Asiatic Herpetological Research

1999

Vol. 8, pp. 60-68

Redescription and Generic Redesignation of the Ladakhian Gecko
Gymnodactylus stoliczkai Steindachner, 1969

Muhammad S. Khan and Herbert Rosler 2
1'12731 SW8th Ct.. Davie, Florida, 33325 USA

Abstract.-

stoliczkai

Gymnodactylus

redesignated to theis2

F Freiligrath Str. 51, 06502 Thale/Harz, Germany

redescribed on the basis of recently collected topotypes, and is
It is found to be morphologically distinct from C. walli and C.

genus the Cyrtodactylus.

yarkandensis with which species it has long been synonymized. Reproduction, ecology and distribution of thishighland gecko is discussed.

Key words.- Cyrtodactylus stoliczkai, taxonomy, reproduction, ecology, zoogeography.

IntroductionIn

1866 Dr. Ferdinand Stoliczka deposited a collec-

tion of animals, collectedfrom different parts of the

Indo-Pakinsubcontinent,

Naturhistoristorisches

Museum Wien, Austria (NMW). The collection contained a gecko from Karoo, north of Dras. Kashmir,apparently collected during geological survey of western Himalayas, which was described by Steindachnerin1867 as a new taxon, Gymnodactylus stoliczkai,

honoring its collector. The specimen is still available
in the museum under registry number
16756

NMW

(Tiedemann et al., 1994). Ever since the description ofthis taxon,ithas become a habit with herpetologists

working on collections from northern Pakistan, to try
to place almost every angular-toed gecko encountered, in the synonymy of G. stoliczkai without going

into details of morphological comparisons:

Smith,

1935 (Gymnodactylus walli Ingoldby, 1922): Minton,

types of this species in Zoologische Staatssammlung
Munchen (ZSM: Table 1: collected by Gruber, 1981),

while Dr. G. R. Zug (National Museum of Natural
History, Washington) has kindly taken data on topotype

MCZ 7132 for us. Moreover, present redescrip-tion is further supplemented with the data available inliterature(Steindachner,

1867:15:

Boulenger,

1890:63; Smith, 1935:57; Constable, 1949:84; Szcz-

erbak and Golubev, 1986:205). Photographs of topotypes (Fig.

1;

ZSM 124.77) and (Fig. 2; ZSM 45.77),

and type NMW 16756 (Khan, 1994; Fig. 2) havehelped us immensely to understand morphology ofthis important taxon.

Taxonomic Notes
Sprix's (1825:17) genus Gymnodactylus included all
known non dilated angular-digited geckos

the then

1966 (Cyrtodactylus mintoni Golubev and Szczerbak,(Boulenger, 1885:22, 1890:59; Annandale, 1913:309;
Smith, 1935:37). Until Underwood (1954) restricted1981) and Mertens, 1969 (Cyrtodactylus dattanensis

this genus to South American angular-digited geckos,

Khan, 1980), creating taxonomic chaos. Recent collections from circum-Himalayan region has shownplacing all south Asian geckos in the genus Cyrtodactylus Gray 1827. Most of the subsequent workers onthat G. stoliczakai

does not belong to the fauna of

the herpetology of Pakistan have followed

Under-

ymy are themselves valid independent taxa (Khan,

wood's view point (Minton, 1966; Khan, 1980; Khanand Mirza, 1977). However, Mertens (1969) is ortho-

1992, 1993, 1994; Khan and Baig, 1992).

dox and a bit cautious by retaining Gymnodactylus as

Pakistan, moreover, all the geckos placed in its synon-

A long, detailed morphological redescription of
Gynnodactylus stoliczkai

isurgently due, tomakea genus and placing Pakistani geckos in the subgenus

Cyrtodactylus.

comparisons possible and to come out of

A recent break through towards a solution comesDue to lack of working material

from Szczerbak and Golubev (1984, 1986): the genus

the present project has considerably been delayed.
The type specimen
16756) and syntype in the

Tenuidactylus is erected to include Palearctic angulardigited geckos. It is divided in three subgenera to

Museum of Comparative Zoology, Cambridge (MCZ

accommodate the rest of the southeast Asian gekkota:

intertaxal"stoliczkai myth".

(NMW7132) are not allowed to be loaned for study. There-

subgenus Tenuidactylus includes two Pakistani speT. montiumsalsorum and T. kohsulaimanai, andfore the data for present study is drawn from several

cies

sources: one of us (HR) has studied series of 14 topo-

the group of Tibeto-Himalayan species:T.

tibetanus.

Asiatic Herpetological Research

1999

Table 1.

Vol. 8, p. 61

Range of scale counts and measurements of topotype series of Tenuidactylus stoliczkai in Zoologische

Staatssammlung Munchen (ZSM) collection (data from juvenile ZSM 1 19/77 not taken into account).

Character

Range

Supralabials

9-11

Infralabials

7-9

Nasals3-3Internasals

1

Postmentals

2-2

Loreals

10-15

Scales between eye-ear

15-20

Interorbitals

17-19

Tubercle rows across body middorsum

8- 1 2

Sale rows across midbelly

25-30

Midventrals

110-133

Lamellae under first toe

11-14

Lamellae under fourth toe

22-27

Cloacal spines

2-3

Granular scale rosette around dorsal tubercle

7-9

SVL

27-49.4mm

TL

25-49.7mm

Head length

8-1 1 .5mm

Head width

5-10.7mm

Head height

3-6.1

mm

Nostril-eye distance

2-4.4

m

Eye-ear distance

2-4.1

mm

Body length10-23.6

Eye diameter

2-2.8

Ear diameter

0.4-0.9

Head length/head width

0.99-1 .43mm

SVUbody length

2.09-2.64mm

Head length/head width

0.99-1 .43

mm

Distance nostril-eye/eye-ear

0.98-1.29mm

Eye/ear diameter

2.33-5

-3

mm

mm
mm

mm

1999

Asiatic Herpetological ResearchVol. 8, p. 62

thS/1977/tfDACH4L

Figure 1. Cyrtodactylus stoliczkai (ZSM 124.77), adult

Figure 2. Cyrtodactylus stoliczkai (ZSM 45.77), adultfemale, with regenerated tail.

female, with unregenerated tail.

rected as a second subgenus to include four Pakistani

body: Cyrtodactylus pulchellus, interrnedius,
consobrinoides, angularis, khasiensis, rubidus, triedrus, nebulosus, collegalensis, dekkanensis, albofas-

forms: agamuroides, scaber, watsoni and kachhensis.

ciatus and jayporensis.

A third subgenus Medioductylus

Circum-Himalayan group:T.

mintoni,T.

chitralensis,T.

stoliczkai and T.kirman-

ensis; the genus Cyrtopodion Fitzinger,

is left

1

843 is resur-

floating (Gol-ubev, pers. comm.. 1996).

Undoubtedly southeast Asian cyrtodactylid geckos
are morphologically distinct from Palearctic tenuidactylids (Leviton and Anderson, 1970: Khan, 1988,
1989, 1991; Khan and Tasnim, 1990). Khan (1993)

thansubtropical, highlandforms, mainly extending between lat. 34-4° N, long
75° 50-50'E, tubercular, beady, scarcely imbricate

granular dorsal scales, interspersed by 2-3 times
inlarger oval keeled or keelless tubercles arranged

more or less in 12-13 longitudinal rows; body and tailsubequal and subcylindrical, subcaudals small in sevinconspicuous dorsal pattern of transverse

maintains the genus Cyrtodactylus Gray, 1827, to
include Tibeto-Himalayan species group and all theeral rows,

southeast Asian species (Smith, 1935). The southeast

bands, spots or reticulation.

Asian cyrtodactylids are a very heterogeneous assem-

Further distinguished in three subgroups:

blage of closely allied species. Their shared charactersare: smooth tubercular granular scales with scatteredstoliczkai subgroup: body and tail rather flat in crossround-oval smooth or slightly keeled tubercles onhead and body dorsum; more than 25 heterogeneoustailsubcylindrical and subequal body andvivid
dorsaltail;pattern; subdigital lamellae about
twice as broad as high with a pair of lateral row ofinterorbitals;

granular scales, lamellae not swollen at the digital
femoralangles; 2-10 preanal pores in male, raresmall
subcaudals
caudalblunt
small
tubercles,
pores;
broad. In the past there have been several
attempts to arrange them in a logical array (Annanrarely

dale, 1913; Smith,1935: Khan, 1993). Consideringmorpho-ecogeography of these geckos, weguish two lineages:

distin-section, caudal tubercles flat, smooth; anterior half ofsegmented, segments laterally lobulated in older
animals, regenerated tail flattened and abnormallyswollen, no preanal and femoral pores. Includes highland species: Cyrtodactylus lawderanus, C. stoliczkai,
C. yarkendensis and C. baturensis.

tibetinus subgroup: Body and tail round in cross sec-

segments not distinct, 4-10 preanal pores,
keeledcaudal tubercles, regenerated tail not
feeblyflattened. Dorsal pattern of vivid cross bars, spots ortion, tailreticulation. Includes Tibeto-

submountaingeckos:

Himalayan low altitude

Cyrtodactylus

tibetinus,C.

himalayanus, C. mintoni, C. dattanensis and C. battalensis.

Circum-oceanic group: tropical, scattered along sub
continental coastal strip and oceanic islands, confined

between lat. 7-32° N, long 75-105° E; dorsal patternof vivid cross bars or spots, dorsal granular scales
mixed with larger rounded, smooth or slightly keeledtubercles, tailand body cylindrical, tail often longer

walli

subgroup: Body

flatter,tailquadrangularincross section, distinctly segmented, caudal tubercles
translarge slightly keeled, median row of subcaudals
versely enlarged; 4-6 preanal pores. Species
are:Cvrtodactvlus kinnanensis and C. walli.

included

1999

Asiatic Herpetological ResearchVol. 8, p. 63

Table 2. Comparison of morphology of 1= Tenuidactylus yarkandensis (J. Anderson, 1872) with its closest congeners: 2= T. stoliczkai (Steindachner, 1867), 3= T. iva///(lngoldby, 1922) and 4= T. baturensis Khan and Baig, 1992;
vl=vertical, (measurements in mm).

CharacterVol. 8, p. 64

Asiatic Herpetological Researchdark bands with heavier posterior margin, broader
than interspaces.

Description of holotype

NMW 16756, Fig.

1,(state-ments in parenthesis are from paratype and topotypes
adding up to the original description of holotype

NMW 16756 by Steindachner, 1867:15): (habitus
depressed); rostral scale big (7-angular,

broader thandeep), slightly convex at the upper edge, and forked in
the middle reaching the anterior end of the snout(median dorsal rostral longitudinal furrow narrowly
misses anterior border of the rostral scale); nasal

opening (small, dorsolateral) bordered in front by rostral plate,(ventrally by) second upper lip shield (first

1999(Gular scales flat, juxtaposed or slightly imbricate,mostly arranged in transverse rows interspersed withlarger scattered tubercles,

becoming flatter and pen-tagonal on chest, at abdomen flat, hexagonal, broader
than long, slightly imbricate rarely juxtaposed,inarrangedtransverse

rows,

30-32

scalesacrossmidabdomen; slightly marked lateral abdominal folds,
5-6 rows of lateral abdominal scales differ little fromdorsals, however clearly marked from abdominals;
femoral and preanal pores absent (even their site notmarked by distinct scales; 142-149 midventral scales
between first pair of submental and anterior anal lip. A
pair of lateral cloacal tubercles, no postanal bulge).

by(Limbs medium sized, covered with imbricatesmooth scales arranged in transverse and longitudinal

three small nasal shields, of which upper most is therows, without tubercles; subfemorals in 5-6 transverse

largest (separated from that of other side by a pair ofrows, as large as abdominals: median-subtibial scales

granular scales, rostral area with heterogeneous tuber-largest, imbricate, arranged in trans verse rows; gran-cular scales mostly arranged in longitudinal rows,

ular postfemorals extend on to the sides of preanals,

supralabial. Notesupralabialinrostral shield is regarded asoriginal

description),

firstposteriorly

11-

12 tubercular loreal scales between posterior nasals
and anterior rim of orbit; head with heterogeneous
tubercular

granularscales,

19-20

betweenorbitsarranged in longitudinal rows, smaller on eye bulgings; a series of sharp supraciliary scales jutting out

from posterior half of the upper eyelid border followed by a row of large tubercular round scales running along the eyelid; temporals and neck with small
tubercular granular scales: ear opening round, muchwith no tubercles; tips of finger-claws extend to ante-of eye, when forelimbs are adpressed forward,
those of toes a little anterior of axilla; subdigital

rior

23-24

lamellaeunder

fourthtoe,equally

not enlarged under basal

throughout,parts of digits).

broad

and angular

Tail, as it appears in the specimen examined by us,isregenerated in the anterior, somewhat longer thanthe body. Lined on each (dorso- lateral) side by threelarger than largest dorsal tubercle).

rows of large, spike shaped raised tubercles, of which

19-21 upper lip shields (10-11 supralabials, first
three of the same size; suture between first supralabial

those of lower most row are the biggest and conical.

and rostral scale almost equals former's length along(regenerated?,parenthesisby Steindachner),
half
of
the
tail
is
with
uniform
scales (morposterior

oral orifice); 13 lower labial shields (7-9 infralabials,

No largetransverse
thisplatesunder side of the

tail

is

second narrowest while fourth the longest; 2-3 rows

phologically tail of the holotype reminds original tailof sublabials). The anterior most lowerof olderlip shieldvery large, triangular (about twiceasextends
broad,
deep
deep between first pair of post-(mental scale)ismentals), three pairs of chin shields (first pair, largest,in contact

with each other behind posterior

specimens of Hemidactylus flaviviridis.
Doubtless the type specimen has original tail which isnormal for older animals of the species as in ZSM
124.77. The tail is moderately depressed, its stump is

width of body and the segmented

narrowly
tip of mental scale, second pair smaller, less than half

less than half thethe size of first not in contact with each other), thirdventralpair (smallest) almost fully separated from lower labi-

length. Itsals (on right while in contact on left).

ally lobulated segments, while posterior unsegmented

Body dorsum clearly granulated (with flat, juxtaposed,

rarelyslightly

imbricate

granularscales,arranged in transverse rows) with numerous, only little bigger fully rounded tubercles (interspersed with
2-3 times large, flat, smooth, round or oval tubercles,
scattered

evenlyon body dorsum, more or lessfrom eachin longitudinal rows, separated

arrangedother by 3-4 granular scales, a rosette of 7-9 granules

around a tubercle; tubercles on sides slightly conical).

part almost equals body's width, a median dorsal andlongitudinal slight furrow runs along its
anterior half is divided in 10 distinct later-

half gradually narrows to sharp terminal tip; dorsolat-eral^ caudal tubercles are given from the middle ofthe segments, four on 1st to 5th segment, three on 6th
to 10th. The dorsal tubercles are small roundish about

2-3 times smaller than laterals which are elongatedconical blunt, in contact with each other, are gradually

reducedin size, until

almost indistinct

inposteriorhalf of the tail. Dorsally 6-7 transverse rows of hetero-geneous, slightly imbricate, smooth, tubercular scales
cover anterior caudal segments while 4 transverse

Asiatic Herpetological Research

1999

rows of small imbricate subcaudals are present on

Dorsal pattern of bands is vividly distinct in juve-

ventral side of segment. In the posterior quarter dorsal

niles (ZSM

and ventral caudal scales become indistinct from eachgrows older.

other,

getpointedflatter,longer,at

and aretips

strongly imbricate).

Vol. 8, p. 65

119.77), but is gradually lost as the animal

Sex: Though dissection is the sure way of sex deter-

Color: Body dorsum light blue grey, with pink edged

mination of geckos, however, presence of prcanal andfemoral pores in males and their absence in females

transverse bands with denser

arc almost universal sex indicators in these animals. In

wavy posterior edges,

broader than interspaces, three on nape, six on body

NMW

and 13 on tail (in preserved specimens bands are dark

specimen of Tenuidactylus stoliczkai
16756, paratype MCZ 7132 and ZSM series the prea-

and does not extend on subcaudals). (Head), labials

nal and femoral are absent, moreover their position is

the type

and tail plates with fine grey dots, limbs and digits

not indicated by distinctiveness of scales in the area.

barred, ventrum light.

On the other hand, swollen postanal sacs are usually

Measurements (in mm): Snout vent length 48, taildistinct in male less so in female geckos (Smith, 1933,

length 52, trunk length 24, head lengthhead

1935: Khan and Baig, 1992). There is no indication ofhead height 7, eye diameter, not including

postanal sacs in type nor in paratype, however, arewell indicated in two specimens inseries which

width

10,

1

1

.5,

bony orbit 2.5: snout length 5: oculo-orbital space 5,
Variation: Table

presents the measurements and1pholidosic counts in ZSM series which fall within the

NMW6756 and paratype
range of the type specimen
7132, differing in minor details of pholidosic

ZSM

are males. Constable (1949:84) designated

MCZ 7132

as a male specimen, which on dissection is proven to

be an adult female with well developed vitellogen fol-1

MCZlicles (Zug, personal communications, 1989).

morphology: rostral scale is protrubrant in most of thespecimens, however, in some it is flat, the median ros-

Ecology: Ladakh lies around 3000 m, above timber-groove in some specimens extend to middle,
while in other it narrowly misses the anterior end of

etationtral

the scale: the supraciliary pointed scales vary in theirpointedness; dorsal tubercles uniformly scattered on
dorsum, 2-3 times larger than dorsal granular scales,line. It is completely dry snow desert, with sparse veg-of herbs, shrubs and grasses. The area

winter and spring.

Maximum summer July tempera-ture is 24.7° C, minimum 10.2°, while maximum win-

separated from each other by 2-4 granular scales, surrounded by a rosset of 6-9 granular scales; lateral

terabdominal folds strongly or poorly indicated.

received during August,Study of tail morphology of MCZ 7132 and specimens in ZSM series indicates that the tail of the holo-

NMW

16756

is

type
undoubtedly
original.
Steindachner (1867) himself was doubtful about its

being"(regenerated?)".

unregenerated

tail,

the

In
tail

young specimens
is

with

uniformly broad frombasal stump till mid-tail, where it gradually tapers to
its tip(ZSM 45.77:5,

1

19.77, 124.77, 149.77:

1,

2, 4,

As animals get older (NMW 16756 and ZSM
124.77), the anterior half of the original tail becomes
broader, flatter and deeply sected on sides so much soitappears laterally lobulated at segments. From lateral5).

lobes caudal

tubercles

strongly jet

out.

While

in

MCZ 7132 and ZSM 49.77,the tailregenerated
swells up almost round at the base, with no indication
tail,

is

highly arid with sub-tropical continental highlands
cold climate. Heavily snowy winters, getting rain intemperatureinJanuaryis-1.4° C, dropping tominimum -13.3°. Maximum rain fall,
minimum,

1.0

15.0

mm, ismm during

November (Ahmad, 1951).
Gruber (1981) collected T. Stoliczkai from rocky
habitat, where this gecko prefers desert, bare, dry situ-

ations in the non-irrigated areas without or with very

sparse vegetation, apparently avoiding direct neighborhood of human settlements. Other reptiles collectedfrom the area are Phrynocephalus theobaldi.

himalayana, and Scincella ladacensis,
while Bufo latastii is the only amphibian recorded
from waters of the area.

Laudakia

Comparison with congeners: Absence of trihedral
tubercles, preanal and femoral pores, broader subdigitallamellae and peculiar tail morphology differentiate

Tenuidactylus

stoliczkai

from Palearctic group ofC

Pakistaniof segmentation, lobulations and tubercles, while pos-tenuidactylid
Cyrtopodion geckos:
scaber (Heyden, 1827), C. watsoni (Murray, 1892), C.

teriorly it abruptly tapers. Moreover, instead of trans-

kachhensis (Stoliczka,verse bands of original tail, the regenerated tail isspotted with longitudinal spots, scattered all over it.

(Annandale, 1913) and C. kohsulaimanai (Khan,
1991 ). While Pakistani members of Tibeto-Himalayan

ZSM 45.77, 49.77:3, 45.77, 121.77 and ZSM

49.77 represent different stages of tail regeneration ingroup of cyrtodactylid geckos: C. mintoni (Golubev
and Szczerbak, 1981), C. dattanensis (Khan, 1980) C.

Tenuidactylus stoliczkai.

battalensis Khan, 1993 are similar to T. stoliczkai in

Tail in1872), C.

montiumsalsorum

the morphology of subdigital lamellae, body configu-

Vol. 8, p. 66

Asiatic Herpetological Researchration, larger number of interorbitals, subabdominals,

1999

however, they differ markedly from it because of theirlus. yarkandensis, is known from lat. 38° 40'N, long
77° 50'E, along the western border of China's Xin-plump rounded body and tail, tail morphology, dorsal
tuberculation and pattern which extends on the tailjiang Province (Khan, 1994) and Cyrtodactylus stolicc
zkai is confined to Ladak, between lat. 34 -35° 45' N,

ventrum, presence of preanal pores in male.

long 75° 50-76° 70'E (Annandale. 1913; Schmidt,

The tenuidactylid group of Pakistani geckos: T.
indusoani (Khan, 1988), T. rohtasfortai Khan andbaturensis, is reported from Gilgit Agency at lat. 36°

Tasnim, 1990 and T. fortmunroi Khan, 1993 are distinguished from the nominated species by their differsmaller
number ofbodyconfiguration,entinterorbitals,

subabdominal pholidosic counts, dorsal

1922; Gruber, 1981).

A highland Pakistani gecko, C.

20'N, long 74° 50' E (Khan and Baig, 1992). The
western most Pakistani form, C. walli, occurs between
lat.

35-36° N, long 71-72° E. while the western extral-

imital Iranian gecko,

C kirmanensis, is reported from30° N, long 58° E (Szczerbak and Golubev. 1986).

scalation and pattern, caudal morphology with keeledlat.

large tubercles, single row of subcaudals, presence ofWorldwide distribution of some geckos is largely
known to be due to human transportation (Darlington,1957). The circum-oceanic coastal forms are charac-

both preanal and femoral pores.

Following

Annandale(1935:39) groupedT.

(1913:316),

stoliczkai withSmith

low altitude T.

lawderanus (Stoliczka, 1972), despite very obvious
differences from the former: more flattened body andfew and feebly developed dorsal tubercles, incon-

tail,spicuous dorsal pattern, single pair of nasal scales,presence of preanal pores in male, tail morphology(compare Fig. 18, Smith, 1935).
C. yarkandensis (J. Anderson, 1872) and C. walli(Ingoldby 1922), have long been synonymized with
C. stoliczkai and C. chitralensis (Smith, 1935), mostly

due to their similar dorsal pattern, body configuration,
dorsal tuberculation and absence of pores (Boulenger.

1885; Blanford, 1878; Annandale, 1913; Smith, 1935;
Minton, 1966: Mertens, 1969: Szczerbak and Golubev, 1986). C. chitralensis has been found to be conspecific with C. walli (Khan, 1992).

Cyrtodactylus stoliczkai is defined by the following combination of characters: dorsal granular scales

teristically carried by sailors from coast to coast, how-ever,some of thespecies penetrated deep inland.

Similarly, westward dispersal of Cyrtodactylus in thelower Himalayas appears largely due to westward
migrations of Buddhist peoples and seasonal nomadic
high-low-highaltitudemigrations

which

playedimportant role in the distribution of cyrtodactylid geckos of the area. Massive pagoda buildings and temples
still the frequent haunts of these geckos. In thevalley-complex of the sub Himalayan system they
evolved into closely allied forms: C. mintoni, C. dat-aretanensis and C. battalensis.

A parallel example ofhuman transportation of a gecko is presented by the
recent record of Palearctic geckos. Hemidactylus persicus.

from Rohtas Fort, Jhelum valley. Punjab, Paki-

stan (Khan and Tasnim, 1990). The fort was built from1542 to 1550 with rock-blocks transported from
Balochistan. H. persicus is the dominant house-geckoinBalochistan and has never been reported from Pun-smooth, round, beady, juxtaposed, interspersed withjab (Khan 1987; Khan and Ahmed, 1987). The present

oblong, smooth beady tubercles, arranged roughly in
longitudinal rows; normal tail in juveniles uniformly

disjunct population most probably descended frombroad from basal stump till mid-tail, where graduallyobserved individuals are in severe competition withtapering to tip, as animal gets old, the anterior half ofthe original tail becomes broader, flatter and deeply

sected on sides in lateral lobules at segments, regenerated

tailmuch swollen and rounded; subcaudals

few individuals so transported. Presently the few
the common indigenous house gecko, H. flaviviridis,

which is dominant throughout the building. The few
individuals of H. persicus are still holding on inremote recesses of the fort.

small, in several rows; caudal tubercles thick andblunt.

Acknowledgments

Geographical Distribution

We wish to express our indebtedness to Dr. George R.

All evidence goes in favor of the idea that the family

Zug of NMNH for taking pholidosic counts and mea-

Gekkonidae evolved in southeast Asia and dispersed

surements on MCZ 7132 and to Dr. Michael Golebev.

USA for valuable discussions on

westward through southwestern Asia into Indo-Paki-

Seattle, Washington,stan and Africa (Kluge, 1967). Ranges of several circum-Indian oceanic cyrtodactylids overlap, however

the internet and translation of Russian texts.

circum-himalayan highland geckos are widely distributed and the ranges of none of them are known tooverlap: the northern most extralimital, Cyrtodacty-

Vol. 8, p. 67

Asiatic Herpetological Research

1999

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Appendix 2. Additional material examined

BMNH 1990.2; C. collegaBMNH 82.4.14.28-29; C. fasciolatus BMNH
1913.11.1 1.2; C. nebulosus BMNH 82.4.14.32-33; C.
oldhami BMNH 1916.6.22.4; C. pulchellus BMNH
1916.3.27.1-2; C.triedrus BMNH 68.3.17.11-12; C.
dattanensis MSK 0056.78: C. yarkandensis BMNH
Cyrtodactylus battalensis

lensis

72.3.22.4; C. tibetinus

Appendix 1. Abbreviations used
BMNH= British Museum, Natural History. London:

Gymnodactylus

CAS= California Academy of Sciences, California,

rensisBMNH 1946.8.23.19: Tenuidactylus batuBMNH 1990.3; longipes CAS 115944, SR
T.

USA;

MSK= Herp laboratory, 15/6 Darul Saddar

North,

Rabwah 35460,

T.

Austria;

Pakistan (author's personal

NMW= Naturhistorisches Museum Wien,

SR= Institute of Zoology, Academy of Sci-

ences, Kiev-30, Ukraine; UF= Florida State Museum,
Gainesville, USA;

USNM= National Museum of Nat-ural History, Washington, D.C.

CAS 196850, CAS 196854;

BMNH 1910.7.12.1; G. chi-tralensis

307:3267-68;

collection);

walli

T.

longipes voraginosus CAS 130323;

montiumsalsorum BMNH 1904.11.19.1 and MSK

indusoani MSK 0467.86;
mhtasfortai
USNM 284133; Gymnodactylus stoliczkai (photo-014.86;graph)T.

T.

NMW 16756; Tenuidactylus fedtschenkoi SR

1078:8837-8;

T.

caspius SR 2546:16713-14; T. turc-

menicus SR 961:8016-17.