Volume 6, No. 2, May-August 1999 ISSN 1026-2296 Russian Journal of Herpetology

Article reprints

Folium Publishing Company Russian Journal of Herpetology Vol. 6, No. 2, 1999, pp. 1 2 5 -1 4 2

ZOOGEOGRAPHIC ANALYSIS OF THE HERPETOFAUNA OF SOUTH-WESTERN TURKMENISTAN

Boris Tuniyev,1 Goran Dusej,2 and Borje Flardh3

Submitted August 20, 1998.

This article gives a complete overview about the reptiles of western Turkmenistan and their zoogeographi- cal relation. The data were obtained by several expeditions during the last years and completed by compi­ lation of the specific herpetological literature about this area. Ten general landscape units are described along a profile from the Central Karakum desert throughout the Malyi Balkhan chains, the Kyurendag Corridor, the Western Kopetdagh to the state border of Iran. The mountain and plain reptile fauna is ana­ lyzed. After discussing the faunal element composition of various landscapes, the Western Kopetdagh was included into the South-West Asian province while the Turanian plain and the Malyi Balkhan were in­ cluded into the Turanian province of the Palearctic.

Key words: Herpetofauna, South-Western Turkmenistan, Biogeography.

INTRODUCTION herpetofauna of the Western Kopetdagh (Skalon 1982; Atayev 1987; Atayev et al. 1991). Due to a long period of investigation the During the last 40 years some new reptile taxa herpetofauna of western Turkmenistan is quite well from western Turkmenistan have been described: known. The first data were published at the end of the Euphlebaris turcmenicus Darevsky 1978 from 19th century by O. Boettger (1888) and N. Zarudnoy the Monjukly ridge; (1889- 1890). Nikolsky (1915, 1916) presented ma­ Eremias strauchi kopetdaghica Shcherbak 1972 terial after collection and examination. Chernov from the southern slope of the Western Kopetdagh; (1934) published the first general survey on the rep­ Coluber atayevi Tuniyev and Shammakov 1993 tiles of Turkmenistan. Later investigations were from the Sayvan-Nokhur plateau of the Western added in a fundamental monograph by Bogdanov Kopetdagh. (1962). A number of articles about reptiles of the The following new findings were recorded: Malyi Balkhan and other western marginal mountain 1951: Lacerta strigata in the Atrek river valley, ridges of Turkmenistan were published by Shamma­ western Turkmenistan (Bogdanov 1956); kov (1964a, 1966, 1969). In 1968 a vast publication 1964: Cyrtopodion spinicauda in the Central of Anderson described the lizard fauna of Iran and the Kopetdagh, 1968 in the Western Kopetdagh and area adjacent to Turkmenistan. In the early nineties 1985 in the Malyi Balkhan (Atayev et al. 1968; two monographs were published: Shammakov Rustamov and Atayev 1976; Shammakov and Atayev (1981) wrote about the lowland reptile fauna and 1987); Atayev (1985) about the mountainous reptiles. The Coluber schmidtii was found in the early sixties monographs of Shcherbak (1974), Shcherbak and in the Western Kopetdagh (Shammakov 1964b); Golubev (1986) and Eremchenko and Shcherbak 1970: Elaphe dione in the Atrek river valley (1986) provide additional taxonomical and ecologi­ (Bogdanov 1970) and Phrynocephalus maculatus at cal data about species distributed in Turkmenistan. the Bami railway station (Bogdanov et al. 1974). Some recent articles are devoted specifically to the Despite the extremely various and changeable 1 Caucasian State Nature Biosphere Reserve, ul. K. Marksa 8a, natural conditions of Western Turkmenistan, nobody Sochi 354341, Russia. investigated and compared the herpetofauna along a 2 Bureau for Ecological Research, KaserlistraBe 12, CH-8919 profile crossing Western Turkmenistan from the sand Rottenschwil, Switzerland. desert Karakum southward to the ridge Monjukly 3 Kasmyra v. 54, S-14733 Tumba, Sweden. which borders Iran. Therefore, the zoogeographical

1026-2296/99/0602-00125 © 1999 Folium Publishing Company 126 Boris Tuniyev et al.

KAZAKHSTAN

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Kaplankur Darganata

lSNOVODSK ^nguz Karakum B. Balkhan .CHARDZHOU

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Stabilized Semi-stabilized Draft sands The Karakum Investigated sands sands Canal area

Fig. 1. A schematic map of Turkmenistan (from Babaev, 1994).

status of this region as a whole and of each of its dis­ We compared the zoogeographical areas of the her- tinct landscapes has not been studied yet. petofauna of Western Turkmenistan according to the different representation of these groups in the studied MATERIAL AND METHODS areas.

Comparison of the ten landscape units bases of PHYSIOGRAPHY AND DESCRIPTION several expeditions in this area during the last 5 years OF THE STUDIED AREA by the first author and a joint Swiss-Swedish-Rus- sian-Turkmenian expedition from April to May 1994, According to the climatic peculiarities and vege­ which has covered the main areas of the western tation the arid regions of Turkmenistan were divided Turanian lowland and the Turkmeno-Khorasan into northern deserts of the Central Asian type and mountains (Fig. 1). into southern deserts of the Mediterranean type Examination sites were chosen according to (Gvozdezky and Mihaylov, 1987). Our study covers altitudinal and ecological principles and were de­ the southern zone of the Turanian lowland as well as scribed by vegetation typology: the whole region af­ the higher area of the Western Kopetdagh mountains ter Korovin (1934), the Western Kopetdagh after Fet and Malyi Balkhan. and Kamahina (1982) and Neshatayeva (1985), the The Western Kopetdagh is situated in the north­ Karakum desert after Kalenov and Muhamedov western part of the Turkmeno-Khorasan mountains, (1992). (We made corrections for local conditions in westward of the village Nokhur (Babayev and Dur- each vegetation type in all plots). Plants were defined dyev, 1982). It differs from the Central and Eastern after the guide of Nikitin and Geldikhanov (1988). Kopetdagh in a relief and landscape of comparatively The reptile species were divided into ecological- young origin and in complex geological and geomor- geographical (faunal) groups according to modem phological structures. Small mountains and broad chorology (which one) and ecological characteristics. longitudinal valleys (Hodjakala, Sumbar, Chandyr) Zoogeographic Analysis of the Herpetofauna of South-Western Turkmenistan 127 cover the area westwards to the highest peak (Uch- from 148 mm near the Malyi Balkhan up to 328 mm kui, 1900 m) of the eastern part of Western Kopet­ in the Sumbar river valley (Babayev and Durdyev, dagh. The Peredovoy range is steep on its northern 1982; Orlovsky, 1992). side and has a broad plateau-like crest on the southern The studied area was divided into 10 landscape slope (Sayvan-Nokhur plateau). The range Monjukly units (Fig. 2) which are described below: with an altitude of about 1000 m lies behind the Unit I and I*. The northern slope of the range Sumbar river valley and near the borders of Iran. Monjukly is up to 1200 m and the southern slope of High belts of vegetation are observed in the Western the Vodorazdelny ridge up to 800 m. The vegetation Kopetdagh (from wormwood deserts in the foothills consists of subtropical Submediterranean shibliaks up to a mountain-steppe on the upper limits of the which are formed by Paliurus spina-christi, Punica ridges). granatum, Celtis caucasica, and partly of Zyzyphus According to its geological peculiarities, the jujuba, Ficus carica, Jasminum fruticans. This is a Malyi Balkhan must be directly included in the quite well watered area with several springs and mountain system of the Kopetdagh like its marginal small creeks. Near the watercourses a dense vegeta­ northwestern link (Gvozdezky and Mihaylov, 1987). tion exists consisting of Prunus divaricata, Cratae­ The Malyi Balkhan reaches 779 m of altitude and is gus sp. var., Rubus anatolicus, and Vitis sylverstris. covered by wormwood-salwort deserts of the south­ The woodless space is covered with xerophylous ern type. shrubs. Between the northern slope of the Malyi Balkhan Unit II. The Sumbar river bed, in 350 - 400 m and the dried-out river-bed of the Uzboy (the ancient altitude. Derivates of the so-called tugay (riparian or river-bed of the Amu Darya) lies the western part of gallery-forest) are found here. A formation of Popu- the Central Karakum, the biggest sand desert of Mid­ lus euphratica with Elaeagnus orientalis, Ulmus car- dle Asia. This desert was formed by alluvium of the pinifolius and grass floor of Lolium rigidum develops ancient river Amu Darya. Its aridization increased on the first terrace. On the second terrace grow Ta- about 70,000 years ago, when the Amu Darya turned marix meyeri formations with partly Tamarix laxa, northward to the Aral sea (Babayev and Zonn, 1992). T. florida, Phragmites australis and Arundo dortax. Various versions of ephemera and ephemera-desert- On the third terrace are units of semideserts consist­ bushes present the modem vegetation of the Kara­ ing of distributed sand and clay. They have an anthro­ kum desert (Kalenov and Muhamedov, 1992). pogenic origin and are represented by Haloxylon Southwards, the Kyurendag Corridor separates aphyllum and Aellenia subaphyla. the Malyi Balkhan from the Western Kopetdagh. Unit III. The upper belts of the Vodorazdelny This is a clay plain which continues further to the east ridge (1200 - 1600 m). The crest of the Vodorazdel­ along the whole Kopetdagh. The major part of the ny ridge is covered with mountain-steppe vegetation. clay lowland is characterized by wormwood-salwort Dominating species are Festuca vallessiaca, Stipa and wormwood-ephemera desert. The central lowest crassiculmis, S. capillata, S. lessingiana, Elytrigia part of the Kyurendag Corridor consists of so called trichophora, Eremurus subalbiflorus, and Bongardia takyrs — bare clay spots almost without vegetation. chrysogonum. Both slopes of the Vodorazdelny ridge Along the main part of the profile the climate is have microtherm shibliaks with Acer turcomanicum of a sharply continental dry type, only in the valley of as prevalent species and several subdominant species the Sumbar river it is subtropical. The mean annual such as Celtis caucasica, Crataegus sp. var., Cotone- temperature is 15.9°C in the north and 16.2°C in the aster nummularius, Lonicera floribunda, Colutea south. The prevailing monthly mean temperatures in buhsei, and Rhamnus coriacea. On the grass level June to August are 30°C in the Turanian lowland and Cousinia umbrosa, Allium paradoxum, and others 26 - 30°C in the Sumbar river valley. The ground were found. Woodless slopes are covered with traga- surface temperature can reach 50°C. Winter is often canthic formations of phrygana and units of tomil- mild and snowless in the lowland and at the foot of lares consisting, e.g., of Acantholimon sp. var., Thy­ the hills, the mountains are snow-covered. The iso­ mus afghanicus, and Onosma dichroantha. This therms of January are about 0°C in the north and whole area is well watered by numerous springs and +5°C in the valley of the river Sumbar, although the brooks near which single specimens of Juglans regia, temperature can fall to -29°C in the whole study area. Platanus orienatalis, and Elaeagnus orientalis are The volume of precipitation increases southward preserved. 128 Boris Tuniyev et al. Zoogeographic Analysis of the Herpetofauna of South-Western Turkmenistan 129 Landscape Landscape distribution of reptiles along the profile across western Turkmenistan. Fig. Fig. 2. 130 Boris Tuniyev et al.

Unit IV. The Sayvan-Nokhur plateau (1000 Unit IX. Malyi Balkhan. Parallel located chains - 1200 m). This is a district of cultivation and pas­ (400 - 700 m). The outer ridges are covered with ture. A big area is covered with crops. The primary semidesert vegetation whereas the inner ridges are vegetation is preserved island-like on the slopes of mainly badlands completely without vegetation. small hills and shows intermediate formations be­ Along the dried river bed Ferula oopoda, Leontice tween semideserts and mountain-steppes. The corre­ ewersmannii, Rheum turkestanicum, Amberboa am- sponding species are Artemisia oliveriana, Onosma berboi, Glaucium elegans, and Asparagus persicus dichroantha, Iris ewbankiana, Bothriochloa ischae- are found. mum, and Hypecoum parviflorum. There is one creek Unit X. Western end of the Central Karakum. on the plateau which dries out during the summer Sand desert with moving dunes, sandy hills and grass period. covered valleys. The grass vegetation is formed Unit V. The southern slope of the Peredovoy mainly of perennial, annual and monocarpical ephe­ ridge (1200 - 1500 m). The main vegetation type, Ju- meras (Poa bulbosa, Carex physodes, Eremopyrum nipereta turcomanica, consists of light juniper forests bonapartis, Arnebia decumbens, Silene папа, Strigo- and units of mountain-steppe. This district is very sella grandiflora, and Nonea caspica). Specialized steep, dry and without springs. trees and bushy psammophyts are represented by Ha- loxylon persicum, Ammodendron sp., Aellenia sub- Unit VI. Gorges of the northern macroslope of aphylla, Salsola richteri, Calligonum microcarpum, the Western Kopetdagh (600- 1000 m). Rocky C. caput-medusae, and C. leucocladum. gorges with steep slopes and big stony-broken areas. The main vegetation type is thermophyllous shibliak with Celtis caucasica, Acer turcomania, and Swida FAUNISTIC NOTES meyeri prevailing on the slopes and Cercis griffithii Class REPTILIA and Berberis densiflora in the steep places. Quite big Order Chelonia associations are formed by Hymenocrater bitumino- Family Emydidae sus and Ephedra sp. The corresponding grasses are Fritillaria raddeana and Hyacintus litwinowi. A few 1. Emys orbicularis (Linnaeus 1758) was only springs are present in this area. recorded along the river Sumbar, where in some places it was common but not numerous. Unit VII. Submountaneous inclined clay and 2. Mauremys caspica (Gmelin 1774) inhabits gravel plain of the Kopetdagh (up to 400 m). The like the former species only the valley of the Sumbar, covering of vegetation is very thin consisting of scat­ however in a much bigger population density than tered annual ephemeras, under-shrubs and low shrubs E. orbicularis. with the main component Artemisia. Its center is formed by specialized gypsophyts of different fami­ Family Testudinidae lies. The short vegetation period is determined by a 3. Agrionemys horsfieldi (Gray 1844). Speci­ spring rain and by summer dryness. mens were observed along the whole investigated Unit VII*. Clay plain near the southern foothills profile except of the bare takyrs of the Kyuren Dag of the Malyi Balkhan (Kyuren Dag Corridor). Clay Corridor. This species is abundant on the Monjukly lowland with single small ravines and sandy spots range, in the Sumbar river valley, on the southern with grass (up to 100 m). The desert vegetation is macroslope of the Western Kopetdagh and on its formed mainly of Artemisia badhysi and Salsola crest zone (units I - III). It is quite common on the richteri mixed with Arnebia linearifolia, Koelpinia Malyi Balkhan (unit IX) and comparatively rare in all macrantha, Zygophylum macrophyllum, Gagea gra- other units. minifolia, Hypecoum trilobum, etc. The vegetation Order Squamata on the clay consists of fragments of psammophylous Suborder Sauria associations. Family Agamidae Unit VIII. Takyrs (clay spots) of the Kyuren 4. Phrynocephalus helioscopus (Pallas 1771) Dag Corridor. Central part of the Kyuren Dag Corri­ was found in the clay plain of the Kyuren Dag Corri­ dor which has bare areas, so-called takyrs with primi­ dor (unit VIII), where it occurs on takyr and poorly- tive communities of Salsola gemmascens-Salsola grassed places of the clay desert. It penetrates into the rigida. first chain of hills of the Malyi Balkhan along the Zoogeographic Analysis of the Herpetofauna of South-Western Turkmenistan 131

broad dried out valleys but habitually it occurs on the fers the burrows of rodent colonies, mainly of Rhom- open plain clay unit. bomys opimus and Meriones erythrourus. 5. Phrynocephalus interscapularis Lichtenstein 13. Cyrtopodion russowi (Strauch 1887) lives 1856, was only observed along the profile in the on big bushes and desert-trees like Ammodendron, Karakum sand desert near the village Ahchakuyma Haloxylon, and Calligonum of the sand deserts, (Unit X). which grow rarely in too dense soils. We found 6. Phrynocephalus mystaceus (Pallas 1776) C. russowi near the village Ahchakuyma (unit X). It was found in the Karakum sand desert near the vil­ is also known to live in the valley of the river Sumbar lage Ahchakuyma. It is common on the slopes of big­ (Shammakov, 1981; Skalon, 1982; Atayev, 1987). ger sand dunes. 14. Cyrtopodion spinicauda (Strauch 1887) is a 7. Laudakia caucasia (Eichwald 1831) is found sporadically distributed species of the small-detritus along the whole Western Kopetdagh and Malyi gray shale badlands of the foothills of the Kopetdagh. Balkhan a widely distributed species. The population It is rarely found in the upper mountain belts of the on the Malyi Balkhan is much smaller than that on Kopetdagh. We observed it on the Malyi Balkhan and the Kopetdagh. Animals were observed in rocky up to 900 m on both macroslopes of the Western gorges as well as along the slopes and plateau-like Kopetdagh. grounds with big stones or blocks. 15. Eublepharis turcmenicus Darevsky 1977, is 8. Trapelus sanguinolentus aralensis Lichten­ known on the low belts of both macroslopes of the stein 1823. These agamas were observed along the Western Kopetdagh. Normally, it occurs in places whole profile with the exception of the takyrs in the with detritus and big blocks and shrubs not far away central part of the Kyuren Dag Corridor. They were from springs. most abundant in the clay and sand lowlands around 16. Teratoscincus scincus (Schlegel 1858) was the Malyi Balkhan. The population density decreased seen in the Karakum sand desert near the village Ah­ up to the mountains and increased again in the valley chakuyma, where it was quite common in open sand of the river Sumbar. patches. Family Anguidae Family Lacertidae 9. Pseudopus apodus (Pallas 1775) was only 17. Eremias grammica (Lichtenstein 1823) is found in the Western Kopetdagh where it was com­ the biggest-sized desert lacertide observed by us. paratively rare on the northern macroslope (units This species was common on the big sand dunes near VI - VII) and quite abundant in the upper belts of the the village Ahchakuyma. mountains and on the southern macroslope. As a 18. Eremias intermedia (Strauch 1876) was the xero-mesophylous species it prefers shibliaks and most prevalent reptile species on the clay plain near bushes or the vicinity of springs, but it also occurs in the southern foothills of the Malyi Balkhan. The the units of the mountain-steppe and semideserts. animals were also comparatively common on sandy patches with grass near the northern edge Of the Family Gekkonidae Malyi Balkhan (units VIII, X). 10. Alsophylax laevis Nikolsky 1907, lives in 19. Eremias lineolata (Nikolsky 1896) occurs the central part of the clay plain in the Kyuren Dag on open sand plains with few shrubs. We observed it Corridor. It prefers the termitariums among poor as well in the sand desert near Ahchakuyma as on the semishrubs оf Artemisia and Salsola. small sandy spots inside the clay plain near the south­ 11. Crossobamon eversmanni (Wiegmann ern foothills of the Malyi Balkhan (units VII*, X). 1834) is a sand-loving gecko of the plains, very com­ 20. Eremias scripta (Strauch 1867) is a typical mon in the sand desert near the village Ahchakuyma. sand-loving species. It inhabits the crests of high Above that it was found on the small sandy spots near aeolian sand dunes, mainly near the sand grass Stipa- the foothills of the Malyi Balkhan. grostis sp. We did not find this species but, according 12. Cyrtopodion caspius (Eichwald 1831) is a to Bogdanov (1962) and Shcherbak (1974), it is species widely distributed throughout the profile, known from Ahchakuyma. with the exception of the takyrs (unit VIII). In the 21. Eremias strauchi kopetdaghica Szczerbak mountains it lives in various rocky and stony habitats. 1971, is a common species in the Western Kopet­ In the hills and plains (including sand plains) it pre­ dagh. It is dominant in the upper mountain belts 132 Boris Tuniyev et al.

(units III — V) but relatively few specimens are found 1985) and from the inclined plain of the Western in the lower belts of the northern macroslope. Kopetdagh (Shammakov, 1981). 22. Eremias velox (Pallas 1771) spreads along Suborder Serpentes the whole profile from the sand deserts and takyrs Family Boidae round the Malyi Balkhan southward to the ridge Monjukly. The biggest population density is found in 29. Eryx elegans (Gray 1849) was found on the the foothills of the Kopetdagh and in the Malyi Bal­ profile on the Sayvan-Nokhur plateau (unit IV) and khan whereas it is smaller in the sand deserts and the on the crest of the Vodorazdelny ridge (unit III) with high mountain belts. corresponding mountain-steppe vegetation and inter­ 23. Mesalina guttulata watsonana Stoliczka mediate semidesert-steppe vegetation. Normally it 1872, was only found on the profile on the inclined prefers rodent colonies and places with big flat lowland of the northern foothills of the Kopetdagh. It stones. is quite common but not numerous in the gravel 30. Eryx miliaris (Pallas 1773) is a widely dis­ (gypsophylous) wormwood semidesert. tributed species with the highest population density in the sand deserts. On the profile, many specimens Family Scincidae were seen in the Karakum sand desert near Ahcha- 24. Ablepharus pannonicus (Lichtenstein 1823) kuyma, only occasionally in other places and none on is a common species in the Western Kopetdagh. the crest of the Kopetdagh (unit III). This snake can Skinks were only abundant among Phryganas from even live on takyrs where it finds refuge under the the foothills up to the crest of the Kopetdagh. Appar­ crest of the clay’s crust. ently, it is possible to find this species also in the up­ Family Colubridae per parts of the Malyi Balkhan. It is known from the more northerly located ridge of the Bol’shoi Balkhan. 31. Boiga trigonatum melanocephala Annan- Unfortunate weather conditions during our work on dale 1904. A species of the plains which is spread the main crest of the Malyi Balkhan were character­ over sand and clay deserts round the Malyi Balkhan ized by storm winds resulting in the absence of reptile and near the northern foothills of the Western surface activity (except for single specimens of Lau- Kopetdagh. dakia caucasia in the cracks of rocks). 32. Coluber atayevi Tuniyev and Shammakov, 25. Eumeces schneideri princeps Eichwald 1993, was earlier included in C. najadum. It was 1839, is a common species of the foothills and known as one of the rarest snakes of the Kopetdagh low-mountain belts on both slopes of the Western (Nikolsky, 1916; Chernov, 1934; Bogdanov, 1962; Kopetdagh. It prefers grassy soft grounds with big Atayev and Shammakov, 1990) and accordingly it flat stones. was included in the Red Data Book of Turkmenistan 26. Eumeces taeniolatus Blyth 1854, is a widely (1985). Actually this snake is locally common in the distributed skink along the whole Western Kopet­ upper belts of the mountains, up from 750 - 800 m dagh from the foothills to the highest limits. Most above sea level. It is a dominant species at 1000 specimens were observed in stony and rocky gorges - 1200 m which is, in comparison with other snakes, with steep slopes covered with bushes and shibliak frequently found. vegetation. 33. Coluber ravergieri Menetries 1832, was found along the whole Western Kopetdagh. This 27. Mabuya aurata (Linnaeus 1758) is as fre­ snake prefers humid shady places not far away from quent in the Western Kopetdagh as the former spe­ cies. It reaches down to the inclined plain near the springs and creeks. The largest density was observed Kopetdagh foothills but it does not form populations in the small gorges of the Vodorazdelny ridge of the Kopetdagh (unit III). of high density as all other skinks of the Kopetdagh. 34. Coluber karelini Brandt 1838, is a species of Family Varanidae the plains. It is more common in sand deserts but can 28. Varanus griseus caspius Eichwald 1831, also live on the clay units. We observed it around the populates the plain and reaches the foothills. We Malyi Balkhan and near the northern foothills of the found it in the clay lowland of the Kyuren Dag Corri­ Kopetdagh. dor. Traces of it were seen in the sand near Ahcha- 35. Coluber rhodorhachis (Jan 1865) (= C. la- kuyma. In the literature these monitor lizards were re­ dacensis [Anderson 1871]). This species is not com­ corded from the valley of the river Sumbar (Atayev, mon in the mountains of the Western Kopetdagh and Zoogeographic Analysis of the Herpetofauna of South-Western Turkmenistan 133 the Malyi Balkhan. It inhabits stony and rocky bio­ both in sand and in clay deserts (except for takyrs). It topes of gorges and occurs also on slopes near the does not occur on real mountains but is found in the burrows of rodent colonies. valley of the river Sumbar (Atayev, 1985). 36. Coluber schmidtii Nikolsky 1909, is com­ Family Elapidae mon in the Sumbar river valley. It is found southward 45. Naja oxiana (Eichwald 1831) is a widely from the village Bami on the northern foothills of the distributed species along the whole profile, except for Western Kopetdagh (Atayev et al., 1991). We found takyrs (unit VIII). The number of findings increases it on the Sayvan-Nokhur plateau, where it prefers ro­ in the Sumbar river valley and on the slopes of the dent colonies in Artemisia bushes along the plain range Monjukly and of the Kopetdagh which are ex­ places near watercourses. posed towards that valley. 37. Eirenis medus (Chernov 1949) was only caught on the Sayvan-Nokhur plateau (unit IV), Family Typhlopidae where the species was locally common under the 46. Typhlops vermicularis Merrem 1820, is stones in semidesert-steppe areas on the slopes of common and locally numerous on the submountain small hills. inclined plain on the north of the Kopetdagh south­ 38. Lycodon striatus bicolor Nikolsky 1903 is a ward to the ridge Monjukly (in all mountain belts of thermophyllous species which inhabits semidesert the Western Kopetdagh). and shibliak biotopes of the Western Kopetdagh. The Family Viperidae few findings are due to the nocturnal life-style of this species. 47. Echis multisquamatus Cherlin 1981 lives in 39. Lythorhynchus ridgewayi Boulenger 1887 the plains. It was recorded round the Malyi Balkhan was only seen on the clay plain between the Western and near the foothills of the Kopetdagh. It is also Kopetdagh and the Malyi Balkhan, except the central found in the Sumbar river valley (Skalon, 1982; Ata­ takyrs. It is a comparatively rare, but in suitable habi­ yev, 1985). tats and at optimal activity seasons locally common 48. Vipera lebetina (Linnaeus 1758) is common snake. It was also observed near the southern foot­ on both macroslopes of the Western Kopetdagh, hills of the Malyi Balkhan in 1992. where it mainly occurs on stony and rocky gorges with big blocks and steep slopes among the shibliaks. 40. Natrix tessellata (Laurenti 1768) is sporadi­ Since if is not caught for serpentariums in the gorges cally distributed along the whole Western Kopet­ of the northern macroslope it occurs there in big num­ dagh, except for light Juniper forests. It lives in habi­ bers. The taxonomic status of this population is not tats near constant watercourses (springs, creeks, clearly defined but we believe it to be closer to V. le­ rivers), but it might be found in hibernation places quite far away from the water. betina obtusa (although somewhat smaller) than to the form “cernovi” described from the eastern 41. Oligodon taeniolatus (Jordan 1853) was not Kopetdagh (Chikin and Shcherbak, 1992). encountered by us during our expedition but it is known from the studied area, according to other au­ Family Crotalidae thors (Scherback, 1979; Atayev et al., 1991; Atayev, 49. Agkistrodon intermedius caucasicus Nikol­ 1985, 1987). It exists in the Western Kopetdagh syn- sky, 1916 inhabits the Western Kopetdagh only on topically with Lycodon striatus. the highest part of the ridge up from 1200 m. Snakes 42. Psammophis lineolatum (Brandt 1838) is a were found on the steep slopes of gorges covered widely distributed species on the plains. Beside the with microtherm shibliaks and with big colonies of Kyuren Dag and the Balkhan Corridors it lives on the Microtus afghanus. Malyi Balkhan, in the valley of the river Sumbar and on the Sayvan-Nokhur plateau. DISCUSSION 43. Pseudocyclophis persicus (Anderson 1872) is a common but not numerous species of low belts We found 49 reptile species which account for on both slopes of the Western Kopetdagh. In the 62% of the known list of reptiles of Turkmenistan in north it can also live on the inclined lowland not far spite of the comparatively small area. This is due to from hills. the various landscapesour of profile: from the far 44. Spalerosophis diadema schiraziana Jan west end of the Central Karakum desert, across some 1865, is a species of the plains which was observed parts of the Turkmeno-Khorasan mountains, to the 134 Boris Tuniyev et al. subtropical districts on the border of Iran. The chosen 4. Eurytopic mountain species. units correspond to different landscapes of the pro­ These are widely distributed reptiles in all or al­ file. They are non-equivalent in species numbers. The most all mountain belts (Laudakia caucasia, Pseudo­ richest fauna was found in the valley of the river pus apodus, Eremias strauchi, Ablepharus pannoni- Sumbar (30 species) and the poorest reptile list was cus, Mabuja aurata, Eumeces taeniolatus, Coluber found in the takyrs of the Kyuren Dag Corridor (5 ravergieri, C. rhodorhachis, Natrix tessellata, Typh- species). If only the number of original (correspond­ lops vermicularis, and Vipera lebetina). ing) species is considered, both the Sumbar valley and the takyrs of the Kyuren Dag Corridor have but 2 5. Ubiquist species. original species correspondingly. Therefore, Emys Representatives of this group might be typical in orbicularis and Mauremys caspica are only present some landscapes but live as well in lowlands as in on the profile in the Sumbar valley because they need mountains at all or almost all ecological alti- comparatively large constant water courses. The ta­ tude-belts. (Agrionemys horsfieldi, Trapelus sangui- kyrs on the other hand are the unique biotop of Phry- nolentus, Cyrtopodion caspius, Eremias velox, Eryx nocephalus helioscopus and Alsophilax laevis. miliaris, and Naja oxiana). The species composition for each landscape unit It can be seen in Fig. 2 that several of the plain depends on the differences in the ecological tolerance species are also present in the Sayvan-Nokhur pla­ of the reptiles. We have distinguished several ecolog­ teau higher than 1000 m. Firstly these are Psammo­ ical groups of reptiles in western Turkmenistan ac­ phis lineolatum and Coluber schmidtii. This is ex­ cording to their character of distribution and altitude plained by the distribution of the two species through range. wide weakly inclined valleys from the south-west up 1. Stenotopic plain species. to the Sayvan-Nokhur plateau. Many springs and creeks in combination with semidesert plain units This group consists of 3 subgroups, la — provide favorable conditions for mesophylous spe­ stenotopic species of sand deserts (Phrynocephalus cies like Coluber schmidtii and desert inhabitants like interscapularis, Ph. mystaceus, Crossobamon evers- Psammophis lineolatum and Eryx miliaris. In the manni, Cyrtopodion russowi, Teratoscincus scincus, same way, several desert animals like Cyrtopodion Eremias grammica, E. lineolata, and E. scripta): lb russowi, Varanus griseus, Psammophis lineolatum, — stenotopic species of clay deserts (Phrynocepha­ Spalerosophis diadema, and Echis multisquamatus lus helioscopus, Alsophylax laevis, Mesalina guttula- have reached the Sumbar river valley from the near ta, and Lythorhynchus ridgewayi)', lc — stenotopic Caspian deserts. All these species are confined to the hydrophylous and mesophylous species (Emys orbi­ small units of corresponding habitats (i.e. sand or cularis, Mauremys caspica, and Coluber schmidtii). badlands) which was well illustrated by Bogdanov 2. Eurytopic plain species. (1962) and Atayev (1987). All these species are not encountered in the tugay-forests and in the humid These reptiles occur both in sand and clay deserts biotopes of the Sumbar valley. and are as a rule only absent in large takyrs (Eremias intermedia, Varanus griseus, Boiga trigonatum, Co­ Another interesting aspect arises from compari­ luber karelini, Psammophis lineolatum, Spaleroso- son of the number of species for mountains and low­ phis diadema, and Echis multisquamatus). lands in general and for each subdivision particularly. We noted 22 species for plains, 21 for mountains, and 3. Steno- and oligotopic mountain species. 6 ubiquists. The number of species of the plains is This group includes animals, which occur in cer­ slightly more than that for mountain species even if tain mountain belts. We distinguish two subgroups. we include several local findings of other reptiles 3 a — species of foothills and lower belts of moun­ near our profile like Lacerta strigata from the rivers tains (Cyrtopodion spinicauda, Eublepharis turcme- Chandyr and Atrek on the south (Atayev, 1985) or nicus, Eumeces schneideri, Lycodon striatus, Oligo- Phrynocephalus maculatus near the village Bami, don taeniolatus, and Pseudocyclophis persicus); Ph. raddei from the northern foothills of the 3b — species of middle- and high-mountain belts Kopetdagh (Shammakov, 1985), and Telescopus rhy- (Eryx elegans, Coluber atayevi, Agkistrodon inter- nopoma from the station Iskander (Rustamov and medius, and Eirenis medus). Atayev, 1976). Zoogeographic Analysis of the Herpetofauna of South-Western Turkmenistan 135

A comparison of the herpetofauna from the Besides the daily niche segregation between diur­ Western Kopetdagh and the Malyi Balkhan shows a nal and nocturnal species, there exists a strong poor list of mountain elements from the latter. On the microhabital preference (Fig. 3) which is evolution­ Malyi Balkhan we observed Cyrtopodion spinicauda, ary fixed. It was well described by Ananjeva (1976) Laudakia caucasia, and Coluber rhodorhachis. Ap­ by the example of the genus Eremias. parently these species entered the Malyi Balkhan in Now we come to a more detailed discussion of the past more humid periods, when gorge-like water the microhabital occurrence of sand desert reptiles. courses ran into the Kyuren Dag Corridor from both High moving sand dunes along crests with sand-grass sides (i.e., from the Kopetdagh and the Malyi (Stipagrostis sp.) are the habitat of Eremias scripta. Balkhan) and formed suitable conditions for the en­ Open grassless slopes of high dunes are populated by trance of mountain reptiles. It is difficult to estimate ■ Phrynocephalus mystaceus. On the same dunes in the now, if these were the only species who could reach units with a few shrubs (Calligonum sp.) Eremias the Malyi Balkhan or if the rigorous modem climatic grammica and Teratoscincus scincus can be ob­ conditions, mainly cruel winds (for about 200 days served. Along the open grassless valleys between per year), have preserved only a small part of the for­ high dunes two species are common: Phrynocephalus mer mountain inhabitants while the rest became ex­ interscapularis and Eremias lineolata. In more tinct. This might be true for our finding of Mabuja grassy interdune valleys and on sandy hills live Cros- aurata and Ablepharusjpannonicus which are known sobamon eversmanni and Eremias intermedia. The in more northerly areas like the Bol’shoi Balkhan, the main spectrum of moving and semifixed sands form Uzboy valley and the Caspian shore. the habitats of Eryx miliaris, Boiga trigonatum, and At the Kopetdagh the most various fauna was ob­ Echis multisquamatus. Species like Agrionemys served at the lower belts of both macroslopes (23 spe­ horsfieldi, Eremias velox, and Trapelus sanguinolen- cies). The species composition was more or less iden­ tus appear on sands with the most vegetation cover­ tical, thus we could not pick out corresponding ele­ ing. Cyrtopodion caspius, Psammophis lineolatum, ments for the described landscape units (I, I , VI). Coluber karelini, and Spalerosophis diadema live But generally, the number of original reptiles is still near the burrows of rodent colonies. All types of large at the foothills of both slopes of the Kopetdagh microstations in the sand desert are under the control (Eumeces schneideri, Eublepharis turcmenicus, Ly­ of Var anus griseus and Naja oxiana. On the desert codon striatus, Oligodon taeniolatus, and Pseudocy- trees and big shrubs like Haloxylon, Ammodendron, clophis persicus). and Calligonum lives the gecko Cyrtopodion russowi. The herpetofauna of the upper belts in the Kopet­ The clay desert between the Malyi Balkhan and dagh is poor but has its own corresponding species. the Kopetdagh has been separated into two units by Thus 17 species were found in the mountain-steppe the takyrs. Near the Kopetdagh on the inclined plain a belt (including the microtherm shibliak) and two of row of mountain species get more abundant towards them live only there (Eryx elegans and Agkistrodon the foothills. Moreover, Mesalina guttulata is a cor­ intermedius). The reptile list of the Sayvan-Nokhur responding species of this part of the gypseous plateau is a little bigger (19 species) due to the above gravely clay desert, which is not found near the Malyi mentioned causes of invasion of some species of Balkhan. At the same time the small sand spots in the plain landscapes. The original reptile of this plateau clay desert near the Malyi Balkhan are populated by is Eirenis medus. The smallest number of high moun­ some sand preferring species (Eremias intermedia, tain reptile species occurs in the light juniper forests E. lineolata, and Crossobamon eversmanni) which with very dry microclimatic characteristics. are not encountered near the Kopetdagh. Comparison of the fauna of different desert types One of the most debatable aspects is the zoogeo- shows that the sand desert is richer and more distinc­ graphical status of the investigated area. Perhaps the tive than the clay desert. This is at first sight a para­ main cause of the traditional uniting of the heteroge­ dox, which can be explained by a broader variation of neous fauna of the plains and mountains of Middle the microbiotopes (microstations) in the sand than in Asia has been the study of this region according to the clay plain. Botanists connect the increasing the political boundaries, ignoring the natural geo­ speciation process of plants in the Karakum with the graphical units. But the majority of authors have rec­ mobility of the sand (Kolenov and Muhamedov, ognized the originality of the sand desert and of the 1992). This rule is the same for all biota. mountain fauna. 136 Boris Tuniyev et al. Microhabital distribution of reptiles in sand desert. Fig. 3. 3. Fig. Zoogeographic Analysis of the Herpetofauna of South-Western Turkmenistan 137

One of the first conceptions was advanced by Ni­ “Turanian”): Phrynocephalus helioscopus, Ph inter- kolsky (1916). According to him, the Middle Asian scapularis, Ph. mystaceus', Crossobamon eversman­ fauna is young and was formed during the postglacial ni, Cyrtodactylus (Cyrtopodion) russowi, Eremias period by migrants, mainly from Central Asian grammica, E. intermedia, E. lineolata, E. scripta, and deserts. This opinion was refuted by Chernov ( 1949) E. velox. Species like Clemmys (Mauremys) caspica, who pointed out the existence of special speciation Emys orbicularis, and Ophisaurus (Pseudopus) apo- centers of sand desert, clay, gravel and foothill desert dus are called Mediterranean. Agama agilis (= Tra- fauna in Middle Asia. According to Chernov, the last pelus sanguinolentus), Eumeces schneideri, Mabuya three faunas had connections with the south-west aurata, and Varanus griseus were called Iranian/Sa- Asian (Middle East) fauna, but they are not identical. haro-Sindian faunal elements. And finally, the so Among the mountain speciation centers in Middle called “Iranian” (Anderson 1968) faunal elements Asia Chernov noted only the Pamirs-Alay and Tien consist of Testudo (Agrionemys) horsfieldi, Agama Shan mountain systems. It is necessary to point out caucasica (Laudakia caucasia), Alsophylax (Cyrto­ that Chernov ( 1935) reported earlier also special liz­ podion) spinicauda, Cyrtodactilus (Cyrtopodion) ard forms corresponding to the Turanian plain, such caspius, Teratoscincus scincus, Eremias (Mesalina) as Teratoscincus scincus, Gymnodactylus russowi guttulata watsonana, Eremias velox strauchi (= Cyrtopodion russowi), Agama aralensis (= Trape- (= E. strauchi), Ablepharus pannonicus, and Eume­ lus sanguinolentus), Eremias scripta, E. lineolata, ces taeniolatus. E. grammica, Phrynocephalus mystaceus, and Ph. in- The examination of the zoogeographical connec­ terscapularis. Later in the zoogeographical chapter tions between reptiles from Middle Asia and the Cau­ Terentyev and Chernov (1949) separated the plains casus allowed Rustamov (1981) to relate Gymnodac­ of Middle Asia from the Kopetdagh and the Pamirs- tylus (Cyrtopodion) caspius, G. russowi, Phrynoce­ Tien Shan into subdivisions. But this symbolic sepa­ phalus helioscopus, Ph. mystaceus, Ablepharus pan­ ration has not been analyzed and explaned and has no nonicus, Agama (Trapelus) sanguinolenta, Eremias ranks to the biochores. This subdivision had a rather velox, Elaphe dione, Psammophis lineolatum, Agkis- physiographic character. At the same time, the au­ trodon halys, and Eryx miliaris to the Turanian spe­ thors discussed the origin of several Middle Asian cies. According to Rustamov, these species have their reptiles. Besides the lizards mentioned above, Terent­ origin in the center of reptile speciation in the Turani­ yev and Chernov ( 1949) classified Eremias interme­ an desert. Other species were segregated by Rusta­ dia, Crossobamon eversmanni, and Eryx miliaris as mov into the following faunal groups: Mediterra­ Middle Asian authochtons; Eremias strauchi, Vara- nean: (Mauremys caspica, Pseudopus apodus, Natrix nus griseus, Coluber najadum (= C. atayevi), and Ly- tessellata), European: (Emys orbicularis, Coluber thorhynchus ridgewayi as Middle East (South-west schmidtii), Irano-Afghanian: (Laudakia caucasia, Asian) species; Eublepharis macularius (= E. turc- Mabuya aurata, Eumeces schneideri, Eremias menicus), Oligodon taeniolatus, Lycodon striatus, strauchi, Typhlops vermicularis, Coluber najadum and Naja naja .(= Naja oxiana) were included in the (= C. atayevi), C. ravergieri); Saharo-Sindian: ( Vipe- Indian faunistic elements. Testudo horsfieldi (Agrio- ra lebetina)-, Minor Asian or Caucasian-Minor Asian: nemys horsfieldi) was called a product of tropical (Lacerta strigata). Africa. Shcherbak (1982) has separated a special arid Anderson (1968) reports in his zoogeographical Mediterranean-Asian subregion (“Aridisches Medi- analysis of the lizard fauna of Iran about the original­ terranian-Asiatisches Untergebiet”) in the Palearctic ity of the Iranian plateau and suggests that even in the subdivision. Later he gave additional evidence for it Iranian part of the Turkmen steppe (low plain near (Shcherbak, 1984). According to his map (p. 229), the Kopetdagh) “only 43.5% of the lizard fauna are the Turkmenian part of the Kopetdagh is together forms which can be considered truly Aralo-Caspian, with the sand deserts in the Karakum sand district of the remainder being species from the Iranian plateau “Karakum-Bezirk” of the Turanian desert province. and species confined primarily to mountain slopes...” Situated northward from the Kopetdagh, the Kara­ (p. 319). And right in the Iranian plateau the number kum takyr district belongs to the same province of Aralo-Caspian species forms only 8.3%. Accord­ (Shcherbak, 1982). ing to Anderson the following species belong to the Finally, one of the recent zoogeographical ana­ Aralo-Caspian faunal elements (which be called lysis of the mountain herpetofauna of Turkmenistan 138 Boris Tuniyev et al. was conducted by Atayev (1985). The reptiles'of the the elevated regions of South-West Asia and their re­ Western Kopetdagh belonged to the following faunal cent distribution largely spreads from Eastern elements: Frontal Asian (Middle East): (Mauremys Anatolia and the Armenian Highland towards the Ira­ caspica, Eumeces taeniolatus, Ablepharus pannoni- nian plateau with irradiation into arid and semiarid cus, Eremias strauchi, Typhlops vermicularis, Eryx districts of the Near East and Eastern Transcaucasia elegans, Coluber rhodorhachis, C. ravergieri, Eire- northward up to Daghestan. The species of the nis meda (E. medus), E. (Pseudocyclophis) persicus, Turkmeno-Khorasan subgroup are endemics of these Oligodon taeniolatus, Telescopus rhynopoma); Eu- ridges and they can penetrate along the continuums ropean-Mediterranean: (Emys orbicularis, Ophisau- of this mountain system westward to Alburz and east­ rus (Pseudopus) apodus, Natrix tessellata, Coluber ward to the Hindu Kush. The Irano-Afghanian sub­ jugularis (= C. schmidtii))', Endemic for Middle Asia group consists mainly of species from the intermedi­ and Eastern Iran: (Agrionemys horsfieldi, Eublepha- ate foothill and low mountain zone of the eastern part ris turcmenicus, Gymnodactylus (Cyrtopodion) cas- of the Iranian plateau and the mountain system of the pius, G. (C.) spinicauda, Naja oxiana)', Sub-endemic Hindu Kush with insignificant irradiation to the inner for Middle Asia and Eastern Iran: (Agama (Trapelus) districts of Middle Asia. sanguinolenta, Eremias velox, Psammophis lineola­ Mediterranean species (including European-Me- tum)', Saharo-Sindian: (Mabuya aurata, Eumeces diterranean) have their origins from areas which are schneideri, Spalerosophis diadema, Vipera lebetina); located around the modem Mediterranean sea. They Indian: (Lycodon striatus) and Caucaso-Minor have often a disjunct distribution in that area and Asian: (Agama (Laudakia) caucasia, Lacerta more eastwards along the regions connected with the strigata). ancient Tethys. Thus, we can see the absence of a common ap­ The Oriental group includes Paleotropical spe­ proach to the zoogeographical subdivision of the cies of Southern and South-Eastern Asia with recent Turkmenistan area and it is even impossible to unite distribution from Sri Lanka and the Hindustan penin­ opinions about which faunal elements the various sula up to eastern Hindu Kush. It has an insignificant species belong to. In addition, these points of view irradiation northward to the foothills of Gissaro-Dar- are often diametrically opposite to each other. vaz and the Kopetdagh mountains. On the basis of recent chorology and ecological The Saharo-Sindian' group includes reptiles tolerance (generally on habitat preferences), we which are separated from the ancient Mediterranean picked out 5 main ecological-geographical groups of group due to the conditions of aridization on the reptiles from the investigated area. These are the southern border of the Palearctic. These are mainly Turanian, South-West Asian, Mediterranean, Orien­ inhabitants of arid and extra-arid plains of northern tal, and Saharo-Sindian groups. Africa eastwards to the deserts of India, part of which, The Turanian group includes species with origin has penetrated to the deserts of the Turanian plain. and modern distribution in the sand and clay deserts The zoogeographical analysis of the herpeto- of the Turanian plain with insignificant irradiation to­ fauna of the Western Kopetdagh, the Malyi Balkhan wards the neighboring areas of Eastern Iran, South­ and the Turanian plain (Fig. 4) shows the progressive ern Kazakhstan and the Fergana valley. All these spe­ decreasing number of South-West Asian species and cies (see Table 1) are animals of the plains and their the increasing number of Turanian representatives findings in elevated districts are determined by the from elevated areas towards the plain. presence of suitable habitats (big plain surface with The Western Kopetdagh has only 19.3% Turani­ sand or clay deserts). In our opinion, a more detailed an elements whereas the number of South-West subdivision of the Turanian group (based on edaphic Asian species is 54.8%. The Mediterranean fauna factors) is not necessary at least not in this article be­ which is not found on the Malyi Balkhan and the Tu­ cause the whole Turanian plain consists more or less ranian plain is present only on the Kopetdagh of a mosaic of sand and clay units. (19.4%). The South-West Asian and the Mediterra­ The South-West Asian group is of a complex nean species form more than 74% of the herpeto- composition. It was divided into 3 subgroups accord­ fauna of the Western Kopetdagh. Among the South- ing to the species distribution patterns and altitude West Asian group of reptiles in the Western Kopet­ range: Iranian, Turkmeno-Khorasanian, and Irano- dagh are 36.7% of Iranian elements, 16.7% endemics Afghanian. The Iranian elements have their origin in of the Turkmeno-Khorasan mountains, and only Zoogeographic Analysis of the Herpetofauna of South-Western Turkmenistan 139

TABLE 1. The Distribution of Reptiles of Western Turkmenistan within the Main Physiography Subdivisions

Species Faunal group Western Kopetdagh Malyi Balkhan Turanian plain

Emys orbicularis E/M + - -

Mauremys caspica M + -_ Agrionemys horsfieldi T + + + Phrynocephalus helioscopus T - + +

Ph. interscapularis T - - + Ph. mystaceus T - - + Laudakia caucasia I + + _ Trapelus sanguinolentus T + + +

Pseudopus apodus M + -- Alsophylax laevis T -- + Crossobamon eversmanni T - + + Cyrtopodion caspius T + + + C. russowi T -- +

C. spinicauda KH + + _

Eublepharis turcmenicus KH + + — Teratoscincus scincus T - _ + Eremias grammica T - _ + E. intermedia T - . + + E. lineolata T - . _ + E. scripta T - _ + E. strauchi I + __ E. velox T + + + Mesalina guttulata SS -_ + Ablepharus pannonicus I + ? _ Eumeces schneideri I + __ E. taeniolatus I + __ Mabuya aurata I + __ Varanus griseus T + + + Eryx elegans KH + - _ E. miliaris T + + + Boiga trigonatum 0 - + + Coluber atayevi KH + __ C. karelini T _ + + C. rhodorhachis I + + _ C. ravergieri I + - _ C. schmidtii I + __ Eirenis medus KH + _ _ Lythorhynchus ridgewayi AF - + + Lycodon striatus 0 + _ _ Natrix tessellata M + _ _ Oligodon taeniolatus 0 + __ Psammophis lineolatum T + + + Pseudocyclophis persicus I + - _ Spalerosophis diadema SS - + + Agkistrodon intermedius KH + - _ Naja oxiana AF + + + Typhlops vermicularis M + _ _ Echis multisquamatus T - - + Vipera lebetina M + - - Note. +) known presence; M) Mediterranean faunal element; I) Iranian faunal element; E) European faunal element; KH) Turkmeno-Khora- sanian faunal element; AF) Irano-Afghanian faunal element; SS) Saharo-Sindian faunal element; O) Oriental faunal element; T) Turanian faunal element. 140 Boris Tuniyev et al.

Western Malyi Turanian Turanian plain. The Peredovoy ridge as well as the Kopetdagh Balkhan plain foothills are often an insurmountable barrier for pen­ etration of the Turanian plain species towards the Ko­ petdagh. On the other hand, the plain conditions of sand and clay deserts have not been suitable habitats for mountain South-West Asian and Mediterranean reptiles. Regarding this, it is necessary to include the Ecological-geographical groups: Western Kopetdagh into the South-West Asian prov­ ince and the Turanian plain into the Turanian prov­ | Turanian | I South-West Asian ince. The border between these two provinces lies be­ | Mediterranian Oriental tween the northern foothills of the Kopetdagh and its | Saharo-Sindian inclined plain. Apparently, the Malyi Balkhan should Fig. 4. Pie diagrams representing the chorotype composition of be united with the Turanian province as a separate herpetofauna of western Turkmenistan. district. Finally, it is necessary to mention that zoogeo- graphists which based their examples on other classes 3.3% of Irano-Afghanian species. Quite a high level of vertebrate animals in their zoogeographical subdi­ of local endemism, described earlier by Darevsky visions of Turkmenistan came to the same conclu­ (1981), and the presence of Mediterranean species sions (Zykov, 1991; Reshetnikov and Shakirova, place the Western Kopetdagh closer to the Alburz 1993). and Talysh than to the Eastern Kopetdagh which has As mentioned above, the zoogeographical bor­ an interconnection with the Hindu Kush through ders prove to be correct according to us, due to the Badhyz. The only endemic species in the Eastern species composition of the inclined lowland. Turani­ Kopetdagh are C. spinicauda and E. turcmenicus. At an (37.5%) and Saharo-Sindian (8.3%) elements the same time there are Irano-Afghanian species like which form the backbone of the plain desert fauna, Cyrtopodion longipes and Laudakia erythrogastra totally represent about half of the reptiles of this spot. and Saharo-Sindian elements like Psammophis The other half consists of South-West Asian (25%), schokari. Mediterranean (16.0%) and Oriental (12.5%) faunal More than half of the species list of the fauna of elements. the Malyi Balkhan consists of .the Turanian group (61.0%). The number of South-West Asian species is Acknowledgments. We thank Dr. Sakhat Shamma­ kov, Dr. Chary Atayev, and Dr. Lidia Marinina, Zoological still high (but less than 28%), whereas the endemics Institute, Ashgabat, Turkmenistan, for their assistance in of the Turkmeno-Khorasan mountains are only repre­ organizing and conducting the field work in Turkmenistan. sented by C. spinicauda. The numbers of Iranian and Heinz Bolzem assisted in the field and critically discussed Irano-Afghanian species reach 11.8% each. There is the fito-landscapes along the profile. Irina Marchukaytis il­ 1 representative of the Saharo-Sindian and 1 of the lustrated this publication. Oriental group (each 5.6%). The majority of the Turanian plain species are REFERENCES from the Turanian ecological-geographical group (80.0%). The South-West Asian group represents a Ananjeva N. B. (1976), “The biotopic distribution of five minority here (8%), moreover it consists totally of species of desert lizards (Sauria, Eremias) in the south­ Irano-Afghanian species. The Saharo-Sindian (8%) ern Balkhash region,” Byull. Mosk. Obshch. Estestvo- and Oriental (8.3%) faunal elements are slightly isp., 81(1), 65-71 [English abstract, in Russian], increasing. Anderson S. C. (1968), “Zoogeographic analysis of the lizard fauna o f Iran,” in: W. B. Fisher (ed.), The Cam­ Thus we can suggest significant differences be­ bridge History o f Iran. I. The Land o f Iran, Cambridge tween the herpetofauna of the Western Kopetdagh Univ. Press, pp. 305 - 371. and the Turanian plain. These differences are associ­ Atayev Ch. (1985), Reptiles of the Mountains of Turk­ ated with the unequal age of genesis of these areas menistan, Ylym, Ashgabat [in Russian], and with the generally modem landscape-ecological Atayev Ch. (1987), “Comparative aspects of the herpeto­ isolation of the Kopetdagh from the desert of the fauna of the river valleys of the Sumbar and the Chan- Zoogeographic Analysis of the Herpetofauna of South-Western Turkmenistan 141

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