Taxonomy and biogeography of the Iranian species of Laudakia (Sauria:...

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Taxonomy and biogeography ofthe Iranian species of Laudakia(Sauria: Agamidae)Nasrullah Rastegar-Pouyani a & Göran Nilson ba Department of Biology, Faculty of Science, RaziUniversity, Kermanshah, Iran E-mail:b Department of Zoology, Gothenburg University, Box463 SE-405 30, Gothenburg, SwedenVersion of record first published: 28 Feb 2013.

To cite this article: Nasrullah Rastegar-Pouyani & Göran Nilson (2002): Taxonomy andbiogeography of the Iranian species of Laudakia (Sauria: Agamidae), Zoology in theMiddle East, 26:1, 93-122

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Taxonomy and biogeography of the Iranian species of

Laudakia (Sauria: Agamidae)

by Nasrullah Rastegar-Pouyani and Göran Nilson

Abstract. At least 5 species of the genus Laudakia Gray, 1845, as one of the representatives of the western radiation of the Agamidae, occur on the Iranian Plateau. Based on extensive field re-search, collecting material from most parts of the Iranian Plateau, examination of material from museum collections, employing statistical methods, and by referring to almost all the available relevant literature, we discuss the taxonomy and biogeography of the Iranian species of Laudakia. From the standpoint of body stature and scale morphology L. caucasia and L. microlepis are distinguishable based on several species-specific characters. Furthermore, descriptive statistics and ANOVA-based pairwise comparison of metric and meristic data show that in most characters they are significantly different. Based on scale morphology, descriptive statistics as well as ANOVA it is shown that L. nupta nupta and L. n. fusca are at least subspecifically different, and this is also the case with L. melanura melanura and L. m. lirata. Different ideas concerning the nomenclature of the Palaearctic ring-tailed, rock-dwelling agamids are discussed and the generic name Laudakia Gray is adopted as the best-fitted alternative. A summary of regional geology and palaeogeographgical events is presented, and different scenarios for the origin, radiation, and sub-sequent evolution of Laudakia species are discussed. Both dispersal waves and vicariant events are proposed to be involved in isolation, radiation, and speciation in Laudakia, and the high mountain ranges of northeastern Afghanistan and southeastern Tajikistan (Pamir) are regarded as their hypothetical centre of origin. A key to the Iranian species of Laudakia is provided.

Abstract. Auf dem iranischen Plateau kommen mindestens 5 Arten der Gattung Laudakia Gray, 1845 vor, die den westlichen Formenkreis der Agamidae repräsentieren. Wir diskutieren die Ta-xonomie und Biogeographie der iranischen Arten von Laudakia, aufbauend auf extensiven Feld-forschungen mit Material aus fast allen Teilen des iranischen Hochlandes, die Untersuchung von Museumsmaterial, die Anwendung von statistischen Verfahren und die Auswertung von Literatur. L. caucasia und L. microlepis lassen sich anhand mehrerer Merkmale der Körperstatur und Be-schuppung unterscheiden. Auch deskriptive Statistiken und ein auf ANOVA basierender paarwei-ser Vergleich von metrischen und meristischen Merkmalen zeigen, dass sich die meisten Merkma-le signifikant unterscheiden. Weiterhin wird gezeigt, dass deskriptive Statistiken und eine ANO-VA-Analyse bezüglich der Beschuppung L. nupta nupta und L. n. fusca zumindest auf subspezifi-schem Niveau trennen. Das gleiche gilt für L. melanura melanura und L. m. lirata. Unterschiedli-che Vorstellungen zur Nomenklatur der paläarktischen felsbewohnenden Wirtelschwanzagamen werden diskutiert und die Genusbezeichnung Laudakia Gray wird als beste Alternative angese-hen. Regionale geologische und paläogeographische Ereignisse werden skizziert und verschiede-ne Szenarien für die Entstehung, Radiation und die sich daraus ableitende Evolution der Lauda-kia-Arten diskutiert. Es wird davon ausgegangen, dass im Verlauf der Isolation, Radiation und Artbildung sowohl Dispersionswellen und vikariierende Ereignisse beteiligt waren. Die Hochge-birge im Nordosten von Afghanistan und Südosten Tadschikistan (Pamir) werden als hypotheti-sche Entwicklungszentren betrachtet. Ein Schlüssel zu den iranischen Arten der Gattung Lauda-kia wird vorgestellt.

Key words. Agamidae, Laudakia, Iranian Plateau, Central Asia, distribution, historical bioge-ography, taxonomy.

Zoology in the Middle East 26, 2002: 93–122. ISSN 0939-7140 © Kasparek Verlag, Heidelberg

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94 Zoology in the Middle East 26, 2002

Introduction

The Agamidae is a morphologically and ecologically diverse family consisting of more than 330 contemporary species and 54 genera. This family is the fourth largest lizard family and is widely distributed throughout various regions of the Old World and Australia. Its morpho-logical and ecological counterpart is the family Iguanidae which is widely distributed throughout the New World (MOODY 1980). Agamids occur in a wide range of different habitats including arid, semi-arid, tropical, and sub-tropical regions. MOODY (1980) split the Agamidae into six groups. Group VI (sensu MOODY) includes the western radiation of the family distributed in central and southern Asia, southeastern Europe, and Africa with about 9 genera (Agama, Brachysaura, Laudakia, Phrynocephalus, Pseudotrapelus, Stellio, Trapelus, Uromastyx, and Xenagama). These genera are mostly rock and desert dwellers. On the Ira-nian plateau, which extends from the highlands of eastern Turkey and Iraq to western Af-ghanistan and Pakistan, and zoogeographically is a connecting cross-road between south-eastern Europe and northwestern India and western Pakistan on the one hand, and Central Asia and northeastern Africa and Arabia on the other, five of the above-mentioned genera of agamids consisting of about 25 species occur which are as follows: Calotes (1 species), Laudakia (at least 5 species), Phrynocephalus (7–11 species), Trapelus (at least 3 species), and Uromastyx (3 species). The genus Laudakia comprises about 16 species which mainly occur in the upland and mountainous regions of central and southern Asia; of these, at least five species occur on the Iranian Plateau as follows:

L. caucasia caucasia (Eichwald, 1831): Iran, Iraq, Turkey, Armenia, Georgia, Azarbaijan, Rus-sia, Turkmenistan, Tadjikistan, Afghanistan, Pakistan.

L. microlepis (Blanford, 1874): Iran, Pakistan, Afghanistan. L. erythrogastra (Nikolsky, 1896): Iran, Afghanistan, Turkmenistan. L. melanura lirata (Blanford 1874): Iran?, Pakistan. L. nupta nupta (De Filippi, 1843): Iraq, Iran, Afghanistan, Pakistan. L. nupta fusca (Blanford, 1876): Iran, Pakistan.

The other taxa of the genus are:

L. caucasia triannulata (Ananjeva and Ataev, 1984): Turkmenistan. L. agrorensis (Stoliczka, 1872): Afghanistan, Pakistan, India. L. badakhshana (Anderson and Leviton, 1969): Afghanistan. L. chernovi (Ananjeva et al., 1981): Tadjikistan, Turkmenistan, Uzbekistan. L. himalayana (Steindachner, 1869): Tadjikistan, Uzbekistan, Kyrgyzstan, Afghanistan, Pakistan,

India. L. lehmanni (Nikolsky, 1896): Tadjikistan, Uzbekistan, Kirgizistan, Afghanistan, Turkmenistan. L. melanura melanura (Blyth, 1854): Afghanistan?, Pakistan. L. nuristanica (Anderson and Leviton, 1969): Afghanistan, Pakistan. L. pakistanica (Baig, 1989): Pakistan. L. sacra (Smith, 1935): Tibet (China). L. stellio (Linnaeus, 1758): Greece, Cyprus, Egypt, Israel, Lebanon, Jordan, Syria, Turkey, Iraq. L. stoliczkana (Blanford, 1875): Mongolia, China. L. tuberculata (Hardwicke and Gray, 1827): India, Nepal, Afghanistan, Pakistan.

The taxonomic status of L. caucasia and L. microlepis is problematic, and different workers regard them as being conspecific or as having separate specific status. We have chosen to consider these two taxa as specifically different based on significant differences in most morphological characters.

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Fig. 1. Distribution of the genus Laudakia.

The main objectives of this work are to present a deeper treatment and to open new in-sights into the taxonomy and biogeography of the Iranian species of Laudakia, using distin-guishing morphological characters, employing univariate statistics as well as reviewing almost all the relevant literature. Undoubtedly, further studies and field work in various parts of the plateau and neighbouring regions will throw more light upon the systematic, phyloge-netic, and biogeographic status of its herpetofauna.

Material and methods

The lizards used in this study are listed in the Appendix. These specimens were collected during field work on the Iranian Plateau (in 1995, 1996, and 1998) and were also borrowed from various museum collections (see ‘abbreviations and acknowledgments’); during field work on the Iranian Plateau, the senior author surveyed remote and herpetologically unexplored regions, and identi-fied some new and distinct populations of agamids (e. g. Laudakia nupta nupta and L. microlepis from central and southeastern Iran), noting their mode of adaptation to various habitats. All the specimens (which were adult males and females) were examined and compared for the nine most distinguishing metric and meristic characters. Descriptive statistics was used to explore the means, standard errors, and ranges of these characters in all the species studied (Tab. 1).

In the Laudakia caucasia-microlepis complex, L. melanura-lirata, and L. nupta-fusca, an ANOVA-based pairwise comparison of the metric and meristic data, by use of the Scheffe’s F test for the post-hoc comparison, showed that in most characters the two species (or subspecies) within each complex are significantly different at the level of 0.05 (Tab. 2). Then the obtained data for each pairwise comparison were plotted against the two species (or subspecies) of each complex as interaction line plots to show the graphical display of the mean differences. The homogeneity of variances was tested by means of the Fmax test (FOWLER et al. 1998). All the ana-lyses were run by the Excel (4.0) and Statview (4.5) statistical packages.

In addition, 9 characters pertaining to body stature and scale morphology were studied and the above-mentioned taxa were compared for some of these characters (Tab. 3).

We also reviewed all the available literature concerning the taxonomy and biogeography of Laudakia, with special emphasis on the Iranian species, from the 19th century to the present time. This literature review constitutes an important portion of this work. The results obtained from the morphological analysis of lizard specimens were compared with all the available data from the

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literature as well as other unpublished data. Based on these comparisons, we are attempting in this paper to resolve the taxonomic status and discuss the historical biogeography of Laudakia with special emphasis on the Iranian species. Metric and meristic (scale count) characters

SVL: snout-vent length (from tip of snout to anterior edge of cloaca); – TL: tail length (from posterior edge of cloaca to tip of tail); HL: length of head (from tip of snout to posterior edge of tympanum); – HW: width of head (from the widest part); – HH: height of head (from top of head to the lower jaw); – SQ: scales around midbody (number of scales in a single row around widest part of body); – LVS: large vertebral scales (number of longitudinal rows of enlarged scales on the vertebral region; – AGFS: anus-gular fold scales (number of ventral scales in a single row from posterior edge of gular fold to vent); – FCWS: fifth caudal whorl scales (scales around fifth caudal whorl just behind the vent). Abbreviations BMNH = The Natural History Museum (London, UK); – CAS = California Academy of Sciences (San Francisco, USA); – GNHM Re. ex. = Gothenburg Natural History Museum Reptilia exotica (Gothenburg, Sweden); – FMNA = Field Museum of Natural History (Chicago, USA); – INNHM (= MMTT) = Iran National Natural History Museum (Tehran, Iran); – SMF = Museum und For-schungsinstitut Senckenberg (Frankfurt a.M., Germany); – SMNH = Swedish Museum of Natural History (Stockholm, Sweden); – ZISP = Zoological Institute of the Russian Academy of Sciences (St. Petersburg, Russia).

Results

Genus Laudakia Gray, 1845

Definition: Body depressed; dorsal crest absent (a rudimentary nuchal crest may be present); gular sac absent; head depressed and more or less triangular; tympanum exposed and large, its diameter at least half that of the orbit, more or less superficial; a prominent transverse gular fold and a fold in front of shoulder present; dorsal scales uniform or intermixed with larger ones; toes compressed, fifth extending beyond first; caudal scales forming more or less distinct annuli; males with callose scales on preanal and usually abdominal, and someti-mes, with highly modified scales on the plantar and palmar regions. Chromosome number of all the studied species is 2n = 36 (except Laudakia tuberculata with 2n = 34) (GORMAN 1973, GORMAN & SHOCHAT 1972, MOODY 1980, MOODY & HUTTERER 1978, PECCININI-SEALE 1981, SOKOLOVSKY 1975).

Distribution: Some 18 species of Laudakia are distributed from northern Egypt and some Greek islands in the west to Nepal, China, and Mongolia in the east (see introduction and Fig. 1). Of these, four species are documented in Iran. In addition, one doubtful species most likely occurs in the country.

Laudakia caucasia (Eichwald, 1831) (Fig. 2)

Stellio caucasius Eichwald, 1831: 3: 187

Type locality: Azarbaijan Republic (Baku) and Georgia Republic (Tiflis) (restricted to Baku by ANDERSON 1999).

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Fig. 2. Laudakia caucasia , an adult male from the Zagros Mountains, Kermanshah province, western Iran.

Diagnosis: Body strongly depressed; tail divided into distinct segments, each segment com-posed of two whorls of scales and its length less than twice body length; gular scales smooth, occipital scale not enlarged; tympanum not larger than eye, superficial; enlarged scales on the middle of flank forming patches, or scattered; 120-185 scales around widest part of body; upper head scales smooth; middorsal region with 7-11 longitudinal rows of rather keeled and large, flattened scales; adult males with callose scales on preanal, midventral, and sometimes with highly modified scales on palmar and plantar regions.

Olivaceous or yellowish-brown above, with yellow and black markings, back sometimes with regularly arranged pale yellowish spots margined with black; tail yellowish or oliva-ceous, with or without dark annuli; the throat often heavily marbled with dark, blue, or black; sometimes the whole belly and under surfaces of the limbs dark-bluish; young greyish or olivaceous above with black spots upon the back, and occasionally with cross-bars on dorsal regions (ANDERSON 1966a, 1999).

This is one of the most studied and widespread taxa of Laudakia, mainly associated with the mountainous areas. ANANJEVA & ATAEV (1984) described a new subspecies of this lizard from southwestern Turkmenistan as Stellio caucasius triannulatus. According to BAIG (1992), this new subspecies is more related to microlepis than caucasia, and the same author placed this new taxon under microlepis because of high scale counts. ANANJEVA & ORLOVA (1979) reviewed the distribution of L. caucasia and defined the easternmost range as sou-thern Tadjikistan. They also compared different populations of L. caucasia with respect to some morphometric and morphological characteristics and divided populations of this lizard into western and eastern groups. They doubted that Mzymta River Gorge on the Black Sea coast was within the range of L. caucasia. It is a very humid and unusual locality for this lizard, and the one specimen recorded by DAREVSKY in 1972 (ZISP 18284) was a case of accidental introduction, probably by tourists (ANANJEVA, pers. comm.).

With regard to the controversies still surrounding the taxonomic status of L. caucasia and L. microlepis, we used descriptive statistics as well as analysis of variance (ANOVA) to explore the metric and meristic differences between the two taxa (Tabs. 1-2). It is clear that two taxa are significantly different in all morphological characters except TL (P<0.05).

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Fig. 3. Distribution of Laudakia caucasia. square = type locality: Azarbaijan Republic, Baku.

There are also distinct differences in body stature and scale morphology between the two taxa (Tab. 3). Based on the mitochondrial DNA analysis, the specific status of L. caucasia and L. microlepis was shown to be established (MACEY et al. 1998). Based on morphological and molecular evidence, the specific status of both taxa is therefore confirmed and any taxo-nomic decision regarding them as conspecific is inadvisable.

In addition we examined several specimens of L. c. triannulata from Turkmenistan. The tail segments, which are basically two-whorled, often have a small extra whorl more devel-oped on the sides of tail, but in the dorsal and ventral view, the two-whorled structure is persistent, and in one specimen (CAS 184640) all the caudal segments are bi-whorled. The relatively high number of scale counts around the body (which is also found in some populations of L. c. caucasia), led BAIG (1992) to assume that this new taxon is a subspecies of microlepis (L. microlepis triannulatus). In body stature and scale morphology, however, they are indistinguishable from L. c. caucasia. We thus propose that recognition of the tri-nomen L. c. triannulatus be discontinued. MACEY et al. (1998) came to the same conclusion based on molecular analysis of different populations of the L. caucasia complex. With the resurrection of microlepis by some authors (ANANJEVA & ATAEV 1984, ANANJEVA & TUNIEV 1994, BAIG 1992, present paper), the Pakistan, Afghanistan, and Tadjikistan popula-tions need to be reviewed. BAIG (1992) studied several specimens from Afghanistan. This author writes: „I am very much sure that both caucasia and microlepis occur in Afghanistan. However, it is yet to decide whether they live together or they inhabit separate biotopes in the same locality.“

According to ANDERSON (1968), the range of L. caucasia does not overlap with that of L. microlepis except in the mountains of the northeastern corner of the Iranian Plateau where three large, rock-dwelling taxa of Laudakia occur very close together (L. caucasia, L. micro-lepis, and L. erythrogastra). But with regard to BLANFORD’s record from Kushk-e-Zard (between Shiraz and Esfahan, south-central Iran) (BLANFORD 1874), we do not rule out a limited contact between the two taxa in the mountains around Esfahan city.

Laudakia caucasia is distributed over a wide area (Fig. 3): in Turkey it is limited to a small area in the northeast bordering the Armenian and Iranian frontiers. CLARK et al. (1973)

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documented its westernmost limits of distribution east of Kars, Turkey, where the to-pography changes abruptly from rich grassland through steep, eroded, rocky hillsides down to open stony valleys. Both L. caucasia and L. stellio meet in eastern Turkey; the distribu-tions do not overlap, but are sharply separated from each other. The boundary between both species is the eastern shore of Lake Van (BARAN et al. 1989). Laudakia caucasia also occurs in Transcaucasia west to the Suramask Ridge, and the mountainous parts of Daghestan. The syntypes come from Tiflis, Georgia, and Baku, Azarbaijan Republic [restricted by ANDER-

SON (1999) to Baku]. East of the Caspian Sea, it is a mountainous species in southern and southwestern Turkmenistan, southern Uzbekistan, and southern Tadjikistan. In northern Afghanistan it occupies the area north of the Central Massif, and extends south in northeast-ern Afghanistan to Paghman (ANDERSON 1999) (Fig. 3).

Some authors believe that it also occurs in the western and southwestern parts of Pakistan (KHAN 1980, KHAN & MIRZA 1977, MERTENS 1969, MINTON 1966, SMITH 1935), but BAIG (1992) excludes southern and southwestern Pakistan from its range and restricts the range to a small area in northwestern Pakistan in the vicinity of the Afghanistan border.

LEVITON et al. (1992) believe that earlier records of L. caucasia in Iraq are based on mis-identifications of L. nupta. ANDERSON (pers. comm.) examined specimens from Iraq at FMNH, regarded as L. caucasia by the previous authors (REED & MARX 1959), and determi-ned them as L. nupta.

However, Laudakia caucasia has been collected in the northern parts of Kermanshah pro-vince, along the Zagros Mountains, not so far from the Iraqi border (RASTEGAR-POUYANI, MS). It is therefore most likely that this species occurs in the northeastern regions of Iraq as well, though there is no documented record for its occurrence inside Iraqi territory. KHALAF (1959) reports L. caucasia from Iraq without specifying the locality.

On the Iranian plateau, L. caucasia is nearly confined to the Alburz, Zagros, and Kopet-Dagh mountains. It does not occur in the western foothills of the Zagros nor does it occur in the southern slopes of Zagros, where L. nupta occupies a similar range (ANDERSON 1966a, 1968, 1999).

Laudakia microlepis (Blanford, 1874) (Fig. 4)

Stellio microlepis Blanford, 1874a: 453. Type locality (here restricted): Iran, Khaneh-Sorkh Pass, between Sirjan and Kerman.

Diagnosis: Usually larger and more depressed than L. caucasia; tail shorter and divided into distinct segments, each composed of two whorls of scales, its length often less than 1.35 of body length; flanks with or without enlarged scales; enlarged vertebral scales smooth or feebly keeled; tympanum almost as large as orbit, superficial; gular scales smooth; scales on upper surface of tail not larger than those on upper surface of tibia; usually more than 190 scales round the middle of body; patches of callose scales almost always present in both sexes and strongly developed in males, occupying a large area; gular region often banded with dark-grey stripes; belly pale yellow and sometimes with black spots.

This taxon was first described by BLANFORD in 1874 from two localities in Iran; Khane-Sorkh Pass, about 2750 m elevation, between Sirjan and Kerman, southeastern Iran, and Kushk-e-Zard in south-central Iran. These two localities are separated by a gap of about 440 km. In order to stabilize nomenclature by restricting the name microlepis to a single specimen, we restrict the type locality of this taxon to the Khane-Sorkh Pass, between Sirjan and Kerman, as the place from which BLANFORD (1874) recorded this taxon for the first

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Fig. 4. Laudakia microlepis in its natural habitat, Khane-Sorkh Pass, Kerman province, southeastern Iran.

time, and we designate the only syntype from this area preserved in the BMNH (1946.8.28.74) as the lectotype [BLANFORD collected four syntypes in this area (BLANFORD 1876: 326) but apparently only one of them is deposited at the BMNH (MCCARTHY, pers. comm.)]. Subsequent authors (ANANJEVA & TUNIEV 1994, ANDERSON 1966a, 1968, 1974, 1999, BAIG 1992, BOULENGER 1885a) dealt with it as a distinct taxonomic entity and at the same time they also expanded the range of this taxon from its type locality to the central, south-central, and southeastern parts of the Iranian Plateau, into Afghanistan and Baluchis-tan, Pakistan.

ANDERSON (1966a) writes: “In view of the variation in the number of scales around the body in Agama caucasica (up to 221 in some specimens from Afghanistan) the status of this form needs re-evaluation when adequate material becomes available”. WETTSTEIN (1953) is of the opinion that "microlepis" should be regarded as a subspecies of L. caucasia, and BLANFORD himself (1876: 326–327) remarked on the very close relationship between the two taxa. MINTON (1966) and MERTENS (1969) regarded the Pakistani populations as cauca-sia, contrary to what has been observed during later studies.

Further studies extended the range of this lizard even more, including almost the whole of Afghanistan and the southern regions of Turkmenistan (ANANJEVA & ATAEV 1984, ANAN-

JEVA & TUNIEV 1994, BAIG 1992). The presence of both microlepis and caucasia at several localities in Afghanistan rules out the possibility of treating them only as distinct populations (BAIG 1992).

In comparison with L. caucasia, the scales in L. microlepis are smaller and usually there is no overlap in the number of scales around the middle of the body (usually less than 175 scales in L. caucasia and more than 190 in L. microlepis). BLANFORD (1876) counted 210–220 scales around the middle of body in L. microlepis. In addition, the callose preanal and midventral scales are almost always present in adults of both sexes, occupying a large area in males. BAIG & BÖHME (1991) postulate some functional implications for the callose scales in various species of Laudakia.

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Fig. 5. Distribution of Laudakia microlepis. Square = type locality (restricted): Iran, Kerman Province, Khane-Sorkh Pass between Sirjan and Kerman.

As mentioned before, the statistical analysis of metric and meristic characters as well as distinct differences in body stature and scale morphology show that L. caucasia and L. mi-crolepis are distinguishable at the specific level (Tabs. 1-3).

In 1996 one of us (N. R.-P.) surveyed the type locality of L. microlepis, Khane-Sorkh Pass, between Sirjan and Kerman, Kerman province, and collected 8 specimens of this lizard (topotypes). They were collected on the stony hills and under boulders very close to a moun-tainous stream at about 2700 m elevation with Salix, Fraxinus, Populus as well as shrubby and bushy vegetation. After a precise examination of these topotypes, we are inclined to accept that they are specifically different from L. caucasia.

There are also some differences in colour pattern between the two taxa; the general colour in L. microlepis is dusky olivaceous, the scales in the middle of back never being conspicu-ously paler as in L. caucasia, and frequently being darker than on the sides; the whole colour is more uniform than in the former subspecies; the cross-bands of pale spots are very much less marked and smaller; the ventral surface is yellowish-cream and sometimes with dark spots which have a tendency to form reticulation; the gular area is often banded with dark-grey longitudinal stripes (ANDERSON 1999). Based on molecular studies, this taxon appears to be the sister taxon to an extremely well supported clade containing all L. caucasia and L. erythrogastra populations (MACEY et al. 1998). In the mountains of the southern, central, and eastern parts of the Iranian plateau, from the area around Izad-Khast, L. microlepis re-places L. caucasia. In the region of Torbat-e-Heydariyeh in NE Khorasan (NE Iran) the two species apparently occur at localities very close together. In Afghanistan L. microlepis oc-curs south of the Kabul River Valley in the mountains of eastern Afghanistan. It has also been recorded west of the central Massif, south of Herat, and in the mountains of eastern Iran along the Afghan frontier. In Pakistan, it occurs in Waziristan and Baluchistan (ANDERSON 1999). The areas of probable intergradation with L. caucasia, if any, have not been estab-lished. The distribution of L. microlepis is shown in Fig. 5.

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Fig. 6. Distribution of Laudakia n. nupta . Square = type locality: Iran, Fars Province, Persepolis,

northeast of Shiraz.

Laudakia nupta nupta (De Filippi, 1843)

Agama nupta De Filippi, 1843: 407. Type locality: Iran, Fars province, Persepolis (NE of Shiraz).

Diagnosis: Head distinctly triangular; tail divided into distinct segments, each segment com-posed of more than two whorls of scales, usually three, or, sometimes, segmentation incon-sistent or indistinct; 81–122 scales around body; median dorsal scales large, keeled, broader than long, in 16–22 oblique longitudinal rows; a prominent transverse fold across nape; often a well developed dorsolateral fold on each side of body; adult males sometimes with plantar and palmar callose scales, in addition to the preanals and midventrals; distal half of tail usu-ally dark-brown or black, sometimes with light rings; tail length more than twice the length from gular fold to vent.

Light-brown, olive, pale green, or dark brown above, with numerous scattered black and yellow scales; distal one-third to half black or dark brown, sometimes with narrow light rings; venter light tan, straw coloured; chin, throat, and chest of male dark blue, often mot-tled with white; juveniles with dark transverse markings on back, breaking up into reticula-tions with age (ANDERSON 1999).

DE FILIPPI described this species in 1843, based on material collected from Persepolis, a-bout 45 km NE of Shiraz, Fars Province. The holotype is deposited at the Milan Museum, Italy. Later on, two subspecies of this taxon were identified by BLANFORD: Laudakia nupta nupta and L. n. fusca (BLANFORD, 1876: 317–320). Subsequently BOULENGER (1885) recog-nized these subspecies and separated fusca from the nominate form by having more devel-oped spinose scales on the sides of head and neck. SMITH (1935) did not find any significant differences between L. n. nupta, L. n. fusca, and Stellio carinatus Duméril, and placed all three under the nominate form nupta. MINTON (1966) and MERTENS (1969) again resurrected fusca as a subspecies of nupta based on characters which had already been described by ANDERSON as age dependent (BAIG 1992). Also KHAN & MIRZA (1977) divided the popula-

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tions of L. nupta in Pakistan into two subspecies: L. n. nupta, and L. n. fusca. ANDERSON (1974) differentiated both forms based on the presence or absence of a transverse fold on the nape. Based on the statistical analysis of morphological characters (Tabs. 1-2), as well as scale morphology (Tab. 3), we follow MINTON (1966), MERTENS (1969), and ANDERSON (1974) and discuss these two taxa at the subspecific level; nor do we believe that the ob-served morphological differences are age or habitat dependent. As is evident, the two taxa are significantly different (P<0.05) in characters other than HH, and this giveis great support for recognizing them as two distinct taxonomic entities, at least at the subspecific level. Laudakia n. nupta is one of the largest rock-dwelling agamids occurring in northeastern and eastern Iraq eastwards through Iran, into Afghanistan and Pakistan. It is abundant in the foothill regions, inhabiting rock outcrops where there are deep crevices.

In 1996 one of us (N. R.-P.) surveyed the type locality of L. n. nupta, Persepolis, about 45 km NE of Shiraz, Fars Province, and collected 8 specimens of this lizard (topotypes). It is abundant on the walls and pillars of the ancient palace of Persepolis as well as on the sur-rounding stony hills and boulders. They were quite active when the air temperature was about 37°C.

According to ANDERSON (1966a, 1999), L. n. nupta is primarily a species of the southern and southwestern mountains of the Iranian plateau, distributed mainly on the outer slopes of the Zagros Mountains. During field work in the central and south-central parts of the Iranian Plateau in 1996, N. R.-P. collected specimens of L. n. nupta in the central and east-central regions and it seems that it might have a continuous distribution throughout most parts of the Iranian Plateau. Laudakia n. nupta together with L. caucasia and L. stellio represents the western radiation of the genus. In southeastern regions of the Iranian plateau, L. n. fusca replaces L. n. nupta but their exact distributional limit and the degree of intergradation, if any, is not known.

TUCK (1979) observed this lizard in the Turan Protected Area in the northern Dasht-e-Kavir, north-central Iran. In 1996 an Iranian expedition collected L. n. nupta from NE of Semnan province (ca 56°15’E, 36°55’N). This is the most northern proper record of L. n. nupta to date (E. RASTEGAR-POUYANI, pers. comm.). In addition, the northwestern limits of its distribution in Kermanshah and Kurdestan provinces along the central regions of the Zagros Mountains were determined, where its range comes in contact with L. caucasia (pers. observ.). The main distributional route of L. n. nupta is in a northwest-southeast direction, mainly along the outer regions of the Zagros and its western foothills, where it is the only representative of Laudakia. The distribution of L. n. nupta is shown in Fig. 6.

Laudakia nupta fusca (Blanford, 1876) (Fig. 7)

Stellio nuptus var. fuscus Blanford, 1876: 319-320. Type locality (here restricted): Iran, Sistan-Baluchistan Province, near Jalk.

Diagnosis: Differing from the nominate subspecies in the absence of nuchal fold across neck, or the fold is rudimentary, as well as in colouration of male and in certain details of scalation; often less than 90 scales round middle of the body; spiny excrescences around ear opening and sides of neck larger than in L. n. nupta and more numerous; enlarged middorsal scales in 13–16 rows and generally more strongly mucronate than in L. n. nupta. In addition, scales of base of tail much larger than in the nominal form, often less than 17 on the fifth whorl behind the vent.

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Fig. 7. Laudakia n. fusca, an adult male from southern Pakistan.

Entire head of male (at least in reproduction season) canary-yellow, remainder of body, limbs, and tail dark brown to sooty black above and below save for sparse yellow speckling on back and for callose ventral and preanal scales which are amber; females drab brown to russet, indistinctly speckled with yellow; distal part of tail black; ventral surfaces straw. Juveniles greyish-yellow, with narrow, rather irregular cross bars of brown, in some cases forming reticular pattern, most of head black, with yellow chevron on crown and large yel-low spots on temporal area, occiput, neck, and shoulders; tail banded almost to tip which is black (MINTON 1966).

BLANFORD (1876) found two specimens of this form in SE Iran, Baluchistan province, one at Kalagan and the other in the vicinity of Jalk, near the Iranian-Pakistani border, at 1000-1150 m elevation. In order to stabilize nomenclature by restricting the name fuscus to a sin-gle specimen, we restrict the type locality of this taxon to SE Iran, Baluchistan province, vicinity of Jalk, and designate the syntype (BMNH 74.11.23.11) collected in this area as lectotype.

According to BLANFORD (1876: 320), these two specimens differ from all others in their very dark colour, in the absence of the fold on the back of the neck, and in the scales on this area being not quite so minute as in the typical form. This author feels some doubt about distinguishing these two forms specifically and believes that the dark colouration may be partly due to the season, as is also the cutaneous fold across neck which is to some extent variable in the nominal subspecies (L. n. nupta). As mentioned before, some authors (e.g. BAIG 1992, SMITH 1935) place the two subspecies under the name nupta and regard the differences as related to the seasonal and environmental changes, but others (e.g. ANDERSON 1966a, 1968, 1974, 1999, BOULENGER 1885a) distinguish fusca from nupta by the absence of a transverse fold on the nape and having a yellow head and sooty or black body in males. As noted above, we believe that these two taxa are at least subspecifically distinct, based on the statistical analysis of the metric and meristic data as well as significant differences in scale morphology (Tabs. 1-3).

According to ANDERSON (1999), a specimen from Binak, Persian Gulf coast, southern I-ran, may belong to L. n. fusca. If so, then the range of L. n. fusca extends from southern and southwestern Pakistan, westward to southeastern and southern Iran. According to MINTON

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Fig. 8. The presumed distribution of Laudakia nupta fusca. Square = type locality (restricted): Iran, Sistan-Baluchistan Province, near Jalk.

(1966), L. n. fusca occurs in the southern and southwestern parts of Pakistan, near Khadeji Falls about 30 miles NE of Karachi, near Diwana on the upper Hab River, and in the south-ern part of Pab Hills. KHAN & MIRZA (1977) also reported this subspecies from the southwestern parts of Baluchistan. According to KHAN (1980), L. n. fusca almost covers the whole northern and southern Baluchistan (Fig. 8) and is endemic to this area. This is a very shy lizard, almost always found in rocky areas with vertical or near vertical surfaces and crevices that provide shelter. Other factors such as the type of vegetation affect the distribu-tion. It has been recorded at elevations up to 2000 m in Pakistan (MINTON 1966).

Laudakia erythrogastra (Nikolsky, 1896) (Fig. 9)

Stellio erythrogaster Nikolsky, 1896: 370-371. Type locality (here restricted): Iran, Khorasan province, Fariman.

Diagnosis: Tail with strong ridges and spines, segmented on proximal two-thirds, each seg-ment composed of two whorls of scales; gular scales mucronate, strongly keeled in adults, weakly keeled in small juveniles; 78–129 scales round middle of body; one or two longitudi-nal rows of spiny tubercles on each side of body, 9–12 longitudinal rows of enlarged, keeled, and mucronate scales on middorsal region.

Colour olive with black, narrow, irregular spots and stripes, sometimes slightly distinct or absent; adult males almost black on lower side of body, callose preanal and midventral scales yellowish, and neck with large, light spots; females are orange in colour, more bright on tail.

NIKOLSKY (1896) described this taxon based on two specimens collected by ZARUDNY in the same year from NE Iran, Khorasan province, Fariman [not “Terimun” in NIKOLSKY 1915: 120 (about 70 km SE of Mashad)] and Kalandar-Abad (93 km SE of Mashad). In order to stabilize nomenclature by restricting the name erythrogastra to a single specimen, we restrict the type locality of this species to Fariman and designate the syntype collected in this area as lectotype (ZISP 8760).

30° 40° 50° 60° 70°

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Fig. 9–10. Left (Fig. 9): Laudakia erythrogastra, an adult male from the Kopet-Dagh Mountains, southern Turkmenistan. Right (Fig. 10): Laudakia melanura lirata, an adult male from southwestern Pakistan.

Once it was thought that this species is restricted to the mountains of NE Iran (NIKOLSKY 1915) but the reports of ANDERSON & LEVITON (1969), CLARK (1991), and BAIG (1992) indicate that it is distributed from northeastern Iran (Khorasan province) eastwards into southern Turkmenistan to the northern and eastern mountains of Afghanistan as far as Paktia and Paghman (Fig. 11).

Unlike the other species of Laudakia, it lives in clayey and sandy-loamy soils (L. caucasia triannulata which inhabits a similar biotope in Turkmenistan is an exception, see ANANJEVA

& ATAEV 1984). It has been found basking on small, man-made piles of rocks, under which it takes refuge when disturbed. In Badkhyz, NW Afghanistan and SE Turkmenistan, it occurs in areas with colonies of Rhombomys, the burrows of which serve as retreats for the lizards. Unlike other members of the genus, it avoids vertical slopes and rock outcrops (TERENTJEV

& CHERNOV 1949). CLARK (1991) reported this lizard from Torbet-e-Jam (Khorasan prov-ince, NE Iran) at 1120 m elevation, and also from 60 km SE Mashad at 1380 m elevation. He found it very wary and common at both localities. The habitat comprised deep holes and crevices in earth banks and cliffs and piles of stones. CLARK et al. (1966) remarked that this agamid is rare in collections and only known from the Mashad area in Iran and SE Turkme-nistan. However, since then it has been found by the Street Expedition in Afghanistan from the northwest regions of this country and more significantly from Paghman in eastern Af-ghanistan at 2440 m elevation (ANDERSON & LEVITON 1969). This indicates a much broader distribution than previously suspected and a considerable altitudinal range (CLARK 1991).

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Fig. 11. Distribution of Laudakia erythrogastra. Square = type locality (restricted): Iran, Khorasan province,

southeast Mashhad, Fariman.

TUNIEV et al. (1991) described a new subspecies of this lizard, L. e. nurgeldievi, from eastern Kopet-Dagh in Turkmenistan, on the basis of morphological characteristics. Based on mole-cular studies, however, MACEY et al. (1998) suggest that the recognition of subspecies for L. erythrogastra is inadvisable. Descriptive statistics of the specimens of this taxon that were studied are shown in Tab. 1.

According to ANANJEVA & TUNIEV (1994), L. erythrogastra belongs to the ‘caucasia’ complex, having a tendency to a semi-arboreal mode of life, occupying a considerably wide range of biotopes, but reaching its highest density in the areas with Pistacia vera in Turk-menistan. These authors believe that L. erythrogastra is closely related to L melanura and that their ancestors penetrated to the north from the south of the Iranian High Plateau (see under historical biogeography). On the northern slopes of eastern Kopet-Dagh, L. erythro-gastra inhabits elevations of 600–900 m and L. caucasia inhabits the biotopes at elevations of 800–1500 m (ANANJEVA & TUNIEV 1994). On the southern slopes of the eastern Kopet-Dagh, elevations of 800-1600 m are typical for L. erythrogastra and 960–2575 m for L. caucasia (ANDERSON 1968). L. erythrogastra occurs in NE Iran, Khorasan province and in the vicinity of Mashad, Fariman (type locality), Kalandar-Abad, Langarak, Sarakhs, Torbat-e-Jam, between Torbat-e- Jam and Fariman, and Abghale (Fig. 11). It also occurs in SE Turkmenistan, N Afghanistan north of the Central Massif and south through the mountain passes to Paghman (ANDERSON 1999).

In addition to the species discussed above, the following taxon most likely occurs in the southeastern regions of Iranian territory proper.

Laudakia melanura lirata (Blanford, 1874) (Fig. 10)

Stellio liratus Blanford, 1874a: 453. Type locality: Saman, Dasht Province, Baluchistan, Pakistan.

Diagnosis: Caudal segments each composed of more than two annuli, but annulation often indistinct; tail much longer than the body (2.0–2.6 times); median dorsal scales broader than

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??

Fig. 12. Distribution of L. melanura (the exact range of L. m. melanura and L. m. lirata is not known).

Square = type locality of L. m. lirata: Saman, Dasht Province, Baluchistan, Pakistan.

long, in 7–9 straight longitudinal series, grading into dorsolateral small scales; 120–151 scales round middle of body; scales of back of head and middle of back often strongly keeled; a rudimentary nuchal crest and a dorsolateral cutaneous fold often present. Adult males dark brown with strongly contrasting sandy-yellow head and neck; females dark brown or grey with light dorsal spots, head usually brown or greyish-olive, underside of thighs and anterior part of tail yellowish-cream.

This taxon was described by BLANFORD (1874a) as Stellio liratus based on one female specimen collected in Saman, Dasht Province, west of Gwadar, Baluchistan, Pakistan, about 70 km E of the Iranian frontier. BLANFORD who collected this specimen on the rocks in some barren hills says: “This species is evidently close to S. melanura Blyth, of which I have no specimen for comparison, but judging by the fact that J. ANDERSON was at one time disposed to consider S. melanura as the young of S. dayanus Stoliczka, I can only suppose that the dorsal scales in S. melanura must be considerably smaller than in the present form. S. mela-nura is said to have the tail scales in distinct verticils. I have not seen any species in which the annulation is so indistinct as in the present” (BLANFORD, 1876: 322).

The populations of L. melanura in Pakistan have been divided into two subspecies; L. m. melanura (Blyth, 1854) in which males and females are almost completely black and dorsal scales are weakly keeled or smooth, and L. m. lirata (Blanford, 1874) in which the head is bright sandy coloured and dorsal scales are strongly keeled. Descriptive statistics of the two taxa, based on the specimens examined, is shown in Tab. 1, and an ANOVA-based pairwise comparison shows that in HL, SQ, LVS, AGFS, and FCWS they are significantly different (P<0.05) (Tab. 2). In addition, major morphological differences pertaining to scale morphol-ogy between the two subspecies are shown in Tab. 3.

Laudakia m. melanura occurs in Pakistan from Hab Chauki, Baluchistan, Sind, Waziris-tan, and the Northwest Frontier Province, and L. m. lirata has been reported from Kirthar Range and Hab Chauki (Baluchistan) (MERTENS 1969). According to KHAN (1980: 141), the populations of L. m. melanura have a relatively wide range, penetrating deep through the Sofed Koh corridor, radiating out to the east to Salt Range in the northern Punjab, west to the Pab Hills and south to the plains of Sind. On the other hand, L. m. lirata is an endemic of southern Baluchistan and, most likely, southeastern Iran. In spite of these findings, the exact

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range of the two taxa is not known. In general appearance and habits this species resembles L. nupta fusca and in some localities of south Pakistan they occur sympatrically, but in sev-eral morphological characters they are easily distinguishable: inconsistencies of tail segments are unique for L. melanura (BAIG 1992). Although some species of Laudakia have a ten-dency to change the number of whorls in the caudal segments from the anterior to posterior side (e.g. L. nupta), the differences never exceed one whorl. In this respect, L. melanura is exceptional among all species of Laudakia in having 2, 3, 4 or 5 whorls of scales in each tail segment (L. m. melanura) or the tail annulation often indistinct (L. m. lirata). The second distinguishing character of this lizard is that the patch of callose midventral scales may be rectangular, much wider than long, while in other species of Laudakia it is usually oblong or a thin longitudinal stripe (BAIG 1992). BOULENGER (1885) distinguished lirata from melanura by having partly keeled head scales, 7–9 rows of keeled vertebral scales, and ab-sence of caudal rings.

According to SMITH (1935), enlarged vertebral scales are smooth in the northern range of this species, whereas they are more or less keeled in the southern part of the range. BAIG (1992) believes that this is not a consistent character in this species and that either habitat or age may have some influence on it. MERTENS (1969) again resurrected lirata as a subspecies of melanura on the basis of carinated scales and colour.

This lizard is one of the least known Iranian agamids and apparently there is no proper re-cord of its occurrence inside the Iranian borders but, as pointed out before, the type locality of Laudakia lirata (Blanford, 1874) is very close to the Iranian frontier. Therefore, it is ex-pected to occur along much of the Makran coast of Iran in Sistan-Baluchistan province, south-eastern Iran. SHOCKLY (1949) gives a sight record from Ras Jiunri, west Pakistan, about 16 km east of the Iranian border.

Laudakia melanura mainly occurs in Pakistan from Salt Range in NW Punjab, Waziristan in North Western Frontier Province through Sind and Baluchistan provinces, at least to the vicinity of the Iranian frontier, and most probably, some parts of southeastern Iran (Sistan-Baluchistan province) (Fig. 12). MERTENS (1969) reports both subspecies from localities very close to one another on the right of the lower Hab River in Pakistan. BAIG (1992) also reports this lizard in some localities in eastern Afghanistan. According to ANANJEVA & TUNIEV (1994), L. melanura is found along the eastern foothills of the Iranian High Plateau to the south of the Hindukush. These authors believe that the taxonomic position of L. melanura is debatable but they regard it as related to L. erythrogastra. We believe that based on ecological and morphological features L. melanura is closely related to L. nupta. Further work and the collection of more material, especially inside Iranian territory, will help to throw light upon various aspects of the biology and affinities of L. melanura.

Concluding remarks

Based on the accounts given above, we divide the Iranian Plateau species of Laudakia into two distinct groups:

1. The first group is characterized by having: almost always bi-whorled tail segments, much heterogeneous dorsal scalation, often with large spinose scales on flanks, rather short tail (always less than twice the body length), a superficial tympanum, and a conspicuous colour pattern. Species belonging to this group mainly occur in the northwestern, northern, north-eastern and eastern regions of the Iranian Plateau. This group contains L. caucasia, L. micro-lepis, and L. erythrogastra.

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Tab. 1. Metric and meristic differences (mean, standard error, and range) of all the Iranian Plateau species of Laudakia. Terminology follows Table 2. All measurements in mm.

Characters SVL TL HL HW HH SQ LVS AGFS FCWS

L. caucasia (n =89)

120.98 ±2.184

101-140.5

170.14 ±2.675

144.1-189.9

33.35 ±0.718 27-40

26.50 ±0.670

21.9-32.9

15.25 ±0.468

12-18.7

160.04 ±3.660

123-179

8.35 ±0.205

7-10

102.83 ±1.638 84-113

27.70 ±0.781 25-31

L. microlepis (n=33)

135.77 ±3.029

111.4-160

162.13 ±3.666

102-184.3

38.52 ±1.189 30-50

32.54 ±1.132

23-41.1

21.29 ±1.095

12.5-30.7

199.83 ±2.575

170-217

11.48 ±0.234

9-13

119.13 ±0.920

109-126

29.74 ±0.650 27-34

L. n. nupta (n=51)

133.54 ±3.126

103-157

254.72 ±6.522

205-321

37.48 ±0.899

29.8-44.8

31.35 ±0.870

24.2-39.4

18.31 ±0.638

12.8-23.7

104.92 ±1.703 81-122

18.84 ±0.377 17-22

94.48 ±1.398 84-110

21.48 ±0.494 15-22

L. n. fusca (n=9)

151.88 ±3.167

135.4-165

309.00 ±9.422

250-341

41.08 ±1.169

36.8-47.3

36.63 ±1.421

29.2-42.8

19.93 ±1.177

16.4-27

81.67 ±1.581 74-89

15.67 ±0.333 13-16

78.89 ±2.003 71-91

15.22 ±0.401 14-17

L. m. lirata (n=9)

132.11 ±2.831

120-146

288.67 ±5.876

269-324

29.02 ±1.819

26.1-31.7

23.91 ±0.999 20-28

14.63 ±1.211

12.9-16.9

129.44 ±3.797

120-151

7.75 ±0.250

7-9

99.56 ±1.982 96-105

17.78 ±0.278 17-19

L. m. melanura (n=6)

132.38 ±2.058

125.3-137

301.50 ±12.614 241-321

30.83 ±1.678

28.7-33.1

23.35 ±1.091

20.1-26.4

14.58 ±1.201

12.8-16.3

152.50 ±2.487

145-159

9.50 ±0.342

9-11

110.67 ±2.961

105-115

21.67 ±0.494 20-23

L. erythrogastra (n=4)

119.92 ±4.563

111-130

152.53 ±7.643

139-169

34.47 ±0.976

25.3-38.9

26.79 ±1.023

23.1-29.2

16.39 ±0.891

15.5-18.9

109.00 ±6.363

100-127

9.53 ±0.288

9-10

88.33 ±1.097 81-94

28.11 ±0.577 27-29

Tab. 2. The ANOVA-based pairwise comparison of the Iranian Plateau species of Laudakia (except L. erythrogastra ) using 9 most informative morphological characters. The table gives P-values, significance level and F-value. Significance level = 0.05; the asterisks (*) and (<) denote that the P-value is significant.

Characters SVL TL HL HW HH SQ LVS AGFS FCWS

L.caucasia (n = 89) vs L. microlepis (n = 33)

0.0003* 9.131

0.0846 3.114

0.0006* 13.865

<0.0001 21.090

<0.0001 25.689

<0.0001 79.037

<0.0001 101.286

<0.0001 75.318

0.0011* 16.092

L. n. nupta (n = 51) vs L. n. fusca (n = 9)

0.0024* 10.819

0.0001* 19.548

0.0379* 4.691

0.0036* 9.855

0.2112 1.628

<0.0001 59.643

<0.0001 22.841

<0.0001 35.231

<0.0001 52.553

L. m. melanura (n = 6) vs L. m. lirata (n = 9)

0.9451 0.005

0.3216 1.062

0.0355* 5.504

0.7201 0.134

0.9384 0.006

0.0006* 20.329

0.0011* 18.000

<0.0001 45.296

<0.0001 54.914

2. The second group is characterized by having: a very long tail (often more than twice the body length) with at least three-whorled, inconsistent, or indistinct annuli, lacking large scales on the dorsolateral region and often with a prominent longitudinal fold on this area, larger and deeper tympanum, and a colour pattern which is easily distinguishable from the first group (often sandy grey, dark brown, or black, usually not reticulated). Members of this group mainly inhabit southern, southwestern, and southeastern regions of the Iranian Plateau. This group consists of L. nupta and L. melanura.

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Taxonomic discussion

As pointed out before, MOODY (1980), in an unpublished thesis, subdivided the Agamidae into six groups. The southwest and central Asian and African radiation comprises group VI. Within this group he split the former wide-ranging genus Agama into six genera; the African Agama (about 27 species), the Asian and east African Stellio [= Acanthocercus + Laudakia] (22 species), the Middle Eastern and north African Trapelus (about 8–10 species), Xena-gama from Somalia (2 species), Pseudotrapelus sinaitus from Arabia and Brachysaura mi-nor from India. Together with the Central Asian genus Phrynocephalus, these genera are believed to represent a monophyletic group (JOGER & ARANO 1987). According to MOODY (1982: 252), these taxa are evolutionarily highly derived agamids and are not related to the primitive Uromastyx.

The nomenclatural system of the Palaearctic clade of ring-tailed agamas has been contro-versial and problematic since 1768 when LAURENTI applied the generic name ‘Stellio’ for the first time. This name is one of the oldest vernacular names known for any lizard. It has re-ceived a widespread and varying application both in pre-Linnaean and post-Linnaean times (HENLE 1995). At present, some authors use the generic name „Stellio“ for agamids of the Stellio-group of the formerly more inclusive genus Agama (e. g. ANANJEVA et al. 1990, JOGER 1991, SCHÄTTI 1989). Apart from the fact that the content of Stellio (sensu MOODY) is still debatable, the name Stellio Laurenti, 1768 is not available (BAIG 1992; BAIG & BÖHME

1991, 1997; FROST, MS; HENLE 1995; LEVITON et al. 1992). Some authors (e. g. ARNOLD 1986) retain the genus Agama (sensu lato) and recognize Stellio (and also Trapelus) as a subgenus. BAIG & BÖHME (1991) and CLARK (1990, 1991) also used Agama (sensu lato).

MOODY (1980) included in the genus Stellio not only the Palaearctic group of whorl-tailed agamas, but also the Afro-Arabian species of the Agama atricollis and A. cyanogaster groups mostly on the basis of their similarly segmented, whorl-bearing tails which he considered to be a common derived characteristic, i.e. a synapomorphy (BAIG & BÖHME 1997). According to BAIG & BÖHME (1997), there are some dissimilarities in tail structure of the Palaearctic and the Afro-Arabian clades that have proved to be constant: Afro-Arabian species generally have compressed tails with rather indistinct and sometimes even concealed whorls and seg-ments, whereas in the members of the Palaearctic clade the tail (and also head and body) is mostly depressed and subdivided into distinct whorls and segments which provide stable key characters for distinguishing different species. Based on all the available evidence, the same authors split the genus Stellio into two distinct genera: Acanthocercus Fitzinger and Lauda-kia Gray. The former includes all members of the Afro-Arabian clade and the latter consists of the Palearctic clade.

LEVITON et al. (1992) believe that Stellio is unavailable and suggest adopting Laudakia as the name for agamids of the Stellio-group. FROST (MS, pers. comm.) argues that the nominal generic name Laudakia Gray, 1845 applies to the Asian species if, as JOGER (1991) suggests, A. adramitana, A. yemensis and, by implication, A. cyanogaster are more closely allied to A. atricollis (which JOGER treats as a sister-group of Pseudotrapelus) than to the Asian mem-bers of the genus.

Some authors (e. g. FRANZEN & SCHMIDTLER 1993) followed LEVITON et al. (1992) and used Laudakia for the western Palaearctic rock-dwelling agamids. There are two possible alternatives for the nomenclature of the western Palaearctic agamids: either to retain all species groups within the genus Agama (sensu lato), or, if one prefers to elevate them to full genera, to find another name for the Stellio-group. None of these two approaches is com-pletely satisfactory in view of our current knowledge of the phylogeny of agamids (HENLE

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1995). If one accepts the first approach, then one should incorporate Phrynocephalus in Agama (s. l.), based on the biochemical evidence provided by ANANJEVA & SOKOLOVA (1990) and JOGER (1991). Furthermore, the first approach would conceal the advances gai-ned in determining the phylogeny of agamids. In discussing the problems of the second approach, we should first look at the results of MOODY (1980). As mentioned above, MOODY split the former more inclusive genus Agama into six genera. The genera Agama, Trapelus, and Pseudotrapelus are characterized by a derived karyotype (2n = 44–48 chromosomes) while Stellio retains the ancestral karyotype (2n = 36 chromosomes, except for S. tubercula-tus with 2n = 34). Xenagama and Brachysaura have not yet been examined. All of these genera have their own distinguishing characters: Trapelus is characterized by a small and deeply sunk tympanum which forms an external auditory tube-like meatus, Pseudotrapelus by a swollen tail and a large tympanum, while the Stellio-group is the only genus having several spinose scales located immediately behind the tympanum or on the sides of the head (HENLE 1995), lacking dorsal crest and gular sac. Following on from this discussion, there are several alternative available generic names for the Stellio-group:

1. The first alternative is Acanthocercus Fitzinger, 1843, which is based on Stellio cyano-gaster (Rüppell, 1835), a member of the African clade. This clade is certainly not congeneric with the Palaearctic clade (JOGER 1991, BAIG & BÖHME 1997). According to the phylogenetic tree provided by JOGER (1991), it may be regarded as congeneric with Pseu-dotrapelus Fitzinger, 1843, or as a sister genus of the latter (WELCH 1994).

2. The second available generic name, is Laudakia Gray 1845, which, as mentioned before, has been adopted by LEVITON et al. (1992), BAIG & BÖHME (1997) and FROST (MS). The type species of Laudakia is Agama tuberculata (GRAY 1845: 254) belonging to the Palaearc-tic clade, and the name may be used for the Stellio-group (sensu stricto) although there are problems in using this approach: Laudakia tuberculata differs considerably in hemipenial characters (apomorphic characters: divided hemipenes, forked sulcus, and transverse division of the apex) from the Stellio-group (s. s.) which is closer to Agama (s. s.) (BÖHME 1988). Furthermore, as mentioned above, L. tuberculata has a derived karyotype (2n = 34 versus 2n = 36 which is the ancestral karyotype of the Stellio-group) lacking one pair of microchromo-somes (GORMAN 1973). Furthermore, according to JOGER (1991), based on albumin antisera and isozymes, the genus Phrynocephalus is a sister group of the Stellio-group (s. s.). JOGER (1991) did not have available data for L. tuberculata, and its phylogenetic position relative to Stellio (sensu stricto) and to Phrynocephalus remains unknown. Also, according to BÖHME (1981), Phrynocephalus has a derived karyotype (2n = 44–48) as in Agama, Trapelus, and Pseudotrapelus, but differs considerably from Laudakia. However, it remains unresolved whether the similarities between Phrynocephalus, L. tuberculata, and Trapelus are synapo-morphies or are due to convergence (BÖHME 1988).

3. The third possible substitute name for the Stellio-group (s. s.) is Plocederma Blyth 1854, based on Agama melanura (BLYTH 1854: 738). HENLE (1995) is of this opinion that the best solution is to restrict the use of Laudakia to L. tuberculata, which leaves open the question of a genus name for the Stellio-group. He believes that Plocederma Blyth, 1854, is the oldest available name for the Stellio-group (s. s.) and should be used for it, and species for which it is not known whether they are more related to the Stellio-group or to L. tuberculata may be referred to either Laudakia or Plocederma. Of the Iranian species, this author classifies L. caucasia, L. melanura, L. nupta, and L. erythrogastra in the genus Plocederma but places L. microlepis either in Laudakia or Plocederma (HENLE 1995: 8).

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Tab. 3. Comparison of L. c. caucasia and L. microlepis; L. n. nupta and L. n. fusca; and L. m. melanura and L. m. lirata (based on material examined as well as literature, e.g. ANDERSON 1966a, 1974, 1999, BAIG 1992).

Character L. c. caucasia L. microlepis Callose scales usually only in males often in both sexes Enlarged scales on flanks always present sometimes absent Spinose scales of neck and ear distinctly developed often weakly developed Dorsal pattern often strongly reticulated darker and weakly reticulated Habitat elevation often 1500-2700 metres often 1800-3400 metres

L. n. nupta L. n. fusca Cutaneous fold across neck strongly developed rudimentary or absent Spinose scales around ear distinctly developed exceptionally developed Head colouration in males variable yellowish Dorsal colouration variable often darkbrown to black

L. m. melanura L. m. lirata Caudal rings present almost absent Dorsal scales weakly keeled or smooth strongly keeled Head scales smooth partly keeled Colouration almost always entirely black or

brownish-black often head and shoulders yellow-ish-sandy coloured

Until such time as these systematic and phylogenetic problems are resolved (by studying additional species and specimens from all over the range of this group, and by combining morphological, karyological, biochemical, palaeontological, and zoogeographical ap-proaches), it can be expected that different authors will apply various generic and subgeneric names to the Palaearctic rock-dwelling species of ‘Agama’ lizards. In view of this lack of resolution, and with some trepidation, we have adopted LEVITON’s et al. (1992), BAIG & BÖHME’s (1997), and FROST’s (MS, pers. commun.) approach in this paper, as the phyloge-netically most comprehensive alternative so far, and apply Laudakia Gray, 1845, to the Asian clade of ‘Agama’ lizards until recently placed in the genus or subgenus ‘Stellio’. But we do not reject the possibility once more evidence is provided in favour of another alterna-tive (e.g. Plocederma Blyth) of adopting the most phylogenetically comprehensive substitute name, and we believe that Plocederma Blyth, 1854, is the oldest available and most compre-hensive substitute name for Laudakia, if it can be proved by all the available methods that L. tuberculata has several species-specific autapomorphies. In this case the genus Laudakia would be monotypic, comprising only L. tuberculata as the type species of Laudakia Gray, 1845, and all the other species of the Palaearctic rock-dwelling agamas would come under the generic name of Plocederma Blyth, 1854. For detailed discussion and the historical background for the use of ‘Stellio’ in lizard systematics and nomenclature, readers are re-ferred to HENLE (1995), BAIG & BÖHME (1997), and FROST (MS).

Historical biogeography

The present topography of the Iranian Plateau and adjacent areas is attributed to the fragmen-tation of the Gondwanaland, the northward movements of its fragments, the Indian and Ara-bian Plates, and their collision with the Eurasian continent. This collision caused very inten-

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se orogenies which resulted in the formation of huge mountains from southern Europe in the west, to India, Mongolia, and China in the east. A number of these huge mountain chains (Zagros, Alburz, and Kopet-Dagh) bordering the Central Iranian Plateau, have played a crucial role in the radiation, fragmentation, and subsequent evolution of the Iranian Plateau species of Laudakia.

Soon after the development of acrodont dentition in the upper Cretaceous, about 80 MYBP (BORSUK-BIALYNICKA & MOODY 1984), the Acrodonta subdivided into two clades, Chamae-leonidae and Agamidae. Fossil evidence suggests that this early radiation took place in east Asia, but Chamaeleonidae must have invaded Africa and Madagascar soon afterward (KLA-

VER & BÖHME 1986). According to CRACRAFT (1973b, 1975), agamids and chamaeleonids diverged after the separation of South America and Africa in the middle Cretaceous. In Asia the Agamidae diverged into several groups, two of which, Uromastyx and the Agama group, had their initial radiation in western Asia. When the Afro-Arabian Shield was connected to Asia after subdivision of the Tethys Sea, about 18 MYBP (JOGER 1987), these groups inva-ded Arabia and four genera, Uromastyx, Trapelus, Pseudotrapelus, and Agama finally rea-ched Africa during the Miocene. Phrynocephalus and Laudakia, however, extended their main radiation into Central Asia, the former genus adapted to terrestrial life, and the latter to saxicolous life (JOGER 1991). The fossil records show that the ancestors of Laudakia were widely distributed until the Miocene (25 MYBP), in the west as far as France (Agama galliae Filhol, 1877), and in the east as far as the eastern part of China (Agama sinensis Hou Lian-Hai, 1975). According to ESTES (1983), a primitive late Cretaceous agamid Mimeosaurus was present in Central Asia; Eocene records refer to the primitive agamid Tinosaurus as occurring in southern Asia, and Agama (Stellio) is recorded from the Paleogene on into the Holocene in Europe. BAIG (1992) suggests that these ancestral taxa survived until the Pleis-tocene (1.6 MYBP). During glacial and interglacial times, drastic climatic changes took place on the earth; many parts of the world including northern Europe were covered with glaciers. During the Tertiary, major surface changes occurred and different mountain ranges formed. The collision of the Indian Plate with Asia (45–50 MYBP) extruded blocks in Af-ghanistan to the southwest and blocks in Tibet to the east. The Pamir, Karakorum, Himalaya, Hindu Kush, and Tien Shan all underwent substantial uplifting (MOLNAR & TAPPONNIER 1977). Until the Miocene the mountain ranges were fairly high and gave rise to different types of habitat. These habitats have their own peculiarities which initiated evolutionary changes in three different directions. Laudakia, Trapelus, and Phrynocephalus are the pro-duct of this habitat-oriented evolution (BAIG 1992). During the process of mountain and desert formation, one group developed characteristics suitable for mountain-dwelling (Lau-dakia), the other group adapted itself for life in deserts (Phrynocephalus), and the intermedi-ary habitat was occupied by Trapelus (BAIG 1992).

The centre of origin for Laudakia is hypothesised as the area with the highest degree of endemism (cf. also BAIG 1992, DARLINGTON 1957, MOODY 1980). In the case of Laudakia, we know that the majority of species occur in the high mountainous regions of NE Afghanis-tan, N Pakistan, and SE Tajikistan (around the Pamir Knot). It would thus be logical to re-gard this area as the centre of origin for Laudakia. This area (the Pamir Knot) fulfills almost all the proposed standards for the centre of origin (ANANJEVA & PETERS 1990, ANANJEVA & TUNIEV 1994, BAIG 1992). Almost all herpetologists who have studied the taxonomy and biogeography of Laudakia are unanimous in the opinion that speciation of these agamids has been closely linked with the Hindu Kush, Pamir, and Himalaya Mountains (ANANJEVA et al. 1981, ANANJEVA & PETERS 1990, ANANJEVA & TUNIEV 1994, BAIG 1992, PETERS 1971). BAIG (1992) proposed a dispersal-based scenario for the origin and subsequent evolution of

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Laudakia (including the Iranian Plateau species); he argues that the ancestor of Laudakia originated and evolved around the Pamir Knot and, with time, various waves of dispersal along the high mountain systems of southern and central Asia took place and extended the range towards the northeast (e.g. Laudakia stoliczkana), southeast (e.g. L. tuberculata), southwest (e.g. L. nupta and L. melanura), and western mountainous regions (e.g. L. cauca-sia and L. stellio).

It is noteworthy that based on molecular data MACEY et al. (1998) attributed the fragmen-tation of populations of the Laudakia caucasia species group on the Iranian Plateau to a north-south vicariant event. They believe that the north-south vicariant event was caused by the rise of the Zagros Mountains (10–5 MYBP), and for the divergence of L. microlepis from the northern populations of Laudakia caucasia group they address a date of ~9 MYBP. Though the occurrence of a vicariant event such as the formation of the Zagros Mountains might have been crucial for the divergence of the southern species (e.g. L. microlepis) from the northern ones (e.g. L. caucasia), the present distribution of most species of Laudakia fits very well into a pattern of dispersal along the high mountain chains (e.g. Zagros, Alburz, and Kopet-Dagh). We therefore believe that the origin, radiation, subsequent evolution, and present distribution of various species of Laudakia can be attributed both to vicariant events (e.g. orogeny) and to dispersal waves along the high mountain ranges.

The allozyme-based phylogenetic hypothesis for the agamid genus Phrynocephalus argues for the ancient divergence of agamids of the Afro-Asian radiation including Laudakia (MACEY et al. 1991, 1992). It would be logical to assume that the first stages of this radiation took place in the Miocene on the present territory of Afghanistan, Pakistan, and Tadjikistan.

On the Iranian Plateau, two distinct branches of the genus Laudakia evolved:

(1) A southern, and most likely more ancient, branch including L. nupta and L. melanura species complexes. These two taxa, mainly occurring in the southern parts of the range of Laudakia, are sympatric in some regions of SW Pakistan and probably of SE Iran too. It is very likely that these two taxa have been derived from a single ancestor which, in turn, originated from the ancestors of Laudakia in the hypothetical centre of origin (Pamir Knot), and then radiated along the Hindu Kush towards the south and southwest. After reaching the southernmost part of the range on the Makran coast, speciation occurred in the ancestral agamid giving rise to L. nupta and L. melanura. The latter species has its main distribution in southern and central Pakistan. It barely extends its range into Iranian territory proper and Afghanistan, and this may be due to the presence of the other congeners as strong competi-tors. Laudakia nupta invaded the mountainous regions of southeastern Iran, along the Makran coast, and extended its range further towards the western and southwestern parts of the Iranian Plateau along the Zagros Mountains, as far west as eastern and northeastern Iraq. This invasion might have begun in the late Miocene, and during the Pliocene L. nupta reached eastern and northeastern Iraq as well as southwestern Iran as the only representative of Laudakia in the latter region. The wide range of L. nupta (from eastern Iraq to Pakistan and Afghanistan) is presumably an indication of its long history. In a mitochondrial DNA-based phylogenetic analysis of the west Asian and African clade of Acrodonta, the most parsimonious tree derived from the analysis of the informative DNA sequence characters showed that L. nupta is in a basal position relative to the other Laudakia species of the Irani-an Plateau (MACEY et al., 2000). Unfortunately, no molecular data were available for L. melanura. (2) A second branch,mainly occupying northwestern, northern, and central parts of the range of Laudakia. This branch consists of L. caucasia, L. erythrogastra, and L. microlepis. As

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noted before, almost all these taxa have a bi-whorled and rather short tail [with the exception of the newly described L. c. triannulata (ANANJEVA & ATAEV 1984) which often has an extra whorl on the sides of the ] with much heterogeneous dorsal scalation, smaller and more superficial tympanum, and more depressed body and tail (in comparison with the first branch). It is obvious that the ancestors of this group have undergone different evolutionary changes which, with time, have given rise to these taxa.

Laudakia erythrogastra presumably derived earlier, and occupied the mountain ranges of Kopet Dagh and Hindu Kush (NE Iran, SE Turkmenistan, and NW and central Afghanistan). It does not penetrate into the interior regions of the Iranian Plateau and this may be due to the presence of L. caucasia in the immediate west and L. microlepis and L. nupta in the southern parts of its range as strong competitors.

The present distribution of L. caucasia and L. microlepis illustrates a biogeographic prob-lem, since we can attribute their patterns of distribution both to fragmentation along the high mountain ranges of the northern and southern regions of the Iranian Plateau (due to dispersal waves) as well as to a drastic vicariant event (e.g. the huge orogenic phenomena of the late Miocene and Pliocene) which interrupted the contiguous range of their ancestors, and since then the vicariance-based fragmented populations have remained in isolation giving rise to two distinct taxa, i.e. L. caucasia and L. microlepis. The former taxon occupied the northern parts of the range and by further dispersal invaded towards the west and south; in the west it reached the high mountain systems of the Caucasus and in the south as far as the central Zagros Mountains. The latter taxon mainly invaded the southern and southeastern regions and in turn, via dispersal waves, extended its range towards the west as far as the south-central mountains of the Iranian Plateau where it has a limited contact with the southernmost populations of L. caucasia as well as being sympatric with L. nupta. With regard to the rela-tive importance of both dispersal waves and vicariant events, it is clear that the Alburz, Ko-pet Dagh, and Zagros Mountains have played a crucial role in the isolation, radiation, and subsequent evolution of L. caucasia and L. microlepis as well as L. erythrogastra.

ANANJEVA & TUNIEV (1994) believe that the populations of L. caucasia from the north of the Suleiman Mountains (the southern slope of the Hindu Kush) are likely to be the most ancient, and western populations of this taxon distributed from the Turkmen-Khorasan mountains, westward through Iran, to Transcaucasus seem to be more recent, and probably in the Pliocene L. caucasia reached the northern slopes of the great Caucasus.

According to ANDERSON (1968), the rock-dwelling species of the Iranian Plateau Lauda-kia may illustrate the results of climatic fluctuations on the Iranian plateau and because of these fluctuations, which took place during the Tertiary and Quaternary, certain forms were distributed to the north (e.g. Laudakia caucasia and L. erythrogastra) while the others bor-dered the plateau on the west and south (e.g. Laudakia nupta and L. melanura) and the inner mountains were occupied by L. microlepis.

ANANJEVA & TUNIEV (1994) divided the Asian rock agamids into two complexes of spe-cies with a number of morphological and ecological differences: ‘caucasia’ complex and ‘himalayana’ complex. Among the Iranian Plateau species, L. caucasia, L. microlepis, and L. erythrogastra belong to the former, and L. melanura, and, apparently, L. nupta to the latter complex. These authors proposed a scenario similar to the one adopted by BAIG (1992) for the origin and subsequent evolution of Laudakia species.

In summary, we conclude that the current patterns of distribution of the Asian rock-dwelling agamids are the result of orogenic phenomena and climatic fluctuations which took place during the Tertiary (huge orogenies of the Miocene and Pliocene) and Quaternary (the glacial and interglacial periods of the Pleistocene), and both dispersal waves and vicariant

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events have had an effect on divergence and speciation in the genus Laudakia. Such are the complexities of geological and climatic history on the Iranian Plateau that it is impossible to produce a detailed description of palaeobiogeographic events, especially as there is no known fossil record of present day agamids in this area. All that can be done is to describe the patterns and probable mechanisms which may have produced them.

Key to the Iranian species of Laudakia

1a. Flanks without enlarged scales or tubercles; tail with segments composed of more than two annuli when viewed laterally, or segmentation indistinct. ...................................................................... 2

1b. Dorsal scales heterogeneous, back and usually flanks with scales of varying sizes inter-mixed; tail with segments often composed of two annuli and its length almost always less than 1.8 of body length; dorsal pattern more or less reticulated. .................................................................... 4

2a. Head subcordiform; median dorsal scales in straight longitudinal series, usually strongly keeled, 7–10 across middle of back, grading into small dorsolateral scales; often 120–151 scales around mid-body; a rudimentary nuchal crest often present; caudal segments inconsistent in number or the segments indistinguishable; patch of callose midventrals often rectangular; head and shoulder often brightly coloured; hemipenes of male whitish. L. melanura lirata (Blanford, 1874)

2b. Head distinctly triangular; median dorsal scales in oblique longitudinal series, 14–22 across middle of back, clearly set off from dorsolateral scales; often 81–122 scales around mid-body; caudal segments with three whorls of scales; hemipenes of male black; dorsolateral fold usually well developed; tympanum larger than orbit and weakly sunk. .............................................................. 3

3a. A prominent transverse fold of skin across nape; often more than 90 scales around middle of body; proximal caudals not exceptionally large, often more than 17 on the fifth whorl behind the vent. ................................................................................................ L. nupta nupta (De Filippi, 1843)

3b. No fold of skin across nape, or the fold vestigial; often less than 90 scales around middle of body; proximal caudals very large, often less than 17 on the fifth whorl behind the vent; males often yellow-headed................................................................................ L. nupta fusca (Blanford, 1876)

4a. One or two longitudinal rows of clusters of spiny tubercles on each side of body; 78–129 scales around middle of body; gular scales strongly keeled and mucronate. ............................................................................................................. L. erythrogastra (Nikolsky, 1896)

4b. Body strongly depressed; enlarged scales on flanks not arranged in longitudinal rows, or may be absent; almost always more than 125 scales around middle of body; gular scales smooth, not mucronate; tympanum not larger than orbit, distinctly superficial. ..................................... 5

5a. Body size variable; often 120–185 (usually less than 175) scales around middle of body; often 7–10 longitudinal rows of enlarged vertebral scales; usually only males with callose preanal and midventral scales; tail length often more than 1.35 of body length. .....................................................................................................................L. caucasia (Eichwald, 1831)

5b. Body stout; 180–239 scales around mid-body; often 9–13 longitudinal rows of enlarged vertebral scales; patches of callose midventral and preanal scales almost always present in both sexes, strongly developed in males, occupying a large area; tail length often less than 1.35 of body length ............................................................................................... L. microlepis (Blanford, 1874)

Acknowledgements. We wish to thank the Razi University authorities (Kermanshah-Iran) as well as ESKAN-

DAR RASTEGAR-POUYANI, ALIREZA HASHEMI, SEYFULLAH AZADI, MASOUD ABRISHAMIAN and ALI RASSULI for their unstinting help during field work on the Iranian Plateau. We would also like to thank: Dr. ALAN E. LEVITON, EVON HEKKALA and JENS VINDUM, California Academy of Sciences, San Francisco, California; Dr.

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GUNTHER KÖHLER, Senckenbergmuseum, Frankfurt; OLAVI GRÖNWALL and S. O. KULLANDER, Swedish Museum of Natural History, Stockholm, for the loan of Laudakia specimens; Prof. S. C. ANDERSON, Univer-sity of the Pacific, Stockton, California, for all his recommendations and encouragement, and for sending informative literature, especially his unpublished manuscript concerning the Iranian Plateau agamids, as well as for critically reviewing this manuscript; Prof. Dr. WOLFGANG BÖHME, Museum Alexander Koenig, Bonn, and Dr. KHALID JAVED BAIG, Pakistan Natural History Museum, Islamabad, for providing us with informative literature; Dr. COLIN MCCARTHY, The Natural History Museum, London, and Dr. IVAN INEICH, Muséum National d’Histoire Naturelle, Paris, for providing the checklist of Laudakia .

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Authors’ addresses: Dr. Nasrullah Rastegar-Pouyani, Department of Biology, Faculty of Sci-ence, Razi University, Kermanshah, Iran. E-mail: [email protected] – Dr. Göran Nilson, Department of Zoology, Gothenburg University, Box 463 SE-405 30, Gothenburg, Sweden.

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Appendix: Material examined

Laudakia caucasia caucasia: CAS 182810-13: Buinaksky District (42°58’N, 47°30’E), Daghestan Autonomous Rep., Russia; CAS 194304-8: Lake Seivan (40°30’N, 45°30’E), Armenia. SMF 10160-1: Karabagh, Armenia; SMF 10162-3: Salmas, NW Iran; SMF 44054-5: Turkey; SMF 61198: Tiflis, Georgia Republic; SMF 64686: Tiflis, Georgia Republic; SMF 66251: Shahpasand, NE Iran; SMF 67909-11: Kala-i-Chambar, Badgis, Afghanistan; SMNH 3126(1-6): Tehran, Iran. GNHM Re.ex.5497-5505: Kermanshah prov., 45 km NE Kermanshah on the Zagros Mountains, at about 2400 m elevation; GNHM Re.ex.5506-08: Kermanshah prov., 65 km NE Kermanshah on the Zagros Mountains, at about 2300 m elevation; GNHM Re.ex. 5509-10: Central Iran, Esfahan prov., 20 km S Golpaygan on road to Khansar, Vaneshan region, about 2000 m elevation; GNHM Re.ex.5511-12: Kermanshah prov., 55 km NE Kermanshah on the Zagros Mountains, Siahdarreh village, about 2200 m elevation; GNHM Re.ex.5513-15: Central Iran, Esfahan prov., 3 km N Fereydun-Shahr on the central Zagros Mountains, 2450 m elevation; GNHM Re.ex. 5516-17: Kermanshah prov., 60 km NE Kermanshah on the Zagros Mountains, near Gerdakaneh village; GNHM Re.ex.5518: NE Iran, Semnan prov., northern Dasht-e-Kavir; GNHM Re.ex.5519-20: Kermanshah prov., 40 km NE Kermanshah on the Zagros Mountains, 2100 m elevation. INNHM1245, Alburz Mountains, N Iran; INNHM483: N of Qazvin, N Iran; INNHM1490: 29 km W Firuzkuh to Damavand, N Iran; INNHM189: Jajrud, N Iran; INNHM482: NE Qazvin, N Iran; INNHM112: Alburz Mountains, N Iran; INNHM12: Sarband; INNHM1295-6: Mesh-kin Shahr, NW Iran; INNHM1329-31: 21 km E Maku on the road Marand, NW Iran; INNHM117, 479-80, 541-5,1158-62: Golestan National Park, NE Iran; INNHM 27, 304-10: N of Shahrud, NE Iran; INNHM525: Kalaleh, Gorgan, NE Iran.

Laudakia c. triannulata: CAS 184640-44: 2 km SW of Madau, Krasnovodsk Region, Turkmenistan.

Laudakia microlepis: CAS 143282-5: Baluchistan, Iran. ZISP 20628; GNHM Re.ex. 5135-39 and 5140-41 (topotypes): Khaneh-Sorkh Pass, between Sirjan and Kerman, Kerman prov., SE Iran. SMF 63205: Ziarat, Quetta, Pakistan. SMF 64222-4: Mansehra, Pakistan. INNHM1395-6: Kuh-e-Shah, N Baft, SE Iran; INNHM1398-9: Kuh-e-Taftan, SE Iran; INNHM1169, 1173: 36 km S Bidokht on road to Birjand, NE Iran; INNHM1170-72, 1235: between Birjand and Turbat Haydareyeh, NE Iran; INNHM1163-8: 8 km S Robat-e-Sang on road to Torbate Haydareyeh, NE Iran.

Laudakia erythrogastra: CAS 184400-3: Turkmenistan-Iran border, foothills of the Kopet-Dagh Mountains, Ashghabad Region, Turkmenistan.

Laudakia melanura lirata: SMF 63206-9: Insel Astola, Pakistan; SMF 63210: Band Murad Khan, Pakistan; SMF 63213-16: Kirthar Range, Pakistan.

Laudakia m. melanura: CAS 99861-63 and CAS 99987: Hub Chowki, Lasbela, Pakistan. SMF 63211-12: Hab-Chawki, near Hab River, Pakistan.

Laudakia nupta fusca: SMF 63219-20: Hab-Chauri and Band Murad Khan, Pakistan; SMF63224-5: Siah Kuh, S of Delbandin, Pakistan; SMF63227-30: near Sand, Pakistan. CAS 99986: North Karachi, Khadeji Water Falls, Pakistan.

Laudakia n. nupta: GNHM Re. ex.5521: Kermanshah prov., 5 km E of the Iranian-Iraqi border, W Iran; GNHM Re. ex. 5522-45: Zagros Mountains, Kermanshah prov., W Iran; GNHM Re. ex. 5546: 70 km N of Qum on the road to Tehran, Markazi prov., north-central Iran; GNHM Re. ex.5547-48: Zagros Mountains, Hamedan prov., W Iran; GNHM Re. ex. 5549: 155 km N of Esfahan on the road to Saveh, Markazi prov., central Iran; GNHM Re. ex.5550-52: 65 km E of Ramsheh on the road to Shahreza, Esfahan prov., central Iran; GNHM Re. ex. 5553-60 (all topotypes): 45 km NE of Shiraz, Persepolis (type locality), Fars prov., south-central Iran. INNHM1408: Khorram Abad, SW Iran; INNHM 221: Mohammad Zaman, SW Iran; INNHM 476: Do- Rud, SW Iran; INNHM 561: Muteh protected region, Esfahan, central Iran; INNHM1035: 41 km SE Bandar-e-Rig on road to Borazjan, S Iran; INNHM 815-6: 136 km Bandar Abbas-Kerman road on road to Bandare Lengeh, S Iran. SMF 63217-18: Barshore, Quetta, W Pakistan; SMF 63221-2: Quetta, W Pakistan.

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