Chelonian Conservation and Biology, 2008, 7(1): 70–77 Ó 2008 Chelonian Research Foundation The Geographic Distribution of Turtles in and the Nagorno-Karabakh Republic (Artsakh)

MARINE ARAKELYAN1 AND JAMES F. PARHAM2,3

1Department of Biology, Yerevan State University, Alek Manukyan 1, Yerevan 375025, Armenia [[email protected]]; 2Department of Herpetology, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, USA; 3University Museum of Paleontology, 1101 VLSB, University of California, Berkeley, CA 94720, USA [[email protected]]

ABSTRACT. – The turtles of the Caucasus are poorly studied, and basic data on their geographic distribution in some Caucasian regions are poorly collated, hard to access, or entirely lacking. We present a comprehensive review of turtle localities in such a region, Armenia and Nagorno- Karabakh Republic (Artsakh), based on literature records, personal observations, and museum specimens. The compiled data for the 3 native species (Emys orbicularis, Mauremys caspica, and Testudo graeca) show that their distributions in this region are restricted to 3 areas of low elevation. In the northeast, turtles can be found in the Kura River drainage. In the south and west, turtles can be found in the Arax River drainage. The distributions of species within these areas vary, with E. orbicularis being the most restricted by elevation. We note that the 3 regions here form part of an important transect from the interior Arax drainage (in the Ararat region of western Armenia) to near its confluence with the Kura drainage (in southern Armenia and Nagorno-Karabakh) to the Kura drainage sensu stricto (in northeast Armenia). Future studies of turtles in these areas will play an important role in understanding the historical biogeography of Caucasian turtles.

KEY WORDS. – Reptilia; Testudines; Testudinidae; Emydidae; Geoemydidae; turtle; tortoise; Armenia; Caucasus; Arax River; Kura River; Emys orbicularis; Mauremys caspica; Testudo graeca

Armenia (Fig. 1) is situated in the southern Caucasus point for future studies on the distribution and conservation and covers approximately 10% of the Armenian plateau status of turtles in this region. (29,800 km2). It is bordered by Azerbaijan, Georgia, Iran, and Turkey. Nagorno-Karabakh Republic (also known as MATERIALS AND METHODS Artsakh) is a smaller (4400 km2) autonomous region between Armenia and eastern Azerbaijan. As a result of The data for this article are from 3 kinds of sources: 1) their geographical position and diversity of landscapes, a review of the pertinent literature; 2) personal observation both Armenia and Nagorno-Karabakh support a wide of the authors, as well as other Armenian biologists; 3) range of animal species, including 3 species of turtles. The museum specimens from the Zoological Institute of St. turtles of Armenia are from each of the 3 families of Petersburg, Russia (ZISP), and the Museum of the Testudinoidea: 1) an emydid, Emys orbicularis (Linnaeus, Zoological Institute of Armenia, Yerevan, Armenia 1758); 2) a geoemydid, Mauremys caspica (Gmelin 1774); (MZIA). Personal observations of turtles in the wild were and 3) a testudinid, Testudo graeca Linnaeus, 1758. primarily by MA unless stated otherwise, especially in Testudo graeca is listed in the Red Data Book of 2004 and incidentally from 1994 to 2006. Global Armenia (Movsesyan 1987), which provides it with positioning system (GPS) coordinates were taken at every nominal protection as a threatened species. However, the locality and all turtles were vouchered via photography. baseline data on its geographic distribution, crucial These images were then made into vouchers (both hard components for effective conservation management, are and digital) and deposited into the Museum of Vertebrate lacking. Some locality data are found in the works of Zoology, University of California, Berkeley, California, Chernov (1939) and Dal (1954). But since then, Darev- USA (MVZ). The GPS coordinates are also stored at the sky’s unpublished PhD thesis (1957) is the main report on MVZ and are available to qualified researchers. Additional distribution of turtles in Armenia and surrounding regions. information was collected from interviews with Armenian Because these publications are written in Russian, they are biologists (especially herpetologists) and local people. In overlooked by most readers. The objective of this research every case, we indicated the source of these data, is to gather baseline data on the geographical distribution recognizing that each constitutes a different class of of the turtle fauna in these regions by using personal information that can be confirmed or refuted with later observations, literature records, and museum specimens. It study. In instances where multiple sources report the same is our hope that this report will provide a good starting locality, we listed them in an approximate order of ARAKELYAN AND PARHAM — Geographic Distribution of Turtles in Armenia and the Nagorno-Karabakh Republic 71

cited by Darevsky 1957). Agasyan (pers. comm. to MA) also claimed that populations of E. orbicularis inhabit the Arax River in the Kovsakan region of Kashatagh Province in Nagorno-Karabakh, near the border with Iran. Subse- quently, one of us (MA) traveled there and collected one specimen in June 2004. After interviewing several local residents, it became apparent that E. orbicularis is abundant in this region and probably inhabits a wider distribution here than previously supposed. In June 2006, one of us (MA) discovered 2 new localities for this species in Nagorno-Karabakh (localities 6 and 7 below). In the studied area, E. orbicularis occurs at altitudes of 290 to 960 m and inhabits ponds, bogs, lakes, rivers, and irrigation channels.

1. Agshtev River valley, Tavush Province (960 m). Dal (1954). Pers. obs. F. Danielyan (2000), A. Agasyan (1998), S. Pipoyan (2001), and E. Yavruyan (2000). 2. Dzhogas River, at the confluence of the Agshtev River, Tavush Province. Darevsky (1957). 3. Vicinity of Yerevan city, Hrazdan River, Yerevan Province. Museum specimen (MZIA 6/1912). Lyajster (1912), Chernov (1939). 4. Arax River Valley, Syunik Province. Bobrinsky (1915), Skryabin (1924), Darevsky (1957). 5. Two km west of Hadjari (Krmen) (village), the bank of the Hakari (Akera) River, at the junction with the Arax River, Kashatagh Province, Nagorno-Karabakh (290 m). Digital voucher (2004; MVZ 250832). Figure 1. Top: Map of Armenia and Nagorno-Karabakh 6. Vicinity of Amaras Monastery, Martuni Province, Republic (Artsakh) and surrounding countries showing the 3 Nagorno-Karabakh (552 m) (pers. obs., 2006). areas of relatively low elevation where turtles occur (A–C): A) 7. Vicinity of Getavan (village), bank of the Tartar River, Western Armenia, Arax River drainage; B) Southern Armenia and Nagorno-Karabakh, Arax River Drainage; C) Northern Shahumian Province, Nagorno-Karabakh (760 m). Armenia, Kura River drainage; Y) Yerevan. Bottom: Political (pers. obs., 2006). map of Armenia and Nagorno-Karabakh Republic (Artsakh) showing localities mentioned in the text. Mauremys caspica (Gmelin, 1774) decreasing reference quality (i.e., museum specimens, photographic vouchers, literature records, and nonvouch- The first report of M. caspica (Figs. 2G–L, 4) in this ered personal observations). We purposefully do not split region was by Me´he´ly (1894), based on a specimen in the these species into subspecies, following our previous MZIA collected in the vicinity of Yerevan (Chernov genetic study of the region (Parham et al. 2006; see also 1939). Chernov (1939) also reported 2 additional sites in Discussion). western Armenia based on other specimens from the MZIA. Since that time, M. caspica has been found in the RESULTS same 3 regions of low elevation as E. orbicularis (Fig. 1), although M. caspica has a much wider distribution (Fig. 4 Emys orbicularis (Linnaeus, 1758) vs. Fig. 3). Mauremys caspica is common in rivers that flow through semidesert habitats (Dal 1954), such as the The first report of E. orbicularis (Figs. 2A–F, 3) in Arax River and the lower reaches of the Akhuryan River. this region was by Lyajster (1912), who captured These observations were echoed by Darevsky (1957), who specimens from the Hrazdan (Zanga) River in the vicinity also noted that, in southern Armenia, M. caspica occupies of Yerevan, as well as the Arax River. Lyajster’s findings a variety of riparian and stagnant aquatic habitats. In some were published by Chernov (1939) and also can be found areas, M. caspica occupies mountain springs where they in the dissertation of Darevsky (1957). However, Dal settle in small bogs or pools. In periods of drought, these (1954) and Darevsky (1957) disagreed with the claim that turtles can be completely isolated from the main drainage. E. orbicularis occurs in the Arax valley (Lyajster 1912; According to our data, M. caspica occurs at an altitude Bobrinsky 1915, cited by Darevsky 1957; Skryabin 1924, from 237 m up to 1800 m. 72 CHELONIAN CONSERVATION AND BIOLOGY, Volume 7, Number 1 – 2008

1. Airum railway station, Debed River, Tavush Province. 20. Fish farm, vicinity of Surenavan (village), Ararat Darevsky (1957). Province (808–818 m) (pers. obs., 2004). 2. Above Uzuntale (Aygehovit) (village), Agshtev River, 21. Erasch (village), Arax River, Ararat Province (818 m) Tavush Province. Darevsky (1957). (pers. obs., 2004). 3. Byurakan (village), Amberd River, Aragatsotn Prov- 22. Khor Virap (village), River, Ararat Prov- ince. Darevsky (1957). ince (836 m). Digital voucher (2004; MVZ 250833). 4. Bagravan (village), at the confluence of the Akhuryan 23. Vicinity of Urtsadzor (village), Vedi River, Ararat and Arax rivers, Shirak Province. S. Pipoyan (pers. Province (994 m). Digital voucher (2004; MVZ 250834). obs., 1985). 24. Azizbekov (Vayk) (town), Arpa River, Vayots Dzor 5. Vicinity of Yerevan City, Yerevan Province. Museum Province (1360 m). Dal (1954). specimen (MZIA 510/1921). Chernov (1939). 25. At the confluence of the Arpa and Terp rivers, Vayots 6. Yerevan, Hrazdan River, Yerevan Province (840 m) Dzor Province. Darevsky (1957). (pers. obs., 2004) (A. Malkhasyan, pers. obs., 2004). 26. Lakes near Rind (village), Vayots Dzor Province 7. Vicinity of Geghard Monastery, Kotayk Province. (pers. obs., 2004). Museum specimen (MZIA 216/1927). Chernov 27. Lake near Srashen (village), Syunik Province (pers. (1939). obs., 2004). 8. Azat River, Kotayk Province (1500 m). Dal (1954). 28. Near Agarak (town), Syunik Province (663 m) (pers. 9. Arzakan (village), Hrazdan River, Kotayk Province obs., 2004). (1460 m). Dal (1954). 29. Lake near Meghri (town), Syunik Province (553 m) 10. Vicinity of Djrvej (village), near Yerevan, Kotayk (pers. obs., 2004). Province. Darevsky (1957). (S. Pipoyan, pers. obs., 30. Lake near Tzobi/Sobi (village), Tzobajur Valley, 1985). Basutajur, Kashatagh Province, Nagorno-Karabakh 11. Aygerlich (village), Province. Museum (453 m) (pers. obs., 2004). specimen (MZIA 511/1928). Chernov (1939). 31. Nine km southeast of Tzobi/Sobi (village), Tzobajur 12. Berkashat (village), . Darevsky Valley, Basutajur, Kashatagh Province, Nagorno-Kar- (1957). abakh (357 m). Digital voucher (2004; MVZ 250835). 13. Five km south of Metsamor (town), Armavir Province 32. Two km west of Midjnavan (town), Arax River, (868 m) (pers. obs., 2004). Kashatagh Province, Nagorno-Karabakh (341 m). 14. Lake near (village), Armavir Province (pers. obs., 2004). (859 m) (pers. obs., 2004). 33. Two km west of Hadjari (Krmen) (village), Hakari/ 15. Fish farm, vicinity of Arax (village), Armavir Akera River at the confluence with the Arax River, Province (857 m) (pers. obs., 2004). Kashatagh Province, Nagorno-Karabakh (290 m). 16. Baiburd (village), Garni River, Ararat Province. Digital voucher (2004; MVZ 230836). Darevsky (1957). 34. Mamark (village), Vorotan River, Kashatagh Prov- 17. Mkhchyan (village), Ararat Province. Darevsky ince, Nagorno-Karabakh (540 m). (pers. obs., 2004). (1957). 35. Two km east of Khudaperin (village), Arax River, 18. Ranchpar (village), Ararat Province (852 m) (pers. Hadrut Province, Nagorno-Karabakh (237 m). (pers. obs., 2004). Darevsky (1957). obs., 2004). 19. Azat Reservoir, Ararat Province (pers. obs., 1998) (A. 36. Aknakhbyur (village), Hudrut Province, Nagorno- Malkhasyan, pers. obs., 2004). Karabakh (568 m). (pers. obs., 2006).

! Figure 2. A) Dorsal view of Emys orbiculari from the Arax River Valley, Nagorno-Karabakh Republic (E. orbicularis locality no. 4, photo by R. Sindaco); B) Head of same individual in A (Photo by R. Sindaco); C) Adult E. orbicularis from the vicinity of Amaras Monastery, Martuni Province, Nagorno-Karabakh (E. orbicularis locality no. 5, photo by R. Sindaco); D) Head of same individual in C (Photo by R. Sindaco); E) Subadult E. orbicularis in a stream near Getavan Village, Shahumian Province, Nagorno-Karabakh (E. orbicularis locality no. 6, photo by R. Sindaco); F) Typical habitat of E. orbicularis, stream with slow-moving water and abundant aquatic vegetation (E. orbicularis locality no. 4, photo by MA); G) Dorsal view Mauremys caspica from Ararat Valley (M. caspica locality no. 23, photo by MA); H) Ventral view of same individuals in B; I) Subadult M. caspica from Aknakhbyur Village, Hudrut Province, Nagorno-Karabakh (M. caspica locality no. 36, photo by R. Sindaco); J–L) Typical habitats of M. caspica in Armenia and Nagorno-Karabakh Republic (Artsakh); J) Lake draining channels, Armavir region, southwest Armenia (M. caspica locality no. 13, photo by MA); K) Fish farms, Ararat region, southwest Armenia (M. caspica locality no. 21, photo by MA); L) Lake near Tzobi Village, Nagorno-Karabakh (M. caspica locality no. 30, photo by MA); M) Testudo graeca from vicinity of Martakert, Martakert Province, Nagorno-Karabakh (T. graeca locality no. 58 photo by R. Sindaco); N) T. graeca from the vicinity of Meghri (T. graeca locality no. 43, photo by MA); O) T. graeca from vicinity of Amaras Monastery (T. graeca locality no. 57, photo by R. Sindaco); P) Subadult T. graeca from Gorovan sand, Ararat region, southwest Armenia (T. graeca locality no. 33, photo by R. Sindaco; Q) Typical habitat of T. graeca in the vicinity of Meghri (T. graeca locality no. 43, photo by K. Milto); R) Typical habitat of T. graeca in the Gorovan sands (T. graeca locality no. 33, photo by MA). ARAKELYAN AND PARHAM — Geographic Distribution of Turtles in Armenia and the Nagorno-Karabakh Republic 73 74 CHELONIAN CONSERVATION AND BIOLOGY, Volume 7, Number 1 – 2008

Figure 3. Distribution of Emys orbicularis in Armenia and Figure 5. Distribution of Testudo graeca in Armenia and Nagorno-Karabakh Republic (Artsakh). Nagorno-Karabakh Republic (Artsakh).

Testudo graeca Linnaeus, 1758 which T. graeca has been found in Armenia is ca. 300 m The first report of T. graeca (Figs. 2M–R, 5, 6) in the (in Nagorno-Karabakh). Caucasus did not include a detailed locality (Pallas 1814). In western Armenia, T. graeca inhabits the Ararat In Armenia, the first specimens were collected by Chantre region to about 1000 m. According to the personal (1882) in the vicinity of Yerevan. Zugmayer (1906) observations of S. Pipoyan, T. graeca could be found near reported the first sightings from the Arax River valley, Yerevan (1100–1150 m) up to the early 1970s. In southern without giving a detailed locality. Nikolskii (1913) added Armenia, T. graeca occupies desert and semidesert areas. another location for Armenia. Chernov (1939), by using From the Arax River up to an elevation of 1300 m, T. materials of Institute of Zoology of Armenia, specified 7 graeca usually inhabits a semidesert landscape with scant additional localities for T. graeca in Armenia. Darevsky (1957) added 5 additional localities in northern Armenia. Since then, little effort has been made to record the distribution of T. graeca in Armenia. Our efforts resulted in 57 localities in Armenia and Nagorno-Karabakh. In this area, T. graeca can be found in mountains up to 1500 m (vicinity of the village of Aragyuh). The lowest altitude at

Figure 6. A) Adult Testudo graeca from the Gorovan sands, Ararat region, southwest Armenia (T. graeca locality no. 33, photo by MA); B) T. graeca from vicinity of Aragyugh, Kotayak Figure 4. Distribution of Mauremys caspica in Armenia and Province, central Armenia (T. graeca locality no. 28, photo by Nagorno-Karabakh Republic (Artsakh). MA). ARAKELYAN AND PARHAM — Geographic Distribution of Turtles in Armenia and the Nagorno-Karabakh Republic 75 vegetation, such as wormwoods (e.g., Artemisia fragrans, 18. Vicinity of Koghb (village), Tavush Province (A. Artemisia araxina) and other associated plants. The Virabyan, pers. obs., 2004). vegetation on the arid and rocky hills is predominantly 19. Akhtala (village), Lori Province. Museum specimen xerophilous, with large abundant tragacanth shrubs. In the (MZIA154/1925, 593/1925, 515/1925). Chernov Meghri region, they are found in rocky gorges, deciduous (1939). woodlands, and arid open juniper woodlands. The forests 20. Shnogh (village), Lori Province. Museum specimen of Meghri are composed of oak, hornbeam, ash, and (MZIA 386/1932). Chernov (1939) (pers. obs., 1999). juniper, in addition to wild fruit trees, such as pear, cornel, 21. Vicinity of Alaverdi (town), Lori Province. Darevsky walnut, hazel, and plum. Various bushes, such as (1957). hawthorn, woodbine, dog-rose, and blackberry bushes, 22. Vicinity of Shamlugh (village), Lori Province. (E. are also prevalent. In these areas, T. graeca is commonly Yavruyan, pers. obs., 2000) found in vineyards, gardens, and agricultural fields where 23. Road between Shamlugh (village) and Akhtala they freely feed on cultivated plants (Taskavak et al. (village), Lori Province. (E. Yavruyan, pers. obs., 2004). 1998). In northeastern Armenia, T. graeca are found in 24. ‘‘Arax valley.’’ (Zugmayer, 1906). forests of the Kura River drainage. In this area, T. graeca 25. Vicinity of Yerevan city, Yerevan Province. (Chantre, inhabits rocky hillsides and flat peaks of oak light forest 1883; Dal, 1954). and fringe of the oak-hornbeam forest at an elevation of up 26. ‘‘Yerevan.’’ Museum specimens (ZISP 9015a, b, c). to 950 m. We found T. graeca in agricultural fields, (S. Pipoyan, pers. obs., ,1970). gardens, and even a cemetery. 27. Vicinity of Yerevan, near Voghdjaberd (village), Kotayk Province (A. Malkhasyan, pers. obs., 2004). 1. Lchkadzor (village), Tavush Province. Museum spec- 28. Vicinity of Aragyugh (village), Kotayak Province imen (MZIA 504/1932). Chernov (1939). (1500 m). Digital voucher (2004; MVZ 230838). 2. Paravakar (village), Tavush Province. Museum spec- 29. Vicinity of Aralikh (Griboedov) (village). Armavir imen (MZIA 505/1934). Chernov (1939) (A. Virab- Province. Nikolskii (1913) yan, pers. obs., 2004). 30. Vicinity of New Shavarut (Hushakert) (village), 3. Between the towns of Kuibishev (Dilijan) and Idjevan, Armavir Province. Dal (1954). Tavush Province. Darevsky (1957). 31. Vicinity of Armavir (town), Armavir Province (S. 4. Vicinity of Noemberyan (town), Tavush Province Pipoyan, pers. obs., 2004). (868 m). Darevsky (1957) (pers. obs., 1998). 32. Vicinity of Vedi (town), Ararat Province. Museum 5. Vicinity of Lambalu (Bagratashen) (village), Tavush specimen (MZIA169/1926, 500/1927). Chernov Province. Darevsky (1957). (1939). 6. Vicinity of Uzuntala (Aygehovit) (village), Tavush 33. Gorovan sand, Ararat Province (911 m). Digital Province. Darevsky (1957). voucher (2004; MVZ 230839). Tuniev and Unanyan 7. Vicinity of Berd (town), Tavush Province. Digital (1986). voucher (MVZ 230837) (A. Virabyan, pers. obs., 34. Vicinity of Davalu (Ararat) (town), Ararat Province. 2004). Chernov (1939). 8. Vicinity of Aigepar (village), Tavush Province (A. 35. Vicinity of Armash (village), Ararat Province. Muse- Virabyan, pers. obs., 2004). um specimen (MZIA 1930). Chernov (1939). 9. Vicinity of Tovuz (village), Tavush Province (A. 36. Near Khor Virap (village), Ararat Province (837 m) Virabyan, pers. obs., 2004). (pers. obs., 2000). 10. Vicinity of Nerkin Karmir (village), Tavush Province 37. Near Urtsadzor (village), Vedi River, Ararat Province (A. Virabyan, pers. obs., 2004). (1062 m) (pers. obs., 2004). 11. Vicinity of Aigedzor (village), Tavush Province (A. 38. Vicinity of Aragats Station (Aragats village), Aragat- Virabyan, pers. obs., 2004). sotn Province. Dal (1954). 12. Vicinity of Chinari (village), Tavush Province (A. 39. Vicinity of Rind (village), Vayots Dzor Province. Virabyan, pers. obs., 2004). (pers. obs., 2004) 13. Vicinity of Choratan (village), Tavush Province (A. 40. Vicinity of Nyuvadi (Nrnadzor) (village), Syunik Virabyan, pers. obs., 2004). Province. Museum specimen (ZISP 15943). Dal 14. Vicinity of Mosesgegh (village), Tavush Province (A. (1954) (pers. obs., 2004). Virabyan, pers. obs., 2004). 41. Meghri (town), Syunik Province (721 m). Museum 15. Vicinity of Tsachkavan (village), Tavush Province (A. specimen (ZIN 23026). Chikhvadze and Barkadze Virabyan, pers. obs., 2004). (1991), Taskavak et al. (2004), Parham et al. (2006). 16. Vicinity of Varagavan (village), Tavush Province (A. 42. Agarak (town), Syunik Province (663 m) (pers. obs., Virabyan, pers. obs., 2004). 2004). 17. Vicinity of Airum (village), Tavush Province (A. 43. Two km east of Meghri (town), Syunik Province (546 Virabyan, pers. obs., 2004). m). Digital voucher (2004; MVZ 230840). 76 CHELONIAN CONSERVATION AND BIOLOGY, Volume 7, Number 1 – 2008

44. Eleven km east of Meghri (town), Syunik Province Armenia, these 3 main turtle areas are not allopatric, (515 m). Museum specimen (ZIN 23025). Parham because they are connected outside of the territory studied et al. (2006) here. 45. Bussakiar (village), Kashatagh Province, Nagorno- From a phylogeographic perspective, the 3 main turtle Karabakh. Pieh et al. (2002). areas form a transect that is in need of further study. For 46. Vicinity of Stepanakert (capital), Askeran Province, example, the Arax populations of T. graeca have been Nagorno-Karabakh. Alekperov (1978). referred to a new taxon Testudo graeca armeniaca 47. Vicinity of Aghdam (town), Aghdam Province, Chkhikvadze and Bakradze, 1991. The populations in Nagorno-Karabakh. Alekperov (1978). northeast Armenia are very close to the type locality of 48. Vicinity of Zangelan (Kovsakan) (town), Kashatagh Testudo graeca ibera (Pallas 1814; near Tbilisi, Georgia, Province, Nagorno-Karabakh. Alekperov (1978). by subsequent designation of Bour 1987 in accordance 49. Vicinity of Martuni (town), Martuni Province, Na- with Eichwald 1831) and might be referred to this taxon. A gorno-Karabakh. Alekperov (1978). recent genetic study (Parham et al. 2006) confirmed that 50. Near Lachin (Berdzor) (town), Kashatagh Province, tortoises from each of the type localities are in separate Nagorno-Karabakh. S. Pipoyan (pers. comm.) mitochondrial clades. Fritz et al. (2007) suggested that 51. Vicinity of Aghbend (village), 8 km from the border of these clades could be considered subspecies; whereas, Armenia on the west bank of the Arax River, Parham et al. (2006) made no such designation. Part of the Kashatagh Province, Nagorno-Karabakh (360 m). problem is that (as pointed out by Parham et al. 2006) Digital voucher (2004; MVZ 230841). tortoises from nearby the type locality of T. g. armeniaca 52. Seven km southeast of Tzobi/Sobi (village), Tzobajur (Meghri, by original designation) can exhibit dramatically River valley, Basutajur, Kashatagh Province, Nagor- different morphologies, despite being in the same no-Karabakh (357 m). Digital voucher (2004; MVZ mitochondrial clade. Clearly more fine-grained studies 230842). (pers. obs., 2006). that integrate morphology with mitochondrial DNA and 53. Four km northwest of Chopadara (village), Tzobajur other genetic markers are needed, especially where River valley, Basutajur, Kashatagh Province, Nagor- different morphotypes and genetic clades come into no-Karabakh (432 m). Digital voucher (2004; MVZ contact. In this respect, the Nagorno-Karabakh populations 250843). of T. graeca are especially interesting because they occupy 54. Five km east of Krmen (village), west bank of the the area intervening the Kura and Arax drainages where 2 Hakari (Akera) River, at the confluence with the Arax different taxa/clades occur. European pond turtles may River, Kashatagh Province, Nagorno-Karabakh (301 exhibit similar genetic differentiation within Armenia. The m). Digital voucher (2004; MVZ 250844). Emys populations in northeast Armenia (in the Kura 55. Nine km north of Varanda (town), Martuni Province, drainage) are referred to E. o. iberica Eichwald, 1831 Nagorno-Karabakh (322 m). Digital voucher (2004; (Fritz 1994); whereas, the populations of the Arax are MVZ 250845). taxonomically unassigned pending further study. The 56. Seven km west of Martuni (town), Martuni Province, details of Arax drainage endemism in turtles is an Nagorno-Karabakh (364 m) (pers. obs., 2003). important area for future study aimed at better defining 57. Two km southwest of Amaras Monastery, Martuni taxonomic units with the Caucasus. Province, Nagorno-Karabakh (729 m) (pers. obs., The historic data are not sufficient to track recent 2006). changes in abundance or distribution. However, we did 58. Four km north of Martakert (town), Martakert record the apparent extirpation of T. graeca from the close Province, Nagorno-Karabakh (556 m) (pers. obs., vicinity of Yerevan (likely because of urbanization). 2006). However, Yerevan is near the altitudinal limit of T. graeca in Armenia (.1000 m), so the tortoise may not have been very abundant there. Our data provide a good baseline for DISCUSSION tracking additional extirpations, as well as for future studies aimed at the conservation status, ecology, or The geographic distributions of Armenian turtles are systematics of Caucasus turtles. defined by elevation. The average altitude of Armenia is 1850 m, which is higher than any turtle in Armenia is ACKNOWLEDGMENTS known to occur. Mauremys caspica was recorded at elevations of up to 1800 m in Armenia; whereas, T. graeca This research was made possible by a Linnaeus Fund is almost always found below 1500 m, and E. orbicularis Research Grant. We thank the Armenian biologists that is found even lower (below 1000 m). Because most of the shared their data on the distribution of Armenian turtles: country is uninhabitable for turtles, populations are Prof Felix Danielyan, Dr Aram Agasyan, Dr Samvel restricted to the lowest portions of 2 distinct drainages: Pipoyan, Dr Eduard Yavruyan, Alexander Malkhasyan, the Kura River drainage in the northeast and the Arax and Andranik Virabyan. Ted Papenfuss (U.C. Berkeley) River valley in the south and west. Although disjunct in also provided invaluable assistance. Special thanks to ARAKELYAN AND PARHAM — Geographic Distribution of Turtles in Armenia and the Nagorno-Karabakh Republic 77

Jarmo Pera¨la¨ for kindly providing essential information on LEHMANN, J., MAZANAEVA, L.F., TU¨ RKOZA, O., AND WINK,M. turtles from the Zoological Institute of St. Petersburg, 2007. Phenotypic plasticity leads to incongruence between Russia. We also thank Uwe Fritz for reviewing an earlier morphology-based taxonomy and genetic differentiation in draft of the manuscript. This is University of California western Palaearctic tortoises (Testudo graeca complex; Testu- dines, Testudinidae) Amphibia-Reptilia 28:97–121. Museum of Paleontology contribution no. 1922. GMELIN, S.G. 1774. Reise durch Russland zur Untersuchung der drey Natur-Reiche. 3:59. LITERATURE CITED LYAJSTER, A.F. 1912. The fauna of Erebuni Province. Reptiles of Erebuni Province. The Records of Erebuni Province for 1912. ALEKPEROV, A.M. 1978. Amphibians and Reptiles of Azerbaijan Yerevan: ‘‘Department of Literature.’’ [cited by Darevsky, [in Russian]. Baku, Azerbaijani: Elm Press, 264 pp. 1957] BOBRINSKY, N.A. 1915. Results of ornithological expeditions to Suriman and Echmiadzin districts of Yerevan Province in the ME´ HE´ LY, L. 1894. Beitrage zur Herpetologie. Transkaukasiens summer of 1911 and 1912 [in Russian]. Report of the und Armenians. Zoologischer Anzeiger 17:1–78 Caucasian Museum 8(3):171–250. MOVSESYAN, S.O. (ED.) 1987. Red Data Book of Armenian S.S.R. BOUR, R. 1987. L’identite´ des Tortues terrestres europe´ennes: Animals [in Russian]. Yerevan, Armenia: Hajastan Press, 124 spe´cimens-types et localite´s-types. Revue franc¸aise d’Aqua- pp. riologie 13:111–122. NIKOLSKII, A.M. 1913. Reptiles and Amphibian of Caucasus [in CHANTRE, E. 1882. Rapport sur une mission scientifique dans Russian]. Tbilisi, Georgia: Caucasus Press, 272 pp. l’Asie occidentale et specialement dans les regions de l’Ararat PALLAS, P.S. 1814. Zoographia Rosso-Asiatici. Tom III [in Latin]. et Caucase. Archives de Mission Scientifiques et Letters, Paris Animalia Monocardia seu Frigidi Sanguinis Imperii Rosso- 3(10):199–263. Asiatica. Petropolis: ex officina Caes. Academiae Scientarum, CHERNOV, S.A. 1939. Herpetofauna of Armenian SSR and 428 pp. Nakhichevan ASSR [in Russian]. – Zoological collection of PARHAM, J.F., TU¨ RKOZAN, O., STUART, B.L., ARAKELYAN, M., articles of Armenian branch of the Academy of Science USSR SHAFEI, S., MACEY, J.R., AND PAPENFUSS, T.J. 2006. Genetic 1:79–194. evidence for premature taxonomic inflation in Middle Eastern CHKHIKVADZE, V.M. AND BAKRADZE, M.A. 1991. On the systematic tortoises. Proceedings of the California Academy of Sciences position of the Recent land turtle of from the Arax valley [in 57(3):955–964. Russian]. Trudy Tbilisskogo Gosudartswenogo Univiversiteta PIEH, A., FRITZ, U., AND BERGLAS, R. 2002. New data on the [The works of Tbilisi State University] Tbilisi 305:59–63. morphology, distribution and nomenclature of Testudo graeca DAL, S.K. 1954. Fauna of Armenian SSR [in Russian]. V1. armeniaca Chkhikvadze and Bakradze, 1991 (Reptilia: Yerevan: Academy Science of Armenian SSR, 415 pp. Testudines: Testudinidae). Faunitische Abhandlungen Staat- DAREVSKY, I.S. 1957. The reptile fauna of Armenia and its litches Museum fu¨r Tierkunde Dresden 22(21):329–345. zoogeographical analysis [in Russian]. PhD Thesis, Zoological SKRYABIN, K.I. 1924. Trematods of Emys orbicularis from valley Institute of the Armenian Academy of Sciences. Yerevan, of Arax River [in Russian]. Trudi Tropicheskogo Instituta Armenia. Armenia, SSR. 1:34–40. EICHWALD, C.E. VON 1831. Zoologica specialis quam expositas TASKAVAK, E., ARAKELYAN, M., AND PARHAM, J.F. 2004. Testudo animalibus. Pars posterior. Vilna, Lithuania: J. Zawadzki, 404 [graeca] armeniaca. Manouria 7(22):7–8. pp. TUNIEV, B.S. AND UNANYAN, E.S. 1986. Species structure and daily FRITZ, U. 1994. Zur innerartlichen Variabilitat von Emys orbicularis (Linneus, 1758). 4. Variabilita¨t und Zoogeographie activity of reptiles of Khosrov State Reserve [in Russian]. im pontoscaspischen Gebeit mit Beschreibung von drei neuen Biological Zhurnal of Armenia 1(7):572–576. Unterarten (Reptilia: Testudines: Emydidae). Faunitische ZUGMAYER, E. 1906. Beitra¨ge zur Herpetologie von Vorder-Asien. Abhandlungen Staatlitches Museum fu¨r Tierkunde Dresden Zoologische Jahrbu¨cher, Abteilung Systematik 23:449–486. 48(4):53–93. Received: 3 July 2006 FRITZ, U., HUNDSDO¨ RFER, A.K., SIROKYI`, P., AUER, M., KAMI, H., Revised and Accepted: 7 April 2008