Asiatic Herpetological Research

1999

Vol. 8, pp. 90-101

Two New Subspecies of Trapelus agilis Complex (Sauria: Agamidae) From
Lowland Southwestern Iran and Southeastern Pakistan
Nasrullah Rastegar-Pouyani
Department of Zoology, Gbteborg University, Box 463, SE 405 30, Goteborg, Sweden, e-mail:nasrallah.pouyani@zool.gu.se

Abstract.- Based on conducting long-term excursions and carrying out extensive field work in various parts of the

Iranian Plateau and studying preserved (museum) material including the syntypes of Olivier's Agama agilis, andparalectotypes of Boulenger's Agama isolepis, two new subspecies of the wide-ranging Asian ground agamid
Trapelus agilis complex are described from the lowland southwestern Iran and southeastern Pakistan (andadjoining northwestern India) respectively. The former of the new subspecies has conventionally been considered
as belonging to T. a. agilis, and the latter to T. a. isolepis. They are distinguishable from the other subspecies of T.
agilisdistinctive morphological characteristics. The distinctiveness of bothconfirmed according to the author's previous extensive studies on this complex using uni- andcomplex by having several

subspeciesismultivariate analyses of morphological characters. Both subspecies mainly occur as geographical isolates in the

periphery of the main range of the complex, and both have mainly been separated from the central continuum (=T. a. agilis) by eco-geographical barriers and are almost entirely restricted in distribution to the lowlands, desert

and semi-desert regions with high annual temperature. A taxonomic and biogeographic account as well as a key
to subspecies of T. agilis complex are presented.

Key words.- Trapelus agilis complex, T a. khuzistanensis, T a. pakistanensis. New subspecies, Agamidae,
Lowland southwestern Iran, Southeastern Pakistan, Systematics, Distribution, Biogeography.

Introduction
The,

taxonomically,

controversial

groundagamidTrapelus agilis is a wide-ranging species complex distributed from extreme southwestern Iran (ca 31° N,

47° E) to eastern Kazakhstan and western China (ca
48° N, 83° E) (Fig. 1 ), encompassing numerous local
populations (Rastegar-Pouyani, 1998). Traditionally,
this complex has been divided into three subspecies;

Trapelus agilis agilis (Olivier, 1804), T. a. sanguino-

T a. isolepis (Boulenger,
1885) (e.g., Anderson, 1974: Welch, 1983; Wettstein,

lentus (Pallas, 1814), and1951).

The

lattertwo subspecies are sometimes

regarded as full species by some authors (e.g., Ananjeva, 1981; Ananjeva and Tsaruk, 1987: Boulenger.
1885; Moody, 1980; Nikolsky, 1915; Zhao and Adler,1993). On the other hand, Anderson (in press) placesalldifferent forms ofinclusiveT.

agiliscomplex under thename "agilis" and believes that without afirm statistical ground, it is not advisable to divide thecomplex into separate taxonomic entities.
In a series of studies, using uni-and multivariate
statistics, the author analysed geographic variation inT.

agilis complex,

lenger) with T.

synonymized T. a.

a. agilis (Olivier),

isolepis(Bou-

designated a new

type locality as "terra typica designata" (central Iranian Plateau, about 1 10 km southeast of Esfahan city)

1999

Asiatic Herpetalogical ResearchVol. 8, p. 91

of adaptation of different populations to the local conditions as well as presence of eco-geographical barriers

which have been involved in differentiation andsubsequent evolution of all subspecies of this complex. Also I studied preserved materials including the
syntypes of Agama agilis Olivier. 1804 and paralecto885, borrowed
types of Agama isolepis Boulenger,1

museum collections around the worldfrom various(see under "Appendix I, Abbreviations, and Acknowl-

edgments").

my previous studies, based on examination of
about 1000 specimens of T. agilis all over the range, IIn

employed uni-and multivariate statistical techniquesand explored the patterns of geographic variation inmorphological characters within this complex (Rastegar-Pouyani, in press, unpublished manuscript).
Indeed, the present paper is the continuation (andFigure 2. Trapelus agilis khuzistanensis, holotype(GNHM Re. ex. 5424); A- dorsal view, B- ventral view.

part of the results) of

Note that almost all dorsal scales are distinctly small
and only slightly keeled.

plex according to which both the lowland southwest-

form (as the central core of the complex) are morpho-

populations do warrant taxonomic recognition.

logically

my previous studies concerningandtaxonomygeographic variation in T. agilis comern

Iranian

as

well

as

southeastern

Pakistani

most similar to the syntypes of Olivier's

Agama agilis.

Subspecies accounts

sanguinolentus (Pallas, 1814) is the northern
representative of the complex, distributed over a wide

Trapelus

area from northeastern Iran into the Central Asian(Figs. 2-6)

countries as far east as western China (Anderson, in

Khuzistan Ground Agama

T.a.

press;

Rastegar-Pouyani,

1998;

Zhao and

Adlcr,

1993). The two other taxa, described in this paper as

new subspecies, are geographically isolated, occurring in the southwestern and southeastern margins of
the main range of T. agilis, and mainly separated fromagiliskhuzistanensis ssp.

nov.1

GNHM

Holotype and type locality: adult female,Re. ex. 5224, collected by the author on 27 July 1996
from Iran, Khuzistan Province, 5
Haft-Gel on the road to Shushtar.

km northwest of

the central continuum (7^ a. agilis) by eco-geographi-

Paratypes: 14 specimens (ten males and four
females) have been designated as paratypes as fol-cal barriers. Both new subspecies arelows:morphologicallydifferent from the syntypes of Olivier's Agama agilis

based on several distinguishing characters (Table 1 ).

GNHM. Re. ex. 5225, same information as the
GNHM.

Re. ex. 5226, collected by theholotype:
author on July 28, 1996 from Iran, Khuzistan Prov-

name "isolepis" has beenince,

38 km south of Masjid-e-Suleiman, Golgir vil-synonymized with "agilis" and is no longer available(Rastegar-Pouyani, in press), new taxonomic names
are designated for these two new subspecies.

lage;

CAS 86342, 86346,

Since the subspecificSo, the main objective of this work is to describe

and introduce these two new taxonomic entities basedon the study of distinguishing morphological characterswhich make them recognizable from the other86390, Iran, Khuzistan

Province, Tul-i-Bazum [31° 55' N, 49° 25' E], about

500 m elevation, collected by S. C. Anderson on 17
1958; CAS 86403-6, Iran,
April and 22 May
Khuzistan Province, along road to lake east of HaftGel, about 300

m elevation, collected by S. C. Ander-

son on 23 May 1958; CAS 86418-19, Iran, Khuzistan

subspecies of T. agilis complex.

Province, along road between Haft-Gel and Masjid-Material and Methods

86556.

Suleiman, by S. C. Anderson on 23 May 1958: CAS
Iran,

Khuzistan Province, Haft-Gel (on golfcourse) [31° 28' N, 49° 30' E], about 300

conducted three long-term excursions and carriedout extensive field work in various parts of the IranianI

Plateau in 1995, 1996, and 1998, collected hundredsof specimens of Trapelus agilis and noted the patternm elevation,

by S. C. Anderson on 5 October 1958; CAS 86464,
Iran, Khuzistan Province, along road south of Shushtar,

byS.

C. Anderson on

18 July

170936, Iran. Khuzistan Province, 85

1958;

FMNH

km southeast of

Vol. 8. p. 92

Asiatic Herpetological Research

1999

Tablel legend. Main morphological differences between T. a. khuzistanensis, T. a. pakistanensis, and the Olivier's.

syntypes (T.a. agilis).
Characters

1999

Asiatic Herpetological ResearchVol. 8, p. 93

Figure 4. T. a. khuzistanensis, one of the maleparatypes (FMNH 170936). Note the presence of astrongly compressed
dorsal scalation.

tailand relatively heterogeneous

Figure 5. T. a. khuzistanensis, posterior ventral regionof a male paratype (GNHM. Re. ex. 5426) with one row
of callose preanal scales.

two paravertebral rows of smaller ones, proximal
upper caudal region with distinct dark-grey bars; ventral

surfaces uniformly whitish.

Measurements (mm): Total length = 206; Snout-vent= 86: Tail length (TL) = 120; Head
length (SVL)=
Head width (HW) = 19; Head depth(HL)
23;
length(HD) = 12.8; Length of forelimb = 45; Length ofhindlimb = 62.
Variation of the type series: all paratypes closely
approximate the holotype both in morphology andmeristics.

Figure 6. Habitat and type locality of T. a. khuzistanensis, 5 km northwest of Haft- Gel on the road to Shush-

The range of the number of scales around body for thewhole series (n = 14) is 82-97 and the mean 88.5; inall paratypes the number ofsubdigital lamellae under

Khuzistan province, southwestern Iran. In thethe fourth toe varies between 16-19; dorsal scales areforeground the author is attempting to capture the holotype from inside of its underground hole.

subequal or heterogeneous (especially in males); theretar,sals relatively larger,

grading into distinctly smallerscales of dorsolateral region

which are only slightly
keeled and mucronate; 94-95 scales round middle of

are

some

differences,

however, between male andfemale paratypes.
-

Male paratypes: all male paratypes (n = 10) arebody; scales of upper surface of limbs larger thanuniformly yellowish-grey-cream dorsally orwith a weakly developed pattern of dark cross bars; asmedian dorsals, distinctly keeled, slightly mucronate:

well, the ventral pattern is either uniformly whitish or

lower surface of digits covered by bi-or tri carinate

with distinct pattern of bluish-brown on the gularregion, chest and flanks; tail strongly compressed inlamellae, 18-19 under the fourth toe; ventral scales

eitheralmost as large as median dorsals, imbricate, veryslightly keeled or smooth, 84-85 scales in a single row

almost all males; preanal callose scales in one or two

from gular fold to the anterior edge of anus; callose

oped, their number varies from 10-21; dorsal scales
more keeled and mucronate and distinctly heteroge-

preanal scales in one row, slightly developed, consisting 10 scales; caudal scales larger than median dorsals,

strongly

keeled,distinctly

mucronate,

34-36

around base of tail just behind vent; tail weakly com-rows, in the latter case the second row weakly devel-

T. persicus in thisrespect; vensac
keeled,weaklygulardistinctly developed;
reverse imbrication of the posterior head and anterior

neous, approaching
trals

pressed at base, distinctly so towards the tip, its length
1 .39 ofbody length.

neck scales relatively more pronounced and the large

Coloration and color pattern: upper surface of headscalesand limbs uniformly yellowish-grey, ground color of

females.

dorsum yellowish-grey-cream with 5 large, broad,light-brown cross bars from nape to sacrum, inter--

rupted by a series of light, large vertebral ocelli andthe preanal pores are only in one row.

scale on posterior end of these reversally-imbricatedmoredistinctive

andmucronate

than

in

Female paratypes: all female paratypes (n = 4)
resemble the holotype in almost all pertinent details;
weakly devel-

Asiatic Herpetological ResearchVol. 8, p. 94

1999

Distribution: the main distributional range of thissubspecies is the Khuzistan Plain which is an extension of the Mesopotamian lowlands (Fig. 7). Also, it

penetrates into the western foothills of the Zagros
Mountains up to 900
elevation. It is the western-mmost representative of the wide-ranging T. agilis spe-

The Zagros Mountains serve

cies complex.

as anstrong barrier to its further eastward distribution, so it

has almost no contact with the central Iranian Plateau

nominal subspecies (T. a. agilis), except in the southeastern regions of Bushehr province, southern Iran

where
Figure 7. Geographic distribution of T. a. khuzistanen-

thetwo taxa occur as parapatric (Rastegar-

Pouyani, Manuscript, a). Its occurrence in the lowland
southeastern Iraq is almost unlikely and, so far, theresis (A) and T. a. pakistanensis (B). Squares represent

isthe type localities.

the Olivier's original record (1804) which is strongly

oped, each occupying the tip of a scale, their number

doubtful]. If it occurs in southeastern lowland regions

variesfrom 8

to

all

11;

scalesthe dorsal

weaklyround or weakly comindorsal pattern occur (indifferences
pressed; slightkeeled and mucronate;some specimens

the

tail

dorsal

cross

bars

more

are

intense).

Habitat:inlowland southwestern Iran and

in

the

western foothills of the Zagros Mountains, this sub-no proper record inside the Iraqi territory [exceptof Iraq, then the Tigris might have served as an effective barrier to its further westward distribution.

In some areas of lowland southwestern Iran (e.g., 85

km southeast of Ahvaz)

it occurs as
sympatric with
Trapelus persicus (Blanford, 1881) but there is noproper record of intergradation, if any, between the

two taxa.
sospecies mainly occurs on sand dunes, alluvial soils,

Etymology:

open plains, low hills, and on dry stream channels

named as it is mainly restricted in distribution to the(Anderson, 1966a,

lowlands of southwestern Iran, Khuzistan province.

in press: personal observations).

Trapelus

agiliskhuzistanensis

is

The type locality is specified by numerous low sanddunes as well as gypseous hills imminented with an

Trapelus

open plain. The vegetation is sparse, mainly Artemisia, Alhagi, Zygophylluni, and Euphorbia association.

(Figs. 8-9)

agilispakistanensis

ssp.

nov.2

Pakistan Ground Agama
Holotype and type locality: adult male, SMF 63258,

The climate conditions being harsh, with hot and longsummers (about 40°- 50°C) and mild, short winters. It

collected by M. G. Konieczny on 31 March 1957 fromseems that this lizard is active throughout the year but

Gaj-River, Kirthar Range, southeastern Pakistan.

Anderson first observed it in the western foothills of
the Zagros in early March, becoming numerous by
mid-April; both newly hatched and half-grown juve-

Paratypes (7 specimens): SMF 63259, 63279, same
information as the holotype; SMF 63256, collected by
R. Mertens on 1st December 1952 from Sonda, Distr.

niles were observed in late October and early Novem-

Thatta, southeastern Pakistan;ber (Anderson, 1963: 446).

7, collected

The holotype and one of the paratypes were collected
about 5km northwest of Haft-Gel on the road to

Shushtar. They were active when air temperature was

45°C and the substrate was 49.5°C. When alarmed,
unlike most of the central Plateau populations whichusually retreat into the base of dense bushes, they
retreated into the underground holes; this may be due
to vegetation scarcity.

inside their

Both specimens were captured

underground holes

(Fig.

6).

The

third

SMF 63243-4, 63286-

by M. G. Konieczny on 26 April 1961

from Old Airport of Karachi, southeastern Pakistan.
Diagnosis: Trapelus agilis pakistanensis differs fromthe other subspecies of T. agilis complex by having a

combination of distinctive characters; body and headsometimes compressed (not depressed) in males;
males almost always with one row of callose preanals
second undeveloped row may be present);
females without callose preanals: dorsal scales rela(rarely atively flat, subequal to homogeneous, distinctly keeled

specimen collected near the mouth of an old well inthe vicinity of the village of Golgir (38 km south of

throughout and mucronate, grading into small dorsolaterals rather abruptly (especially in males), 67-83

Masjid-e-Suleiman) while trying to retreat into thearound body;well.

keeled in males; body and limbs often strongly slen-

ventralscales

also

often

distinctly

der and head distinctly pointed (in adult males); tail

1999

Asiatic Herpetological ResearchVol. 8, p. 95

Figure 9. T. a. pakistanensis, holotype (SMF 63258);
posterior ventral region. Note the presence of only one
row of callose preanal scales and distinctly slender

hindlimbs.

imbricate,almost entire individual scale distinctly

keeled (unlike the eastern populations of T. a. agilis inwhich only the proximal part of scales are keeled) andmucronate; median dorsals

pakistanensis, holotype (SMF 63258); Adorsal region, B- ventral region. Note the presence of a

Figure 8.

T. a.

larger, grading into dissmallerscalesofdorso-lateraltinctlyregion which arestrongly compressed tail, and pointed and slenderslightly keeled and mucronate; 72-73 scales
round the widest part of body: ventral scales rather

body and head.

distinctly keeled, imbricate, large, but slightly smaller

often strongly compressed in adult males, its length
more than 1.55 of body length; the mean number ofsupra- and infralabials significantly lower than thoseof the other subspecies; a rudimentary nuchal crest
often present.

Description of holotype: an adult male, preserved in
70% ethyl alcohol in good condition; head pointed
with a slightly convex forehead, its length 1.36 of its

width and 0.27 of body length and 0.15 of tail length;
canthus rostralis continued as a supraciliary ridge
which is rather strongly developed, composed of 7
scales on each side; nostril as a horizontal slit, slightly

above canthus rostralis, pierced in a triangle-shaped
scale, posteriorly directed;

4 internasals

ina singletransverse row; upper head scales subequal. smooth,or slightly rugose, imbricate and subimbricate; 14-14

upper- and 13-13 lower labials; tympanum horizontally elliptical, smaller than orbit, partly covered

above by 6-8 spinose scales; no reversal imbricationof scales on the posterior part of head and anterior
part of neck; a rudimentary nuchal crest composed of

6-7 spinose scales; gular sac moderately developed
and rather pointed posteriorly; gular region coveredby large and small, imbricate, and slightly keeled
scales intermixed: gular fold and a fold in front of

only

than median dorsals, 70-71 in a single longitudinalrow from gular fold to the anterior edge of anus;
scales of upper surface of limbs almost as large as

median dorsals, strongly keeled and mucronate; lower
surface of digits covered by bi- and tri-carinate keeledlamellae, 24-25 under the fourth toe; preanal callose
scales only in one row, not exceptionally developed,

encompassing 8 scales; all caudal scales distinctly
keeled and larger than median dorsals, 33-35 aroundbase of tail, just behind the vent; tail long and strongly

compressed throughout, its length 1.81 times of bodylength.

Coloration and color pattern: upper surface of headyellowish-grey; dorsum uniformly sandy-grey withnumerous light scales scattered throughout; upper surfaces of limbs olive-brown; upper caudal region as

dorsum in coloration with barely distinct dark-brownrings; gular region, chest, and flanks heavily suffusedby lavender-blue; other ventral surfaces yellowishwhite; no black patch on the shoulder fold.

Measurements (mm): Snout- vent length (SVL) =101.6, Tail length (TL) = 184, Head length (HL) =28.2, Head width (HW) = 20.7, Head depth (HD) =13.5, Length of forelimb = 50, Length of hindlimb =73.

shoulder strongly developed; body slender and distinctly compressed laterally, a prominent vertebral

to the holotype both in morphology and meristics. In

ridge throughout the dorsum; limbs distinctly slender;alldorsal scales subequal to homogeneous, rather large,

tralisVariation of the type series: all paratypes are similar

specimens nostril is almost above the canthus rosand a rudimentary nuchal crest, more or less,Vol. 8, p. 96

Asiatic Herpetological Researchdeveloped. The range of scale counts around body for
the whole series (n = 7) is 69-81 (mean 74.42). The

1999

Distribution: Trapelus agilis pakistanensis, as thesoutheasternmost subspecies of T. agilis complex, isrange of ventral scales from gular fold to the anterior

restrictededge of anus (in a single longitudinal row) varies from67-77 (mean 74.57). In all paratypes the number ofdesert regions of Sind province, southern

subdigital lamellae under the fourth toe varies

from
22-26 (mean 23.85). The number of supra-and infra
labials varies

from 13-16 (mean 14.55) and 12-16

indistribution to thelowland and semiPunjab, andsome regions of eastern Baluchistan (southern andsoutheastern Pakistan), from around Hab River in thewest through Karachi and Thatta to the vicinity ofHyderabad and Mirpur Khas eastward into the Indian(mean 14.25) respectively. Dorsal scales being homo-

Desert (Fig.

geneous or subequal and, more or less, set off fromthis lizard inside the Indian
territory

the small dorsolaterals.

Kolayat, Pugal, Phalodi and some other localities in

However, there are some differences in morphology
between males and females:

distributed along the Kirthar

7).

Biswas and Sanyal (1977) recorded(from Jaisalmir,

Rajastan Desert, northwestern India); to the north, it isRange up to the areas

south of Khuzdar (south-central Pakistan). Apparently

Male paratypes: all male paratypes (n =4) resemble
the holotype in almost all relevant details; the mean
SVL = 90.5 mm, TL = 155 mm; mean TL 1.71 times
as mean SVL; the preanal callose scales almost onlytion of this taxon towards the west; however it isin one row (in the case of two, the second row
rarelygoes up to the
Nagaur District, Rajasthan, northwestern India.

-

developed), encompassing 10-11 scales (mean 10.4);

median dorsal scales enlarged, homogeneous, distinctly keeled throughout and mucronate, forming distinct ridges

along dorsum, grading, rather abruptly,
into small dorsolaterals which are weakly keeled andmucronate; all ventral scales, as in holotype, more orless keeled, rather large, but smaller than median dorsals;

strongly keeled in almost all specimens,
body not as strongly compressed andtailalthoughthe Hab River serves as a barrier for further distribu-

patric with

its

T.

a.

para-

agilis in the eastern regions

Baluchistan province. In the east,ofit

Etymology: Trapelus agilis pakistanensis is so named
it is restricted in distribution to the lowland andassemi-desert regions of southeastern Pakistan and adjacent northwestern India.

Taxonomic and biogeographic account
Detailed discussion concerning systematics and patterns of geographic variation in Trapelus agilis com-

elsewhere

pointed as in the holotype; color pattern almost as inplex

holotype.

Manuscript a-b) and here is not dealt with in details.
As a brief account, however, it can be mentioned that I-

Female paratypes: in females (n = 3), the body andare normal(neither distinctly compressed nor
slender), the dorsal scales are moderately keeled and
tail

is

presented

classified

allpopulations of Trapelus agilis complex

throughout the range into four distinctive taxonomic(and as the most parsimonious definition=
T. a.
agilis (Olivier, 1804), T. a. san-mucronate and median dorsals are not clearly set off

entitiesfrom the small dorsolaterals; as well, ventrals areslightly keeled and distinctly smaller than the mediansubspecies):

dorsals; callose preanal scales absent; gular sac not as(Rastegar-Pouyani,

guinolentus
ssp.nov. 1,(Pallas,1814),

Ta.

khuzistanensis

and T. a. pakistanensis ssp.nov.2.

well developed as in the holotype; in color pattern

An ANOVA-based pairwise comparison showedsome of them are rather different from the holotype in
the presence of, more or less pronounced, dark crossthat in most metric and meristic characters these fourtaxonomic entities are significantly different (P<0.05)

bars and a series of vertebral light ocelli and in the(Rastegar-Pouyani, Manuscript,

absence of ornamentation in the lower parts of body;multivariate statistical techniques (principal

ventral surfaces being uniformly whitish.

nent analysis, canonical variate analysis, and cluster

Habitat: the habitat of this subspecies is characterised by flat alluvial plains as well as some highslopes.

The

vegetation consists of some grass, herbs, andstunted shrubs. Some populations occur on the low-

lands around Karachi, and the eastern part of therange is a typical desert known as Thar Desert in east-em Sind extending into adjacent northwestern India(the great Indian Desert). This desert mainly consistsof sandy

hillswith 100-150
the

main

which vary from small dunes to hills

m elevation. In summer, dust storms arefeature

of

the

area

(Khan,

1980).

a).

Also employing

compo-

analysis), to a great extent, showed the objectivity ofthese four distinct groups within T agilis complex andre -confirmed

my previous taxonomic decisions (Ras-

tegar-Pouyani, Manuscript, b). Of the four subspeciesof T. agilis, the nominal form (T. a. agilis) occurs in awide range of habitats, shows higher degree of morphological variability, occupies the central and southern parts of the species range (these regions might

have served as a refugia during intervals of unfavorable and cold climatic conditions in the Tertiary and
Quaternary), and consists of two western and easternAsiatic Herpetological Research

1999

Vol. 8, p. 97

inrevisionary study has been done on all species of thisWith regard to thesefactors, it could be logical if we consider this form as

genus and this is mainly because of political instability of the region and difficulties in collecting propercomplex and as the parental

material throughout the range. Furthermore, even in agroups of populations (clines) (Rastcgar-Pouyani,press, unpublished manuseript).

the central core of the

population from which the other subspecies have beenfew studies done on a limited number of Trapelus spe-

derived. So, preliminarily,

propose the following
scenario for origination and subsequent evolution ofcies, the results

marginal subspecies which occur in thethe main range of the complex beingof
peripherywiththe central continuum (=T. a. agilis):
parapatricdictory (e.g., Anderson, in press; Joger and Arano,Ithe threeobtained by morphological and theother approaches (e.g., immunological) were contra-

1987; Rastegar-Pouyani, in press, unpublished manuscript).

seems that T. a. sanguinolentus, in spite of being

Trapelus is mainly Saharo-Sindian in distribution,

distributed over a very wide area, is the most recently-

often associated to the lowlands, desert, and semi-

evolved group, originated from the parental populations in the southern parts of the range (apparently) in

desert regions with high annual temperature.

Itthe very Late Pliocene and Pleistocene (2-1.2

MYBP

Key to the subspecies of Trapelus agilis com-=millions of years before present), invading towards

plex

the northern and northeastern regions during intervalsof favorable climatic periods. The very low degree ofvariability observed in this subspecies is indicative of

callose preanal scales

its

recent history. Trapelus a. pakistanensis separated

from the central continuum in the southeastern partsof the range (probably by dispersal or, less likely, due
to a vicariant event) distributed

invasion might have taken place in the Pliocene (5-1.7

MYBP). In the southwestern part of the range, a drasvicariant event (thehuge orogeny of the Zagros)

separated the southwesternmost populations from restof the complex and from the parental populations.
These isolated populations served as founders and,with further divergence, gave rise to T. a. khusistanensis. Apparently, thisthe Late

lb. Tail2

often compressed, often only one row of cal-lose preanal scales, in the case of two, thesecondundeveloped (preanals absent or weakly developed in3females)

towards the east andreached as far east as northwestern Indian desert. Thistic

almost always round, two (or more) rows of

la. Tailvicariant event has taken place in

Miocene or early Pliocene (7-4.5 MYBP).

2a.

Body size variable; 65-91

scales around body;dorsal scales subequal, weakly to moderately keeled,often strongly mucronate; ventral scales

smooth or

weakly keeled; usually 2, sometimes 3 (rarely 4-5)
rows of callose preanals; background coloration variIraniancentralable;

Plateau, central

Afghanistan, southwestern Pakistan.

.

and southern
Trapelus agilis

agilis (Olivier, 1804).
2b.

Body stout; 52-73 scales around body; all dorsal,

ventral, and gular scales larger in size, homogeneous,

keeled andmucronate; almost always 2

Therefore, we can say that both dispersal and vicari-strongly

ance have been involved in radiation and subsequent(rarely 3) rows of callose preanals; background color-evolution of various subspecies of Trapelus agilis
complex though the role of dispersal in evolution of T.

ation of males often dark sandy-grey; northeastern

Afghanistan, Central Asian Republics,
western coast of the Caspian Sea (Daghestan), westIran, northernsanguinolentus and T. a. pakistanensis, and vicariance in evolution of T. a. khuzistanensis are more

ern China.

prominent.

1814).

Trapelus agilis (Olivier, 1804) is the easternmost
representative of an essentially homogeneous and3a.

similar group of about five species complexes whichpressed; head and neck distinctly short; all bodyscales smaller than those of the other subspecies; dor-a.

also include T persicus (Blanford, 1881 ) in the Meso-

.

.

.

Trapelus agilis sanguinolentus (Pallas,

Body and limbs smaller than those of the othersubspecies and sometimes relatively slender, not com-scalestopotamian Plain and lowland southwestern Iran, T. flavimaculatus Riippell, 1835 in Saudi Arabia, Tsalsavignii (Dumeril and Bibron,

97 scales around body; ventral scales slightly keeled;

1837)

in

Israelandsubequalunequalweakly to moderately keeled,(heterogeneous),

weakly mucronate; 80and anterior part ofeastern Egypt, and T. tumevillei (Lataste, 1880) in
north Africa (north of Sahara). Although the mono-scales of posterior part of head

phyly of Trapelus has been shown by Moody (1980)
with a morphological approach and by Joger (1991or rugose; often 14-19 upper- and lower labials; back-neck reversally imbricated; upper head scales keeledusing molecules (neither Moody nor Joger studied all

ground coloration often yellowish grey-cream; lowland southwestern Iran (0-900 m elevation)

the species of Trapelus) but, so far, no comprehensive

Trapelus agilis khuzistanensis, ssp. nov. 1)

Vol. 8, p. 98

Asiatic Herpetological Research

1999

Body and limbs often distinctly slender, some-und Museum Alexander Koenig; Dr. Gunther Kohler,

times compressed in males; dorsal scales subequal to

Forschungsinstitut Senckenberg; Dr. Marta Poggesi,
Museu Zoologico de "La Specola"; Dr. Ivan Ineich,

3b.

homogeneous, distinctly keeled and mucronate, usufrom small dorsolaterals; 67-83

ally clearly set off

scales around body; ventrals distinctly keeled in adultmales; no reversal

imbrication of head andneckMuseum National d'Histoire Naturelle; and Dr. Olavi
Gronwall and S. O. Kullander, Swedish Museum of
Natural History, for loan of Trapelus specimens. Dr.

scales; upper head scales often smooth; 12-16 upper-

N. B. Ananjeva, Zoologicaland lower labials; background coloration often sandygrey: southeastern Pakistan and adjoining northwest-

emy of Sciences, St. Petersburg, who made it possible
for me to visit the herpetological collections of Zoo-em Indialogical Institute and examine specimens of T. agilis.

.

.

.

Trapelus agilis pakistanensis, ssp. nov.2

Institute,

Russian Acad-

This work was partly supported by grants from theAbbreviations

Royal Swedish Academy of Sciences Hiertza-RetAnna Ahrenberg, and Wilhelm Martina(

BMNH = British Museum (Natural History; (London,
UK), CAS = California Academy of Sciences (SanFrancisco, USA), FMNH = Field Museum of Natural
History (Chicago, USA), GNHM = Goteborg Natural
History

MNHN =

Museum(Goteborg. Sweden),Nationald'Histoire
Naturelle

Museum(Paris,France), MZLS = Museo Zoologico de "La Specola"

NMW = Naturhistorisches Museum(Firenze, Italy),

Wien (Vienna, Austria). SMF = Museum und Forschungsinstitut(Frankfurt,

Senckenberg

Germany),

SMNH = Swedish Museum of Natural
ZFMK =

History
Zoologisches Fors-zius Fond),

Foundations.

Literature Cited
Ananjeva, N. B.

1981. structural characteristics ofskull, dentition,and hyoid of lizards of the genus

Agama from the fauna of the USSR. Pages 3-20. In:'The Fauna and Ecology of Amphibians and Reptilesof the Palearctic Asia". Proc. Zool. Inst. USSR Acad.
Sci. 101.

chungsinstitut und Museum Alexander Koenig (Bonn,

Ananjeva, N. B., and O. J. Tsaruk. 1987. The taxonomic status of the steppe agama, Trapelus sanguino-

Germany); ZISP = Zoological Institute St. Petersburg

lenta in the Praecaucasus. Proc. Zool. Inst. Leningrad.

(Stockholm, Sweden),(St.

Petersburg,

MuseumZMUC

Russia),Universityof Copenhagen=

Zoological(Copenhagen,Denmark).

158: 39-46.

Anderson, S. C. 1963. Amphibians and Reptiles from
Iran. Proc. Calif. Acad. Sci.. 31(16): 417-498.

Anderson,

Acknowledgments

C.

S.

1966a.

The Turtles,

Lizards, andwould like to thank my supervisor. Dr. Goran Nil-

Amphisbaenians of Iran. Ph.D. thesis. Stanford University. 660 pp.

son, for all his help and cooperation as well as for crit-Anderson, S. C. 1974. Preliminary key to the Turtles,

I

ically reviewing this paper.

Iwish to thank the Razi

University authorities (Kermanshah-Iran) as well as
Eskandar Rastegar-Pouyani, Alireza Hashemi, Seyful-

Masoud Abrishamian and Ali Rassuli for
their unsparing help during field work on the Iranianlah Azadi,

Plateau.

I

am also grateful to Professor S. C. Ander-son, Biological Science Department, University of the
Pacific, Stockton, California, for all his recommendations,encouragements, and for sending informative

Lizards, and Amphisbaenians of Iran. Fieldiana: Zool.
65: 27-44.

Anderson, S. C. (in press). The Lizards of Iran. Society for the study of Amphibians and Reptiles, Oxford,OH.
Biswas, S. and D. P. Sanyal. 1977. Fauna of Rajastan,
India. Part: Reptilia. Rec. Zool. Surv. India, pp. 254-

255.

literature, especially his unpublished Manuscript con-Blanford, W.

cerning the Iranian agamids.

tiles

My special thanks also go to: Dr. Alan E. Leviton,
Department of Herpetology, California Academy of
Sciences; Dr. Colin McCarthy, Department of Zoology, British Museum (Natural History): Dr. Harold K.

Department of Herpetology, Field Museum of
Natural History; Dr. Jens B. Rasmussen, ZoologicalMuseum, University of Copenhagen: Dr. Franz TiedeVoris,

mann, Naturhistorisches Museum Wien; Professor
Wolfgang Bohme, Zoologisches Forschungsinstitut

T 1881. On a collection of Persian rep-

recently added to the British

Museum.

Proc.

Zool. Soc. London, pp.67 1-682.

Boulenger. G. A. 1885. Catalogue of the lizards in
British Museum. 1: 334-369.

Dumeril, A. M. C, and G. Bibron. (1834-1854). Erpetologie generaleouhistoire naturelle

complete desreptiles. Paris, 8 Vols.

Joger, U. 1991.

A molecular phylogeny of agamid liz-ards. Copeia 1991(3): 616-622.

1999

Asiatic Herpetological Research

Joger, U., and B. Arano. 1987.

Biochemical phylog-

eny of the Agama genus group. In: Van Gelder, Strijhosch, and Berger (eds.). Proc. 4th Ord. Gen. Meet.

1980. Affinities and Zoogeography of
BIOLOGIA, 26(1-2): 113-

S.

Herpetiles of Pakistan.
171.

Lataste, F.

Iraq, vicinity of Baghdad. GNHM. Re. ex. 5213-17: Iran,
Tehran prov. 50 km NE Qum on road to Tehran. GNHM.

Re. ex. 5218-28: Iran, Tehran prov. 75 km NE Qum on road

GNHM. Re. ex. 5229-34: Iran, Tehran prov. 5 km
NE Saveh on road to Tehran. GNHM. Re. ex. 5235-62: Iran,
Tehran prov. (50-60 km W Tehran) between Eshtehard
Saveh. GNHM. Re. ex. 5263: Iran, Tehran prov. 30 km SE
Tehran on road to Garmsar. GNHM. Re. ex. 5264-66: Iran,Central prov. 25km Khomain on road to Mahalat. GNHM.
to Tehran.

Soc. Europ. Herp., Nijmegen, 215-218.

Khan. M.

Vol. 8, p. 99

1880. Diagnoses des reptiles nouveaux-

Re. ex. 5267-81: Iran, Central prov. between Komain-Deli-d'Algerie. Naturaliste 1, pp 299-325.

Moody, M. S. 1980. Phylogenetic and Historical Biogeographical Relationships of the Genera in the Family Agamidae (Reptilia: Lacertilia). Ph.D. Thesis.

jan and Mahalat-Delijan roads. GNHM. Re. ex. 5282-86:
Iran, Esfahan prov.near Naragh, 50km

W kashan on road toNaragh. GNHM. Re. ex. 5287-89: Iran, Esfahan prov. 20
km ES Kashan on road to Natanz on the margin of Dasht-e

University of Michigan. Ann Arbor. 373 pp.

Kavir.GNHM. Re. ex. 5290-92: Iran, Esfahan prov. 80km N

Nikolsky, A. M. 1915. Fauna of Russia and adjacentcountries (Translated from Russian by Israel program

Esfahan on road to Natanz.GNHM. Re. ex. 5293-97: Iran,for scientific translations. 1963). Vol.

1.

Chelonia andSauria. 352 pp.

l'Empire Ottoman,l'Egypte et al Perse. Agasse, Paris. Vol. 4.
P.

payeh city. GNHM. Re. ex. 5298-99: Iran, Esfahan prov.
15km N Shahreza on road to Esfahan. GNHM. Re. ex.
5300-07: Iran, Fars prov. 50 km N Abadeh on road to Shahr-

Olivier. G. A. 1804. Voyage dansPallas.

Esfahan prov. 70km E Esfahan on road to Naein, near Kuh-

S. 1814.

Zoogeographia rosso-asiatica, sis-eza.

GNHM. Re. ex. 5308-16: Iran, Fars prov. 50- 75km S

Abadeh on road to Shiraz. GNHM. Re. ex. 5317-31: Iran,Markazi province, 45 km E Arak on the road to Qum.

Rastegar-Pouyani, N. 1998. Systematica and Distribution of the Iranian Species of Trapelus (Sauria: Aga-

GNHM. Re. ex. 5332-34: Iran.Tehran prov., 65 km NE
Saveh on road to Robat-Karim. GNHM. Re. ex. 5335: Iran,Markazi prov.. 50 km N of Delijan on road to Salafchegan.
GNHM. Re. ex. 5336-44: Iran, Esfahan prov., 45 km E of
Golpaygan. GNHM. Re. ex. 5346-50: Iran, Esfahan prov., 7
km E of Tiran on road to Najaf-Abad. GNHM. Re. ex.

A Review. Russian Journal of Herpetology

5351-57: Iran, Esfahan prov., 25 km SE of Esfahan on roadtens

omnium animalium extenso imperio rossico etadjacentibus maribus observatorum recensionent. 3,
Petropoli, Acad. Scient. 428 pp.

midae):to Dastjerd.

5(2): 127-146.

Rastegar-Pouyani, N. (In press). Analysis of Geo-

GNHM. Re. ex. 5358-68: Iran, Esfahan prov.,

about 1 10 km SE of Esfahan city, on road from Malvajerd to

Ramsheh. GNHM. Re. ex. 5369-70: Iran, Esfahan prov., 20

graphic Variation in Trapelus agilis complex (Sauria:

km W of Ramsheh on road to Shahreza. GNHM. Re. ex.

Agamidae). Zoology in the Middle East.

5371-78: Iran, Fars prov., 50 km N of Abadeh (15 km S of

Rastegar-Pouyani, N. (Unpublished) Taxonomic ReEvaluation of Trapelus agilis complex (Sauria: Aga-

Izad-Khast) on road to Esfahan. GNHM. Re. ex. 5379-91:

midae): A Multivariate Approach.
Ruppell, E. 1835. Neue Wirbeltiere zu der Fauna vonAbyssinienIII.

gehorig.

Amphibien.

SiegmundSchmerber, Frankfurt-am-Main. (2)+18p., 6 pis.

Welch, K. R. G. 1983. Herpetology of Europe andSouthwest Asia, a checklist and bibliography of theorders Amphisbaenia, Sauria and Serpentes. Robert E.

Krieger publishing Co., Malabar, Florida, viii + 135
pp.

Wettstein, O. 1951. Ergebnisse der Osterreichischen
Iran- Expedition

1949 / 1950, Amphibien und Rep160: 427-448.

tilien. Sitzb. Akad. Wiss. Wien.

Zhao. E., and K. Adler. 1993. Herpetology of China.
Soc. Stud. Amph. Rep. pp. 186-187, 288-289, 304.

Appendix 1. Material examined

Iran. Fars prov., 65 km SE of Abadeh on road to Shiraz.
GNHM. Re. ex. 5399: Iran. Kerman prov., 28 km NE of Sirjan on road to Kerman. GNHM. RE. ex. 5400-1: Iran, Kerman prov., 70 km NE of Sirjan on road to Kerman. KhanehSorkh Pass, about 2800 m elevation. GNHM. Re. ex. 5402-

13: Iran, Semnan Prov., 5o

km E of Semnan, North of

Dasht-e- Kavir, Northeastern Iran. GNHM. Re. ex. 5414-23:
Iran, Khorasan Prov.,

1

10 km East of Shahrud, Northeastern

CAS 142230: Iran.Tehran prov. Lar-Damavand.
CAS142189, CAS 142190, CAS 142182, CAS 142192,
CAS 142191, CAS 142187, CAS 142188, CAS 142183,
CAS 142184, CAS 142186, CAS 142185: Iran, Zanjan
prov. Ghazwin. CAS 142231: Iran.Tehran prov. Karadj.
CAS 142093, CAS 142094, CAS 142098, CAS 142097,
CAS 142100. CAS 142092, CAS 142096, CAS 142095,
CAS 142101, CAS 142099: Iran.Tehran prov. Saveh. CAS
Iran.

141293: Iran, Tehran prov.24km

W of Saveh On road toHamadan. [35 07 N, 50 08 E]. CAS 142194, CAS 142197,

CAS 142195, CAS 142199, CAS 142198, CAS 142193:
Iran. Esfahan prov. Mahalat. CAS 141019: Iran, Esfahanprov.6km by road NW Ardestan. [32 26 N, 52 20 E]. CAS

Trapelus agilis agilis (n =541 )141 109: Iran, Esfahan prov. 8km S of Kashan and 6km

MNHN 5708 (2) = 1994.

road between Yazd -Kashan. [33 54 N, 51 30 E].

1178 (2) (Olivier's syntypes):

W of

FMNH

170987.

Asiatic Herpetological Research

100Vol. 8,FMNH 170988: Iran, Gilan prov. 4,2 mi N of Tuke-NMW 33 166-1, Iran: Fars
prov. 9 km N of Abadeh. NMW 33167, NMW 33085-2:
Iran, Hormozgan prov. 14km E Bandar-Abbas. NMW 7276NMW 7276-2: Iran: Kerman prov.? Sabzewaran. NMW
stan (Zanjan prov. Takestan?).

I,7276-3: Iran: Kerman prov.? Rigmati. 150 km SE of Sabze-waran.

NMW 7276-4: Iran, Esfahan prov. 135 km N of
NMW 33159-3, NMW 33159-4, NMW 33159-5,

Esfahan.

NMW 33165-1: Iran, Esfahan prov. Murcheh-Khurt, N of
NMW 33165-2, NMW 33165-3, NMW 33085-1:

Esfahan.

Iran: Esfahan prov. 4

km SE Robai-Tork

165 km(NW of

NMW 33085-3: Iran. Zanjan prov. 26 km SE
NMW 33083-1, NMW 33083-2, NMW 33083-3:
Iran, Zanjan prov. 41 km SE Ghazwin. Elev: 1380 m. NMW
33164: Iran/Tehran prov. 60 km S of Tehran. NMW 33084Esfahan).

Ghazwin.

3: Iran, East Azarbaijan prov. 42

km SE Mianeh. Elev: 1200

1999

NMW

33148-1-10:
IRAN.Tehran prov. 95 km S Tehran.
IRAN: Tehran prov. Zaveyeh, 80 km SW Tehran.
7276-5-6: IRAN, Kerman prov. Sabzewaran.
33175-NMW
NMW

NMW 33170-5, NMW 33170-6. NMW 33170-7: Iran.
NMW 33172-3-4: IRAN: Kerman prov. 138
km S Rafsanjan. NMW 33172-5: IRAN: Kerman prov. 29
km SE Sirjan. NMW 33 165-4: IRAN, Kerman prov. 10 km
SW Kerman. NMW 33172-2, NMW 33175-6, NMW
33175-7: Iran: Yazd prov. 200km SE Yazd. NMW 33174-1IRAN, Khorasan prov. 15-25 km S Qayen. NMW 33163:
Pakistan: Nepandgur (where?). NMW 33162: Pakistan:
Delbandin. NMW 33161-1-2: Iran, Shirgesht Beiteabas.
NMW 33161-3: Iran, eastern Iran, Ozbak-Kuh. NMW
5,

Esfahan prov.

1

8:33172-6: Pakistan. Baluchestan prov. 50 km

W Nushki.

NMW 33150-9: Iran W Sangbast. NMW 24763-1 -3 :Iran, NPersian. Kuh Daschteh (Taj-abad). NMW 24767: Iraq.Bagh-NMW 33166-1: Iran. Kerman prov. 15 km SE Kerman.

dad? (Paris Museum). CAS 120280-1: Afghanistan, 10km

SMNH 3139-1, SMNH 3139-2, SMNH 3139-3, SMNH
3139-4, SMNH 3139-5, BMNH 85. 5. 27. 14, BMNH 85. 5.
27. 15. BMNH 85. 5. 27. 16. BMNH 85. 5. 27. 17, BMNH
85. 5. 27. 18, BMNH 85. 5. 27. 19: Iran: Tehran (exact

NE Darweshan (Central Afg.). CAS 84642-45: Afghanistan,m.

locality?). GNHM. Re.ex. 4437, GNHM. Re.ex. 3324,
GNHM. Re.ex. 4395-1: Iran, Gilan prov. 65 km SSW of

GNHM Re.ex. 4395-2: Iran, Tehran prov. 150 km S
of Tehran. Siah-Kuh. Shah- Abbas post. GNHM Re.ex.

35 mil. down stream from Girishk. Dasht-e-Margo area,
Chah-e-Angir, (Central Afg.). CAS 120242-4 Afghanistan,
12 km S Lashkargah (near Girishk), 2700ft. CAS 97990:
Afghanistan. 20 mil. SE Kandahar, 31 23 N, 65 53 E, 3800

CAS 90762-9: Afghanistan, Sharisafa. 60 km NE Kanda1400m. CAS 90777: Afghanistan,Tarnak river, 75 km
NE Kandahar. 1405m. CAS 120276-9: Afghanistan, 30km Sft.

Rasht.

har.

4396-1, GNHM. Re.ex. 4396-2: Iran, Mazandaran prov. 150

Ghazni-Qalat 7100ft. FMNH 20987-1-10: Iran.Esfahanprov. Yazd-e-Khast .FMNH 20985-90: Iran, Esfahan prov.

km E of Gorgan, Golestan Park. La Specola 30584, La Specola 30585: Iran/Tehran prov. 50 km S Qum? BMNH 1920.
Hamadan prov. Hamadan. Jinjan? BMNH
1936. 10. 12. 1: Iran. Kerman prov. N. Of Kerman, Sekonj.

FMNH 20988-1-2: Iran,Daria-Masila. FMNH 245507-10:

3. 20. 1: Iran,Pakistan.Baluchistan prov. SMF 63226-37, SMF 63262,

7000 ft. BMNH 1934. 12. 16. 3. BMNH 1936. 12. 16. 4:

din.

Iran. Sheakuh, seat (Salt?) desert.

1919.5.2.2-3: Iran, Fars prov. Abadeh.

BMNH 1912. 3. 26. 13, BMNH 1912. 3. 26. 14, BMNH
1912. 3. 26. 15, BMNH 1906. 8. 10. 25, BMNH 1900. 5. 9.
9, BMNH 1900. 5. 9. 10, BMNH 1900. 5. 9. 11, BMNH
1900.5.9. 12, BMNH 74. 11.23. 104, BMNH 74. 11.23.
23. 106. BMNH 74.
23. 107,105, BMNH 74.
BMNH 74. 23. 108, BMNH 74. 23. 109, BMNH 74.
1

1

1.

1.

1

1

1.

1.

11.23. 110.BMNH74. 11.23. 111.BMNH74. 11.23. 112,

BMNH 74.
23.
13: Iran, Sistan-Baluchestan prov.
BMNH 1951. 1.6. 50, BMNH 1951. 6. 51-2. BMNH 94.
II. 13. 4, BMNH 94. 11.13. 5: Iran, Hormozgan prov.
BMNH 87. 12. 20. Iran, Hormozgan prov. Kishim(=Gheshm?) Island, Persian Gulf. BMNH 79. 8. 15. 10-15:
Iran, Fars prov. Dehbid, north of Shiraz. BMNH 1903. 3.
14.
20Iran, Bushehrprov. Bushehr. BMNH 1933.4.
22: Pakistan, Waziristan, N. W. F. P. BMNH 86. 9. 21. 17.
BMNH 86. 9. 21. 18, BMNH 86. 9. 21. 23, BMNH 86. 9.
21. 24.BMNH 86. 9. 21. 25: Afghanistan: Helmand. ZMUC
R36133, ZMUC R36149-54, ZMUC R36160, ZMUC
1

1.

1

1.

1:

1:

1.

SMF 63285, SMF 63255: W-Pakistan, Siah-Kuh, S Delban-

BMNH 1951.1.6.54: Iran, Bandar-e-Lengeh. BMNH
BMNH

1936.10.12.3: Iran, Fars prov.between Quatru-Chah Salz to
Neiriz road.

BMNH 1966-355-57: Iran, Kerman prov. 148

km E Neiriz on Zaidabad road to Sirjan. BMNH
1951.1.2.20-22: Iran, Kerman prov. 20 mil. S of Kerman.

BMNH 1951.1.6.45-48: Iran, Sistan prov. Khash, SE
BMNH 1940.3.1.19-24: Afghanistan, Ghazni, (EJupar.
Iran.

Afgh.l.CAS 141028: Iran, Kerman prov. 17km SSE Minabon inland road to Jask. CAS 141020: Iran, Kerman prov. 13

km E of Eastern edge of city of Kerman. CAS 141051: Iran.
Kerman prov. 19km SE Shagu on road to Minab. CAS
141027: Iran, Kerman prov. 21 km N Rudan on road to
Jiroft. CAS 141097: Iran, Sistan prov. 10km SW Hirmand,
abandoned village, SE of road from Zabol to Dust-e-

Mohammad Khan. CAS 141065: Iran, Baluchistan
prov. 3km southerly of Zahedan on road to Khash. CAS
1102484-90: Iran: Sistan prov. 15 mi

SW Zabol. CAS

102491-2: Iran, Fars prov. Ahram. CAS 141 149: Iran, Fars
prov. 5km northerly from Dalaky on road to Shiraz where

CAS 96270: Iran. Khorasan prov. Tayebat.

R36204-5: Afghanistan: Seistan prov. Faisabad (south offoothills begin.

ZMUC R36155-9, ZMUC R36145-8,
SeistanAfghanistan,prov. Baqrabad. ZMUC R36161:
Afghanistan, Seistan prov. Faisabad-Farah. ZMUC R36206:

about 10 mi from Afghanistan border.

Afghanistan).

Afghanistan, Kandahar prov. Przadah,

W of Kandahar.

Iran. Khorasan pro. 5

NMW 33150-1-6:

km N Taybad. NMW 33166-1

:

Iran:

Kerman prov. 15 km SE Kerman.

ZMUC R36208-9: Afghanistan, Kabul prov. near Kabul?

Trapelus agilis sanguinolentus (n =238)

ZISP 7361: Afghanistan: Harat.

GNHM Re.ex. 4396 l-2):Iran: Mazandaran prov. 150 km E

NMW 33170-12: IRAN.
East Azarbaijan, 42 km SE Mianeh. NMW 33173-1-2:
IRAN, Zanjan prov. 26 km SE Qazwin. NMW 33170-1:(

of Gorgan, Golestan Park. GFN 40: Turkmenistan, Lowland
steppe. East Kopet Dagh.

GNHM. Re. ex. GFN 41: Turk-

1999

Asiatic Herpetological Researchmcnistan. Murgub River, 150 km S of Marv (Mary).

GNHM. Re. ex. GFN 42-50:Turkmenistan, Sahra-BairumAli. 45km NW of Marv, Kara kum desert. CAS 185 104-9.
CAS 185134-5: Turkmenistan, Ashgabad region [38 00 N,
58 00 E], CAS 184570-6: Turkmenistan: Krasnovodsk
region [39 45 N. 54 33 EJ.

GNHM. Re. ex. GFN 35: Turkmenistan. Archenjan village.
GNHM. Re. ex. GFN 36: Turkmenistan. Krasnovodsk
region [39 45 N, 54 33 E]. GNHM. Re. ex. GFN 43 (1-2):
Turkmenistan, Kaka.

GNHM Re. Ex 10622-3: Kazakhstan,at the lake (artificial ) at Illi River, near village Bokter [43 54

Vol. 8. p. 101

pros.. 38 km S of Masjid-e-Sulaiman, Golgir village. CAS
86403-6, 86418-19 (paratypes), Iran, Khuzistan prov.
:

along road to lake east of Haft-Kel. CAS 86464 (paratype):
Iran, Khuzistan an prov. along road south of Shushtar. CAS

86556 (paratype): Iran. Khuzistan prov. Haft-Kel (on golfcourse) [31 28 N. 49 30 E]. CAS 86342, 86346, 86390(paratypes): Iran. Khuzistan prov. Tuli-Bazum road [31 55

N. 49 25 E], FMNH 170936 (paratype): Iran, Khuzistan

CAS 86341, 86343-72,
86374-89: Iran, Khuzistan prov. Tuli-Bazum road [31 55 N.
49 25 E]. CAS 86320. 86323-28: Iran, Khuzistan prov.
prov. 53 mi SE Ahwaz , Mashrageh.

stan, Illi River, near border of China. ZISP 5796: Kazakh-

Masjid-Suleiman [31 57,N, 49 16,E]. CAS 86322: Iran,
Binak. near Persian Gulf at foot of Kuh-e-Bang [29 44,N,

stan.Tardski, Dshungaria, between Kazakhstan-China. ZISP

50 19,E]. CAS 86338-51: Iran, Khuzistan prov. along old

20298 (7745-79): Kazakhstan, near Ilisk city, Illi River. E.
Kazakhstan. ZISP 15143, 17329, 13695, 1 168: Kazakhstan,

Masjid-Suleiman and Ahvaz road [32 N, 49 1 l.E]. CAS
86251: Iran. Khuzistan prov. Masjid-i-Suleiman [31 57 N,

near Ilisk city, Illi River, E. Kazakhstan. ZISP 1 1070

49 16 E], CAS 86373: Iran. Khuzistan prov. road south ofN, 77 16 E]. ZISP5109, 13701-15 Kazakhstan, E. Kazakh-

Kazakhstan:

,

Illi

(

1

-

1

2 ):

River, E. Kazakhstan.close to lake

Shushtar. CAS 102491-2: Iran, Fars prov. Ahram. CAS

Balkhash. ZISP 19115 (7643-84): Uzbekistan, Vicinity of

86625: Iran. Khuzistan an prov. Binak. on Persian Gulf at

Nucus city. Caracal, near Aral sea. ZISP 19398 (7428-48):
Uzbekistan-Tajikestan, Fergan Valley. ZISP 15803 (1-15):

CAS 86487: Iran, Fars prov. Agha Jari. [29 48 N, 49 46 E].

Tajikistan, Fergan Valley. ZISP 20097(371-87), 10715 (3-FMNH 141392-3, 141395-6, 141398: Iran, Fars prov.

13): Tajikestan-Afghanistan border, Termez. ZISP 13588,

Ahram. ZISP 10335: Iran, Kochrud, Irak-Adschemi (exact

foot of Kuh-i-Bang, north of Ganaweh. [29 44 N, 50 19 E].

13590(1-12), 13592.6914(1-10): Kazakhstan-Uzbekistan,

locality?) (Nikolsky's Type of Agama kennanensis , var.

between Aral Lake and Caspian Sea. ZISP 19392 (1-24):

brevicauda ). ZISP 9321: Iran. Kochrud. Irak-Adschemi

Russia (west of Caspian Sea), Daghestan. ZISP 15753: Iran.

(exact locality?) (Nikolsky's Type of Agama kennanensis ).
ZISP 9889: Iran (exact locality ?).

Astarabad (= Gorgan?). CAS 183032-39: Russia, ChechenIngush Autonomous Republic, the lowland between TerekKuma Rivers [43 21 N, 46 06 E], CAS 120249-50: Afghanistan, 30-70

km E of Herat 3700-5350 ft. CAS 1 15922-3:

Afghanistan, Maimana, 35 54 N, 64 43 E. 884m. CAS

Afghanistan, 10km NE Darweshan (Central
CAS
120275: Afghanistan, 25 km E KhanabadAfg.).
(between Mazare-Sharif-Faizabad), 2400ft. CAS 115920:
120280-

1:

Afghanistan, northeastern Afghanistan. 64 mi by road E
Faizabad, 37 05 N, 70 40 E. CAS 1 15921 Afghanistan,:

Paghman Vicinity, 34 36 N, 68 56 E. 2440m. CAS 120251:

NW

Trapelus agilis pakistanensis (n =32)

SMF 63258 (holotype), 63259, 63279 (paratypes): Pakistan,
Gaj-River, Kirthar Range. SMF 63256 (paratype): Pakistan,
Sonda Distr.Thatta. SMF 63236: Pakistan, Old Airport of
Karachi. 63243-4 and 63286-7 (paratypes): Pakistan, Old

Airport of Karachi.
at Goth

SMF 63239, 63264: Pakistan, Hab River

Mauladad. SMF 63263: Pakistan, Jati, Sind. SMF

63276: Pakistan, Karangee at Karachi. SMF 63265: Pakistan, Karachi.

SMF 63267: Pakistan. Karachi area. FMNH

Pul-e-Khumri, 2400ft. (near MazarAfghanistan. 25km
sharif?). CAS 120255: Afghanistan, 20 km E Mazar-e-

224946: Pakistan, Sind Prov. Dadu dist. Ranicot. FMNH

Sharif.

ment.

CAS 120253: Afghanistan, 10 km W Tashkurgan,
Near Mazar-Sharif? CAS 120256-8: Afghanistan, 20-50 km
E Mazar-e-Sharif. CAS 120259-61: Afghanistan, 45 km W
Mazar-e-Sharif, 1500ft. CAS 120273: Afghanistan, 50 km

W Mazar-e-Sharif. FMNH 161 197-9: Afghanistan, Maimana, 35 54 N, 64 43 E 884 m.

FMNH 161 133: Afghanistan,

64mil E Faizabad. 35 05 N, 70 40 E. FMNH 161 191-2:
Afghanistan, Paghman vicinity, 34 36 N, 68 56 E. 244m.

FMNH 141399: Iran, Mazandaran prov.

1

mi N of Pahlavi

Dezh.

Trapelus agilis khuzistanensis (n =97)

GNHM. Re. ex. 5424 (holotype): Iran, 5 km NW Haft-Gel
on the road to Shushtar. GNHM. Re. ex. 5425 (paratype):
Iran, Khuzistan prov., 5 km NW of Haft-Gel on road to
Shushtar. GNHM. Re. ex.5426 (paratype) Iran, Khuzistan:

244977: Pakistan, Sind Prov. Karachi Dist. Malir. conton-

FMNH 244948: Pakistan, Sind Prov. Karachi Dist.
BMNH 1933.7.8.23: Pakistan, Salt
Range. Punjab. BMNH 74. 4. 29. 1432, 80. 1. 10. 25: Pakistan, Sind area. BMNH 98. 12. 22. 7-8: Pakistan: Kurrachee(karachi). BMNH 1964-271: Pakistan, Mirpur Khas.
BMNH 1933.12.7.1: Pakistan, Sind area.
Malir, contonment.

1