 supralabials Cyrtodactylus scaber (Heyden 1827)

42b. 10 dorsal tubercles in longest transverse series across back; width of dorsal tubercles
nearly equal to greatest diameter of ear opening; 9 supralabials

Cyrtodactylus brevipes (Blanford 1874)

43a. Tail cylindrical, very slender, and of almost uniform diameter from base to tip (tip
blunt), no mucronate tubercles on annuli; distal phalanges of digits compressed,
narrower than basal phalanges and strongly angularly bent

Agamura persica (Dumeril 1856)

43b. Tail tapering gradually (tip of original tail sharp), 2 mucronate tubercles on either
side of each annul us; digits cylindrical, not strongly angularly bent

Alsophylax spinicauda Strauch 1887

44a. No paired, symmetrically arranged shields on top of head, which is covered by granules,
small scales, or tubercles 45

44b. Enlarged, paired symmetrical plates on top of head (some granules may be present,
but large shields predominate) 70

45a. Venter covered by small juxtaposed granules or quadrangular scales; tongue deeply
divided, long and slender, smooth, retractile into sheath at base; dorsum covered
with numerous small juxtaposed granules or scales; dentition pleurodont.
VARANIDAE 46

45b. Venter covered by imbricate scales, not granules; tongue broad and short, smooth or
covered with villose papillae, not deeply forked; dorsum covered by imbricate scales
or a combination of imbricate scales and granules; dentition primarily acrodont.
AGAMIDAE 48

46a. Tail compressed throughout its length, with low, double-toothed crest above; abdom-
inal scales in 88-1 10 transverse series from collar fold to groin

Varanus bengalensis bengalensis (Daudin 1802)

46b. Tail round in cross-section, or slightly compressed posteriorly, without double-toothed
crest above; abdominal scales in 110-125 transverse series from collar fold to
groin 47

47a. Tail round in cross-section throughout its length; back with 5-8 (usually 6) gray bars
in addition to 1-2 nuchal crossbars, pattern becoming indistinct in older animals,
pattern of dots predominating; tail patterned nearly to tip with 19-28 dark cross-
bars Varanus griseus griseus (Daudin 1 803)

47b. Posterior half of tail narrow in cross-section, compressed, distinct keel above; back
with 5-8 (usually 6) sepia bars in addition to nuchal crossbar; tail with 13-19 dark
crossbars, end of tail without pattern .... Varanus griseus caspius (Eichwald 1841)

48a. Tympanum concealed or absent 49

48b. Tympanum exposed 57

49a. Large fringed cutaneous fold at angle of mouth

Phrynocephalus mystaceus galli Krassowsky 1932

ANDERSON: KEY TO REPTILE GROUPS 35

49b. No cutaneous fold at angle of mouth 50

50a. Dorsal scales heterogeneous, small scales intermixed with strongly enlarged scales
51

50b. Dorsal scales subequal, homogeneous 54

51a. Enlarged dorsal scales flat, not tubercular, posterior border not sharply upturned;
sides of back of head and neck with long, flat, upturned fringe-like scales; both
sides of 4th toe with long, well-developed fringes

* Phrynocephalus luteoguttatus Boulenger 1887

51b. Some enlarged dorsal scales nail-like, often tubercular, large part of scale raised free
of back; sides of back of head and neck without long flat, upturned fringe-like scales
(but sometimes with short spiny scales); one or both, sides of 4th toe with short
fringe 52

52a. Nasal shields in contact, or rarely separated by a single series of scales; crossbars on
tail most intense (black) ventrally, though usually quite dark dorsally as well; always
present ventrally Phrynocephalus scutellatus (Olivier 1807)

52b. Nasal shields separated by 3-5 (exceptionally 1, usually 3) series of scales; crossbars
on tail usually most intense dorsally, rarely absent, and much lighter or absent
ventrally, sometimes interrupted dorsally, and seen as a series of spots along sides
of tail 53

53a. No longitudinal crest of mucronate scales; a distinct transverse fold of skin across
back of neck; entire nostril not seen when viewed from side of head; width of space
between nostrils considerably smaller than distance between nostril and preocular
ridge Phrynocephalus helioscopus helioscopus (Pallas 1771)

53b. A longitudinal nuchal crest of 3-8 mucronate, tubercular scales; no transverse fold of
skin across back of neck; entire nostril seen when viewed from side of head; width of
space between nostrils equal to space between nostril and preocular ridge

Phrynocephalus helioscopus persicus de Filippi 1863

54a. Sides of head and neck with long, projecting fringe-like scales; row of enlarged up-
raised tubercular scales on posterior margin of thigh and sides of tail forming shorj
fringe; often a row of slightly enlarged scales along flank

* Phrynocephalus inters capularis Lichtenstein 1856

54b. Sides of head and neck without projecting fringe-like scales; no fringe of scales on
posterior margin of thigh and sides of base of tail; no enlarged scales along flank. 55

55a. Nasal shields separated by 1-3 series of scales; ventral surface of tail with indistinct
dark crossbars, or entire tip dark gray

Phrynocephalus maculatus maculatus Anderson 1872

55b. Nasal shields in contact, or partially separated; tail with 4 or 5 jet-black crossbars
ventrally, tip of tail not black nor gray 56

56a. Distinct dark-margined light dorsolateral stripe from posterior angle of eye along
body onto tail; single very elongate suborbital scale, 2 or 3 times as long as adjacent
scales * Phrynocephalus clarkorum Anderson and Leviton 1967

56b. No light stripe along side of body; 3 suborbital scales of about equal size

Phrynocephalus ornatus Boulenger 1887

57a. Femoral pores present; tail strongly depressed throughout most of its length, shorter
than snout-vent length, covered above by whorls of very large, spinous tubercles
which are rounded at their bases 58

57b. Femoral pores absent; tail not strongly depressed, except sometimes at base, longer
than snout-vent length unless broken, without whorls of large spinous tubercles
rounded at base (large keeled mucronate scales may be arranged in annuli, how-
ever) 60

36 F1ELD1ANA: ZOOLOGY. VOLUME 65

58a. Whorls of spinous scaks on upper surface of tail not separated b\ small scales, back
without transverse rows of enlarged spinous tubercles

Uromastvx microlepts Blanford 1874

58b Whorls of spinous scales on upper surface of tail separated by small scales, back with
more or less regular transverse rows of enlarged spinous tubercles 59

59a. 9-15 femoral and preanal pores on each side; 7-10 tubercles across base of tail: 20-25
transverse rows of scales on middle of belly, on space corresponding to length of
head (tip of snout to angle of jaw) Uromastvx asmussi (Strauch 1863)

59b. 15 or more femoral and preanal pores on each side: 12 tubercles across base of tail;
30-40 transverse rows of scales on middle of belly, on space corresponding to length
of head Uromastvx ioruatus (Blanford 1874)

60a. Well-marked dorsel crest, at least on neck Caiotes versicolor (Daudin 1802)

60b. No dorsal crest 61

61a. Caudal scales obliquely arranged, not forming annuli: tympanum small, more or less
decpK sunk 62

61b. Caudal scales forming more or less distinct annuli: tympanum usually larger than
eye, superficial 65

62a. Dorsal scales homogeneous, large scales of back grading into progressive^ smaller
scales of flanks, no distinctly larger scales among them . .Agama agiiis Olivier 1807

62b Dorsal scales heterogeneous, back and usually flanks with scales of varying sizes
intermixed 63

63a. Abdominal scales distinctly keeled: largest dorsal scales about twice width of adjacent
small scales: at least anterior oval vertebral spots linked together to form undulating
gray or lavender vertebral stripe on neck and back, bordered by brown (darker)
stripes extending onto dorsal surface of head: males with distinct gular sac

Agama hlanfordi Anderson 1966

63b Abdominal scales smooth (rarely faintly keeled): largest dorsal scales about 3 times
width of adjacent small scales: oval vertebral spots often indistinct, contained within
dark crossbars, and not linked into longitudinal stripe: males without gular sac 64

64a. Upper surface of thigh with patch of enlarged scales usually distinct, intermixed with
smaller scales, flanks with numerous enlarged scales among smaller scales: "glandu-
lar" callose preanal scales in 2 rows: small patch of scales on neck just posterior to
occiput in which direction of imbrication is reversed, i.e.. these scales have anterior
margins imbricate Agama ruderaia ruderata Olivier 1807

64b Upper surface of thigh usually lacking distinctly enlarged scales, or with an area of
large scales not intermixed with small scales; enlarged scales of back do not extend
onto flanks; "glandular" preanal scales in single row; none of the neck scales show-
ing reversed imbrication Agama ruderaia megalonyx (Gunther 1864)

65a. Flanks without enlarged scales or tubercles: distal two-thirds or more of tail with
segments composed of more than 2 annuli when viewed laterally (anterior portion
of tail up to 2 or 3 head-widths posterior to vent may have only 2 annuli per segment),
or segmentation indistinct 66

65b. Ranks with enlarged scales, arranged in patches or in regular series: segments of
tail composed of 2 annuli throughout length of tail 68

66a Median dorsal scales in straight longitudinal series. 6-10 across middle of back, grad-
ing into dorsolateral scales: hemipenes of male nonpigmented

• Agama melanura iiraia (Blanford 1874)

66b. Median dorsal scales in oblique longitudinal series. 16-20 across middle of back,
clearly set off from dorsolateral scales; hemipenes of male black 67

67«. A prominent transverse fold of skin across nape Agama nupw nupia dc Filippi 1843

67b. No fold of skin across nape Agama nupia fusea (Blanford 1876)

ANDERSON: KEY TO REPTILE GROUPS 37

68a. One or 2 longitudinal rows of clusters of spiny tubercles on each side of body; 90-102
scales round middle of body; gular scales strongly keeled (weakly keeled in small

juveniles), mucronate Agama erythrogastra (Nikolsky 1896)

68b. Enlarged scales on flanks not arranged in longitudinal rows; ISO or more scales round

middle of body; gular scales smooth, not mucronate 69

69a. Males with 1 15-188 (usually less than 170) scales round middle of body, females with

II 9- 174 Agama caucasica caucasica (Eichwald 1831)

69b. Males with 177-235 scales round middle of body, females with 190-239

Agama caucasica microlepis (Blanford 1874)

70a. Abdominal scales similar to dorsals; no femora! or preanal pores; tongue nicked

anteriorly; body with osteodermal plates; premaxillary bones paired. SCINCIDAE

70b. Abdominal scales subquadrangular or quadrangular, much larger than dorsals, in
6-18 longitudinal rows across venter; femoral pores present; tongue deeply forked;

no osteoderms on body; premaxillary bone single. LACERTIDAE 86

71a. Eyelids immovable (spectacle); small species (adults less than 65 mm. from snout to

vent); limbs well developed 72

71b. Eyelids movable; adults more than 65 mm. from snout to vent; limbs well developed

or reduced 74

72a. Prefontals forming a median suture; 2 frontoparietals

Ablepharus bivittatus bivittatus (M£n6tries 1832)

72b. Prefontals separated; usually a single frontoparietal 73

73a. Ear opening distinct Ablepharus pannonicus (Lichtenstein 1823)

73b. Ear hidden Ablepharus grayanus (Stoliczka 1872)

74a. Digits fringed laterally Scincus conirostris Blanford 1881

74b. Digits not fringed 75

75a. Limbs greatly reduced, with less than 5 digits; body elongate, serpentine 76

75b. Limbs well developed, with 5 digits; body robust 80

76a. Fingers 4, toes 3 77

76b. Fingers 3, toes 2 or 3 78.

77a. Scale rows 20 at midbody1 Ophiomorus blanfordi Boulenger 1887

77b. Scale rows 22 at midbody Ophiomorus brevipes (Blanford 1874)

78a. Toes 2 Ophiomorus persicus (Steindachner 1867)

78b. Toes 3 79

79a. Parietals in contact posteriorly, prefrontals not in contact with supralabials (20 scale

rows at midbody) Ophiomorus streeti Anderson and Leviton 1966

79b. Parietals not in contact posteriorly; prefrontals in contact with supralabials (usually
22, occasionally 20 scale rows at midbody) . . Ophiomorus tridactylus (Blyth 1853)

80a. Lower eyelid with transparent shield 81

80b. Lower eyelid without transparent shield 83

81a. Nostril between nasal and rostral, in emargi nation of latter; scales smooth; back with
numerous dark-margined light ocelli irregularly transversely arranged

Chalcides ocellatus ocellatus (Forskal 1775)
81b. Nostril in nasal shield; dorsal scales usually distinctly, but weakly bi- or tricarinate;

back without ocelli 82

82a. Parietal scales usually in contact behind interparietal; nuchals and postnuchals with
3 strongly developed keels; often a distinct light vertebral stripe, usually dark-mar-
gined and clearly set off from ground color Mabuya vittaia (Olivier 1804)

'Counts must be made exactly midway between snout and vent.

38 F1ELDIANA: ZOOLOGY, VOLUME 65

82b. Parietal scales not in contact; nuchals smooth, post-nuchals smooth or very weakly

keeled; no light vertebral stripe Mabuya aurata (Linnaeus I7S8)

83a. 2 median rows of dorsal scales united into single row of broad scales; postnasal shield

present Eumeces laeniolatus (Blyth 1 854)

83b. 2 median rows of dorsal scales broader than those on flanks; no postnasal shield

84a. Dorsum with dark vermiculate or mottled pattern, mid-dorsal spots tending to form

longitudinal lines Eumeces schneiderii variegatus Schmidt 1939

84b. Dorsum without dark vermiculate or mottled pattern 85

85a. Base of tail reddish in life1 Eumeces schneiderii zarudnyi Nikolsky 1899

85b. Base of tail not reddish in life, dorsum with or without orange or reddish flecks.

Eumeces schneiderii princeps (Eichwald 1839)

86a. Eyelids immovable (spectacle) Ophisops elegans Menetries 1832

86b. Eyelids movable 87

87a. Nostril separated from 1st supralabial by nasal shield 88

87b. Nostril in contact with 1st supralabial, or separated from supralabial by very narrow

brim 100

88a. Ventral plates in straight longitudinal series; lower nasal resting on 1st supralabial.

88b. Ventral plates in tessellated or oblique longitudinal series, converging posteriorly;

lower nasal resting on 2 or 3 supralabials 90

89a. Occipital in contact with interparietal, or separated from it by small shield; large
transparent scales of lower eyelid edged with black; ventral plates in 10 longitudinal

series Eremias guttulata (Lichtenstein 1823)

89b. Occipital absent or minute, not in contact with interparietal; transparent shields of
lower eyelid not edged with black; ventral plates usually in 12 (rarely 10 or 14)

longitudinal series Eremias brevirostris (Blanford 1874)

90a. Subocular bordering mouth 91

90b. Subocular not bordering mouth 97

91a. Lateral scales of 4th toe forming a distinct fringe in its entire length 92

9 1 b. Lateral scales of 4th toe not forming a distinct fringe 93

92a. A broad dark dorsolateral stripe from nostril through eye, along body and side of tail,
one or 2 additional narrower dark stripes mediad to these on each side, the remainder
of the dark dorsal stripes interrupted and anastomosing to form a reticulate pat-
tern, evident even in very young specimens; 4th toe with 2 complete rows of sub-
digital scales, i.e., a total of 4 scales counted around toe (except that an extra scale

may be present at a joint) * Eremias scripta (Strauch 1867)

92b. Dorsal pattern consists of 7 dark stripes, the outer dorsolateral stripe broadest, these
stripes persisting unbroken in both adults and juveniles; 4th toe with single row of
subdigital scales, i.e., total of 3 scales counted around toe (except an extra scale may

be present at a joint) Eremias lineolata (Nikolsky 1896)

93a. The 2 series of femoral pores broadly separated, space between the 2 series at least

one-third the length of each Eremias pleskei Bedriaga 1907

93b. The 2 series of femoral pores meeting, or separated by space not greater than one-
fourth length of each 94

94a. Back with 5-1 1 dark stripes, broader than interspaces, none of the stripes containing
light ocelli or spots; stripes persistent in adults, but sometimes indistinct so that

1 Preserved individuals in which the color has faded cannot be identified to the sub-
species level, as far as I have been able to determine.

ANDERSON: KEY TO REPTILE GROUPS 39

back appears almost uniform sandy; 4th toe with 2 complete rows of subdigital
scales and a complete row of sharply pointed lateral scales, i.e., a total of 4 scales
counted around penultimate phalanx; collar scales small, usually only a single
median collar scale distinctly larger than adjacent gulars.

Eremias fasciata Blanford 1874

94b. Light ocelli or spots on upper flanks (rare exceptions), dark stripes of juveniles break-
ing up in adults to form spots or broken lines; 4th toe with single complete row of
subdigital scales, a complete row of somewhat smaller ventrolateral scales, and a few
scattered, much smaller, ventrolateral scales not forming complete row; total of 3
scales counted around penultimate phalanx; usually several collar scales distinctly
larger than adjacent gulars 95

95a. Adults with dark interrupted dorsolateral black stripe forming ocelli with white spots,
this dorsolateral pattern not contrasting strongly with interrupted dark stripes and
spots of dorsum; juveniles with 3 dark stripes on dorsum between white-spotted
dorsolateral stripes, vertebral stripe being black, bifurcated on nape (dark stripes
breaking up into several irregular rows of dark spots with age); ventral surface of
tail carmine red in juveniles (in life) Eremias velox velox (Pallas 1771)

95b. Adults usually with black dorsolateral stripe, more or less continuous for at least
major portion of its length, containing white spots, black stripe contrasting strongly
with dorsal color pattern; juvenile with 4 dark stripes on dorsum between dorso-
lateral white-spotted stripes, vertebral stripe being white (dark stripes breaking
up into 4 more or less regular rows of dark spots with age); ventral surface of tail not
red in juveniles 96

96a. Adults with 4 more or less distinct rows of dark spots on dorsum between dorsolateral
dark stripes; dark dorsolateral stripes usually containing white spots in single row;
distal portion of tail bluish in juveniles (in life)

Eremias persica Blanford 1874

96b. Adults usually without dark stripes or spots on mid-dorsum; dorsolateral region with
alternate rows of light and dark spots, often fusing longitudinally, forming 2-4 longi-
tudinal stripes, often broken, the impression being 3-4 rows of white spots on flanks;
ventral surface of tail yellow in juveniles (in life) . . . Eremias strauchi Kessler 1878

97a. 4th toe with distinct fringe on both lateral and medial sides, formed by complete row
of sharply pointed lateral scales and complete row of similar medial scales; ungual
lamellae of fingers and toes with prominent, flat, lateral expansions

Eremias grammica (Lichtenstein 1823)

97b. 4th toe without distinct fringe; ungual lamellae without prominent lateral expansion. 98

98a. 5th toe with 2 complete rows of subdigital scales and incomplete row of small lateral
scales; 2nd supraocular ( 1st of 2 large, undivided supraoculars) as long as or shorter
than its distance from 2nd loreal Eremias arguta (Pallas 1771)

98b. 5th toe with single complete row of subdigital scales and a few scattered lateral scales
not forming complete row; 2nd supraocular (1st of 2 large, undivided supraoculars)
longer than its distance from 2nd loreal 99

99a. 4th toe with single row of subdigital scales; usually distinct tympanic shield; 4th
supraocular usually distinct Eremias intermedia (Strauch 1876)

99b. 4th toe with 2 rows of subdigital scales, internal much larger; tympanic scale usually
small or indistinct; 4th supraocular usually indistinct

Eremias nigrocellata (Nikolsky 1896)

100a. Digits with lateral fringes 101

100b. Digits without lateral fringes 104

40 FIELDIANA: ZOOLOGY, VOLUME 65

Hil.i 3 scales around fingers; ventrals usually 10 in longest transverse row across belly;
dorsal scales feebly keeled, 48 or more across middle of body

Acanthodactylus micropholis Blanford 1874

101 b. 4 scales around fingers; ventrals 13-18 in longest transverse row across belly; dorsal
scales strongly keeled, 54 or less across middle of body 102

102a. Ventral scales in oblique or irregular longitudinal series, not forming straight longi-
tudinal rows; 18-22 dorsal scales in transverse series between hind limbs

Acanthodactylus fraseri Boulenger 1918

102b. Ventral scales in straight longitudinal rows, at least down middle of venter; outer
series may be somewhat oblique; 10-16 dorsal scales in transverse series between
hind limbs 103

103a. Dorsal color pattern reticulate, not lineate even in young specimens, indistinct in
large adults; 13-18 ventral plates in longest transverse series; 38-54 dorsal scales
across middle of back Acanthodactylus cantoris schmidti Haas 1957

103b. Dorsal color pattern lineate, young specimens with 6 dorsal and one lateral light
longitudinal streaks, with or without round white spots between them; some adults
nearly uniform, no distinct pattern; 12-16 ventral plates in longest transverse series;
34-46 dorsal scales across back Acanthodactylus cantoris blanfordi Boulenger 1918

104a. Lower eyelid with 5-7 transparent shields edged with black; subdigital lamellae
keeled Apathya cappadocica urmiana 1 ant/ and Suchow 1934

104b. Lower eyelid without transparent shields; subdigital lamellae smooth or tuberculate
105

105a. Ventral plates more or less rectangular with rectilinear or nearly rectilinear posterior
margins 106

105b. Ventral plates trapezoidal, with notches between longitudinal rows 108

106a. Dorsal scales strongly keeled, more or less distinctly hexagonal; collar serrated

Lacerta chlorogaster Boulenger 1908

106b. Dorsal scales smooth, granular, round or oval, collar not serrated 107

107a. 5-6 (rarely 4) supralabials anterior to subocular; normally 2 superposed postnasals
(but sometimes fused on one or both sides of head); pterygoid teeth strongly devel-
oped; outer ventrals with small black spots Lacerta hrandtii de Filippi 1863

107b. 3-4 (rarely 5) supralabials anterior to subocular; normally a single postnasal; ptery-
goid teeth absent; outer ventrals without black spots (turquoise blue spots present in
males) Lacerta saxicola Eversmann 1 834

108a. Ventral plates in 10 longitudinal series; 34-37 dorsal scales across middle of body
109

108b. Ventral plates in 6 or 8 longitudinal series; 38 or more dorsal scales across middle of
body 110

109a. Outer row of ventrals (marginals) smooth; 20-22 gulars; 13-17 femoral pores on each
side; lower edge of subocular one-half or less than one-half maximal length of shield

Lacerta princeps princeps Blanford 1 874

109b. Outer row of ventrals (marginals) keeled; 17-19 gulars; 16-21 femoral pores on each
side; lower edge of subocular one-half or more than one-half maximal length of
shield Lacerta princeps kurdistanica Suchow 1936

1 10a. 17-21 femoral pores, row of pores reaches knee; usually less than 20 temporal scales;
5th submaxillary shield always well developed; young specimens usually with unin-
terrupted lateral light line in addition to vertebral and dorsolateral lines

Lacerta strigata Eichwald 1831

1 10b. 1 2-16 femoral pores, row of pores does not attain knee; usually more than 20 temporal
scales; 5th submaxillary small or absent; young specimens with lateral light line
interrupted in its anterior half .... Lacerta trilineata media 1 ant/ and Cyren 1920

ANDERSON: KEY TO REPTILE GROUPS 41

REFERENCES

Anderson, Steven C.

1968. Zoogeographic analysis of the lizard fauna of Iran. In Fisher, W. B., ed.. The
Cambridge history of Iran, vol. I, The land of Iran, pp. 305-371, Cambridge University
Press, London.

Fisher, W. B.

1968. Physical geography. In Fisher, W. B., ed., The Cambridge history of Iran, vol. 1,
The land of Iran, pp. 3-110, Cambridge University Press, London.

Peters, James A.

1964. Dictionary of herpetology. Hafner, New York. 392 pp.

Schmidtler, Josef Johann and Josef Friedrich Schmidtler

1972. Zwerggeckos aus dem Zagros-Gebirge (Iran). Salamandra, 8, pp. 59-66.

Witte, Gaston Fr. de

1973. Description d'un Gekkonidae nouveau de l'lran (Reptilia Sauria) Bull. Inst. r. Sci.
nat Belg. Biologie, 49, pp. 1-6.

FIELDIANA: ZOOLOGY, VOLUME 65

Table I. Summary of distribution of turtles, lizards, and amphisbaenians among the
political divisions of Iran (see fig. 1).

Key: ♦ definite record; ? doubtful record; P probable occurrence, but no record. Species
preceded by an asterisk (*) have not been recorded definitely from Iran.

Species

I 2 3 4 5 6 7 8 9 10 II 12

1. Emys orbicularis

♦

2. Mauremys caspica caspica

♦

•

♦

3. Testudo graeca ibera

♦

•

♦

4. Testudo graeca zarudnyi

♦

•

5. Testudo horsfieldii

•>

♦

6. Trionyx euphraticus

•

7. Agama agilis

♦

•

<•

•

♦

8. Agama blanfordi

9. Agama caucasica caucasica

•

♦

10. Agama caucasica microiepis

♦

•

II. Agama erythrogastra

* Agama melanura lirata

1 2. Agama nupta nupta

•

♦

13. Agama nupta fusca

14. Agama ruder at a ruder at a

1 5. Agama ruderata megalonyx

16. Calotes versicolor

♦

* Phrynocephalus clarkorum

1 7. Phrynocephalus helioscopus helioscopus

1 8. Phrynocephalus helioscopus persicus

* Phrynocephalus interscapularis

* Phrynocephalus luteoguttatus

19. Phrynocephalus maculatus maculatus

20. Phrynocephalus mystaceus galli

♦

21. Phrynocephalus ornatus

♦

22. Phrynocephalus scutellatus

23. Uromastyx asmussi

24. Uromastyx loricatus

25. Uromastyx microiepis

26. Anguis fr agilis colchicus

•>

♦

27. Ophisaurus apodus

♦

28. Agamura persica

29. Alsophylax spinicauda

30. Bunopus aspratilis

•

31. Bunopus crassicauda

•

32. Bunopus lubercutatus

•

♦

33. Crossobamon eversmanni

♦

34. Cyrtodactylus agamuroides

35. Cyrtodactylus brevipes

■>

♦

36. Cyrtodactylus caspius

♦

37. Cyrtodactylus fedtschenkoi

•

38. Cyrtodactylus gastropholis

♦

39. Cyrtodactylus heterocercus heterocercus

40. Cyrtodactylus kachhensis

♦

(continued)

ANDERSON: KEY TO REPTILE GROUPS

Species

1 2 3 4 5 6 7 8 9 10 II 12

41. Cyrtodactylus kirmanensis

*Cyrtodactylus kotschyi

42. Cyrtodactylus russowii

43. Cyrtodactylus sagittifer

44. Cyrtodactylus scaber

45. Eublepharis angramainyu

46. Eublepharis macularius

47. Hemidactylus flaviviridis

48. Hemidactylus garnotii

49. Hemidactylus persicus

50. Hemidactylus turcicus turcicus

5 1 . Asaccus elisae

52. Asaccus griseonotus

53. Pnsturus rupestris

54. Ptyodactylus hasselquistii

55. Stenodactylus affinis

56. Stenodactylus doriae

57. Teratoscincus bedriagai

58. Teratoscincus microlepis

59. Teratoscincus scincus

60. Tropiocolotes helenae

61. Tropiocolotes heteropholis

62. Tropiocolotes latifi

63. Tropiocolotes persicus persicus

64. Tropiocolotes persicus bakhtiari

65. Acanthodactylus cantoris blanfordi

66. Acanthodactylus cantoris schmidti

67. Acanthodactylus fraseri

68. Acanthodactylus micropholis

69. Apathya cappadocica urmiana

♦

70. Eremias arguta

♦

71. Eremias brevirostris

♦

72. Eremias fascial a

73. Eremias grammica

74. Eremias guttulata

♦

75. Eremias intermedia

76. Eremias lineolata

77. Eremias nigrocellala

♦

78. Eremias persica

♦

79. Eremias pleskei

* Eremias script a

80. Eremias strauchi

81. Eremias velox velox

82. Lacerta brandtii

83. Lacerta chlorogaster

♦

84. Lacerta princeps princeps

85. Lacerta princeps kurdistanica

86. Lacerta saxicola defilippii

(continued)

FIELDIANA: ZOOLOGY. VOLUME 65

Species

87. Ijneria sa.xicola raddei

88. iMceria singula

89. Ijjcerta irilineaia media

90. Ophisops elegans

91. Ablepharus hivit talus hi villains

92. Ahlepharus grayanus

93. Ahlepharus pannonicus

94. Chalcides ocellatus ocellaius

95. Eumeves schneiderii princeps

96. Eumeves schneiderii variegatus

97. Eumeces schneiderii zarudnvi

98. Eumeces taeniolatus

99. Mahuva aurata

100. Mahuva villala

101. Ophiomorus hlanfordi

102. Ophiomorus brevipes

103. Ophiontorus persicus

104. Ophiomorus street i

105. Ophiomorus tridactylus

106. Scincus conirostris

107. Varanus hengalensis hengalensis

108. Varanus griseus griseus

1 09. J aranus griseus caspius

1 1 0. Diplometopon zarudnvi
TOTAL (species and subspecies)

I 2 3 4 5 6 7 8 9 10 II 12

23 18 13 13 14 32 21 41 30 43 39 26

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