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Online Publication Date: 01 Aug 2008

The Geographic Distribution of Turtles in Armenia and the Nagorno-Karabakh Republic (Artsakh)

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Article Category: Research Article
Page Range: 70 – 77
DOI: 10.2744/CCB-0664.1
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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.

Keywords: 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 and covers approximately 10% of the Armenian plateau (29,800 km2). It is bordered by Azerbaijan, Georgia, Iran, and Turkey. Nagorno-Karabakh Republic (also known as Artsakh) is a smaller (4400 km2) autonomous region between Armenia and eastern Azerbaijan. As a result of their geographical position and diversity of landscapes, both Armenia and Nagorno-Karabakh support a wide range of animal species, including 3 species of turtles. The turtles of Armenia are from each of the 3 families of Testudinoidea: 1) an emydid, Emys orbicularis (Linnaeus, 1758); 2) a geoemydid, Mauremys caspica (Gmelin 1774); and 3) a testudinid, Testudo graeca Linnaeus, 1758.

Figure 1. Top: Map of Armenia and Nagorno-Karabakh Republic (Artsakh) and surrounding countries showing the 3 areas of relatively low elevation where turtles occur (A–C): A) Western Armenia, Arax River drainage; B) Southern Armenia and Nagorno-Karabakh, Arax River Drainage; C) Northern Armenia, Kura River drainage; Y) Yerevan. Bottom: Political map of Armenia and Nagorno-Karabakh Republic (Artsakh) showing localities mentioned in the text.Figure 1. Top: Map of Armenia and Nagorno-Karabakh Republic (Artsakh) and surrounding countries showing the 3 areas of relatively low elevation where turtles occur (A–C): A) Western Armenia, Arax River drainage; B) Southern Armenia and Nagorno-Karabakh, Arax River Drainage; C) Northern Armenia, Kura River drainage; Y) Yerevan. Bottom: Political map of Armenia and Nagorno-Karabakh Republic (Artsakh) showing localities mentioned in the text.Figure 1. Top: Map of Armenia and Nagorno-Karabakh Republic (Artsakh) and surrounding countries showing the 3 areas of relatively low elevation where turtles occur (A–C): A) Western Armenia, Arax River drainage; B) Southern Armenia and Nagorno-Karabakh, Arax River Drainage; C) Northern Armenia, Kura River drainage; Y) Yerevan. Bottom: Political map of Armenia and Nagorno-Karabakh Republic (Artsakh) showing localities mentioned in the text.
Figure 1. Top: Map of Armenia and Nagorno-Karabakh Republic (Artsakh) and surrounding countries showing the 3 areas of relatively low elevation where turtles occur (A–C): A) Western Armenia, Arax River drainage; B) Southern Armenia and Nagorno-Karabakh, Arax River Drainage; C) Northern Armenia, Kura River drainage; Y) Yerevan. Bottom: Political map of Armenia and Nagorno-Karabakh Republic (Artsakh) showing localities mentioned in the text.

Citation: Chelonian Conservation and Biology 7, 1; 10.2744/CCB-0664.1

Testudo graeca is listed in the Red Data Book of Armenia (Movsesyan 1987), which provides it with nominal protection as a threatened species. However, the baseline data on its geographic distribution, crucial components for effective conservation management, are lacking. Some locality data are found in the works of Chernov (1939) and Dal (1954). But since then, Darevsky's unpublished PhD thesis (1957) is the main report on distribution of turtles in Armenia and surrounding regions. Because these publications are written in Russian, they are overlooked by most readers. The objective of this research is to gather baseline data on the geographical distribution of the turtle fauna in these regions by using personal observations, literature records, and museum specimens. It is our hope that this report will provide a good starting point for future studies on the distribution and conservation status of turtles in this region.

MATERIALS AND METHODS

The data for this article are from 3 kinds of sources: 1) a review of the pertinent literature; 2) personal observation of the authors, as well as other Armenian biologists; 3) museum specimens from the Zoological Institute of St. Petersburg, Russia (ZISP), and the Museum of the Zoological Institute of Armenia, Yerevan, Armenia (MZIA). Personal observations of turtles in the wild were primarily by MA unless stated otherwise, especially in 2004 and incidentally from 1994 to 2006. Global positioning system (GPS) coordinates were taken at every locality and all turtles were vouchered via photography. These images were then made into vouchers (both hard and digital) and deposited into the Museum of Vertebrate Zoology, University of California, Berkeley, California, USA (MVZ). The GPS coordinates are also stored at the MVZ and are available to qualified researchers. Additional information was collected from interviews with Armenian biologists (especially herpetologists) and local people. In every case, we indicated the source of these data, recognizing that each constitutes a different class of information that can be confirmed or refuted with later study. In instances where multiple sources report the same locality, we listed them in an approximate order of decreasing reference quality (i.e., museum specimens, photographic vouchers, literature records, and nonvouchered personal observations). We purposefully do not split these species into subspecies, following our previous genetic study of the region (Parham et al. 2006; see also Discussion).

RESULTS

Emys orbicularis (Linnaeus, 1758)

The first report of E. orbicularis (Figs. 2A–F, 3) in this region was by Lyajster (1912), who captured specimens from the Hrazdan (Zanga) River in the vicinity of Yerevan, as well as the Arax River. Lyajster's findings were published by Chernov (1939) and also can be found in the dissertation of Darevsky (1957). However, Dal (1954) and Darevsky (1957) disagreed with the claim that E. orbicularis occurs in the Arax valley (Lyajster 1912; Bobrinsky 1915, cited by Darevsky 1957; Skryabin 1924, 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. Subsequently, 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).

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).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).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).
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).

Citation: Chelonian Conservation and Biology 7, 1; 10.2744/CCB-0664.1

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

Citation: Chelonian Conservation and Biology 7, 1; 10.2744/CCB-0664.1

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).

  6. Vicinity of Amaras Monastery, Martuni Province, Nagorno-Karabakh (552 m) (pers. obs., 2006).

  7. Vicinity of Getavan (village), bank of the Tartar River, Shahumian Province, Nagorno-Karabakh (760 m). (pers. obs., 2006).

Mauremys caspica (Gmelin, 1774)

The first report of M. caspica (Figs. 2G–L, 4) in this region was by Méhély (1894), based on a specimen in the MZIA collected in the vicinity of Yerevan (Chernov 1939). Chernov (1939) also reported 2 additional sites in western Armenia based on other specimens from the MZIA. Since that time, M. caspica has been found in the same 3 regions of low elevation as E. orbicularis (Fig. 1), although M. caspica has a much wider distribution (Fig. 4 vs. Fig. 3). Mauremys caspica is common in rivers that flow through semidesert habitats (Dal 1954), such as the Arax River and the lower reaches of the Akhuryan River. These observations were echoed by Darevsky (1957), who also noted that, in southern Armenia, M. caspica occupies a variety of riparian and stagnant aquatic habitats. In some areas, M. caspica occupies mountain springs where they settle in small bogs or pools. In periods of drought, these turtles can be completely isolated from the main drainage. According to our data, M. caspica occurs at an altitude from 237 m up to 1800 m.

Figure 4. Distribution of Mauremys caspica in Armenia and Nagorno-Karabakh Republic (Artsakh).Figure 4. Distribution of Mauremys caspica in Armenia and Nagorno-Karabakh Republic (Artsakh).Figure 4. Distribution of Mauremys caspica in Armenia and Nagorno-Karabakh Republic (Artsakh).
Figure 4. Distribution of Mauremys caspica in Armenia and Nagorno-Karabakh Republic (Artsakh).

Citation: Chelonian Conservation and Biology 7, 1; 10.2744/CCB-0664.1

  1. Airum railway station, Debed River, Tavush Province. Darevsky (1957).

  2. Above Uzuntale (Aygehovit) (village), Agshtev River, Tavush Province. Darevsky (1957).

  3. Byurakan (village), Amberd River, Aragatsotn Province. Darevsky (1957).

  4. Bagravan (village), at the confluence of the Akhuryan and Arax rivers, Shirak Province. S. Pipoyan (pers. obs., 1985).

  5. Vicinity of Yerevan City, Yerevan Province. Museum specimen (MZIA 510/1921). Chernov (1939).

  6. Yerevan, Hrazdan River, Yerevan Province (840 m) (pers. obs., 2004) (A. Malkhasyan, pers. obs., 2004).

  7. Vicinity of Geghard Monastery, Kotayk Province. Museum specimen (MZIA 216/1927). Chernov (1939).

  8. Azat River, Kotayk Province (1500 m). Dal (1954).

  9. Arzakan (village), Hrazdan River, Kotayk Province (1460 m). Dal (1954).

  10. Vicinity of Djrvej (village), near Yerevan, Kotayk Province. Darevsky (1957). (S. Pipoyan, pers. obs., 1985).

  11. Aygerlich (village), Armavir Province. Museum specimen (MZIA 511/1928). Chernov (1939).

  12. Berkashat (village), Armavir Province. Darevsky (1957).

  13. Five km south of Metsamor (town), Armavir Province (868 m) (pers. obs., 2004).

  14. Lake near Yeraskhahun (village), Armavir Province (859 m) (pers. obs., 2004).

  15. Fish farm, vicinity of Arax (village), Armavir Province (857 m) (pers. obs., 2004).

  16. Baiburd (village), Garni River, Ararat Province. Darevsky (1957).

  17. Mkhchyan (village), Ararat Province. Darevsky (1957).

  18. Ranchpar (village), Ararat Province (852 m) (pers. obs., 2004). Darevsky (1957).

  19. Azat Reservoir, Ararat Province (pers. obs., 1998) (A. Malkhasyan, pers. obs., 2004).

  20. Fish farm, vicinity of Surenavan (village), Ararat Province (808–818 m) (pers. obs., 2004).

  21. Erasch (village), Arax River, Ararat Province (818 m) (pers. obs., 2004).

  22. Khor Virap (village), Metsamor River, Ararat Province (836 m). Digital voucher (2004; MVZ 250833).

  23. Vicinity of Urtsadzor (village), Vedi River, Ararat Province (994 m). Digital voucher (2004; MVZ 250834).

  24. Azizbekov (Vayk) (town), Arpa River, Vayots Dzor Province (1360 m). Dal (1954).

  25. At the confluence of the Arpa and Terp rivers, Vayots Dzor Province. Darevsky (1957).

  26. Lakes near Rind (village), Vayots Dzor Province (pers. obs., 2004).

  27. Lake near Srashen (village), Syunik Province (pers. obs., 2004).

  28. Near Agarak (town), Syunik Province (663 m) (pers. obs., 2004).

  29. Lake near Meghri (town), Syunik Province (553 m) (pers. obs., 2004).

  30. Lake near Tzobi/Sobi (village), Tzobajur Valley, Basutajur, Kashatagh Province, Nagorno-Karabakh (453 m) (pers. obs., 2004).

  31. Nine km southeast of Tzobi/Sobi (village), Tzobajur Valley, Basutajur, Kashatagh Province, Nagorno-Karabakh (357 m). Digital voucher (2004; MVZ 250835).

  32. Two km west of Midjnavan (town), Arax River, Kashatagh Province, Nagorno-Karabakh (341 m). (pers. obs., 2004).

  33. Two km west of Hadjari (Krmen) (village), Hakari/Akera River at the confluence with the Arax River, Kashatagh Province, Nagorno-Karabakh (290 m). Digital voucher (2004; MVZ 230836).

  34. Mamark (village), Vorotan River, Kashatagh Province, Nagorno-Karabakh (540 m). (pers. obs., 2004).

  35. Two km east of Khudaperin (village), Arax River, Hadrut Province, Nagorno-Karabakh (237 m). (pers. obs., 2004).

  36. Aknakhbyur (village), Hudrut Province, Nagorno-Karabakh (568 m). (pers. obs., 2006).

Testudo graeca Linnaeus, 1758

The first report of T. graeca (Figs. 2M–R, 5, 6) in the Caucasus did not include a detailed locality (Pallas 1814). In Armenia, the first specimens were collected by Chantre (1882) in the vicinity of Yerevan. Zugmayer (1906) reported the first sightings from the Arax River valley, without giving a detailed locality. Nikolskii (1913) added another location for Armenia. Chernov (1939), by using materials of Institute of Zoology of Armenia, specified 7 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 which T. graeca has been found in Armenia is ca. 300 m (in Nagorno-Karabakh).

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

Citation: Chelonian Conservation and Biology 7, 1; 10.2744/CCB-0664.1

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 Province, central Armenia (T. graeca locality no. 28, photo by MA).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 Province, central Armenia (T. graeca locality no. 28, photo by MA).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 Province, central Armenia (T. graeca locality no. 28, photo by MA).
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 Province, central Armenia (T. graeca locality no. 28, photo by MA).

Citation: Chelonian Conservation and Biology 7, 1; 10.2744/CCB-0664.1

In western Armenia, T. graeca inhabits the Ararat region to about 1000 m. According to the personal observations of S. Pipoyan, T. graeca could be found near Yerevan (1100–1150 m) up to the early 1970s. In southern Armenia, T. graeca occupies desert and semidesert areas. From the Arax River up to an elevation of 1300 m, T. graeca usually inhabits a semidesert landscape with scant vegetation, such as wormwoods (e.g., Artemisia fragrans, Artemisia araxina) and other associated plants. The vegetation on the arid and rocky hills is predominantly xerophilous, with large abundant tragacanth shrubs. In the Meghri region, they are found in rocky gorges, deciduous woodlands, and arid open juniper woodlands. The forests of Meghri are composed of oak, hornbeam, ash, and juniper, in addition to wild fruit trees, such as pear, cornel, walnut, hazel, and plum. Various bushes, such as hawthorn, woodbine, dog-rose, and blackberry bushes, are also prevalent. In these areas, T. graeca is commonly found in vineyards, gardens, and agricultural fields where they freely feed on cultivated plants (Taskavak et al. 2004).

In northeastern Armenia, T. graeca are found in forests of the Kura River drainage. In this area, T. graeca inhabits rocky hillsides and flat peaks of oak light forest and fringe of the oak-hornbeam forest at an elevation of up to 950 m. We found T. graeca in agricultural fields, gardens, and even a cemetery.

  1. Lchkadzor (village), Tavush Province. Museum specimen (MZIA 504/1932). Chernov (1939).

  2. Paravakar (village), Tavush Province. Museum specimen (MZIA 505/1934). Chernov (1939) (A. Virabyan, pers. obs., 2004).

  3. Between the towns of Kuibishev (Dilijan) and Idjevan, Tavush Province. Darevsky (1957).

  4. Vicinity of Noemberyan (town), Tavush Province (868 m). Darevsky (1957) (pers. obs., 1998).

  5. Vicinity of Lambalu (Bagratashen) (village), Tavush Province. Darevsky (1957).

  6. Vicinity of Uzuntala (Aygehovit) (village), Tavush Province. Darevsky (1957).

  7. Vicinity of Berd (town), Tavush Province. Digital voucher (MVZ 230837) (A. Virabyan, pers. obs., 2004).

  8. Vicinity of Aigepar (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  9. Vicinity of Tovuz (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  10. Vicinity of Nerkin Karmir (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  11. Vicinity of Aigedzor (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  12. Vicinity of Chinari (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  13. Vicinity of Choratan (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  14. Vicinity of Mosesgegh (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  15. Vicinity of Tsachkavan (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  16. Vicinity of Varagavan (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  17. Vicinity of Airum (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  18. Vicinity of Koghb (village), Tavush Province (A. Virabyan, pers. obs., 2004).

  19. Akhtala (village), Lori Province. Museum specimen (MZIA154/1925, 593/1925, 515/1925). Chernov (1939).

  20. Shnogh (village), Lori Province. Museum specimen (MZIA 386/1932). Chernov (1939) (pers. obs., 1999).

  21. Vicinity of Alaverdi (town), Lori Province. Darevsky (1957).

  22. Vicinity of Shamlugh (village), Lori Province. (E. Yavruyan, pers. obs., 2000)

  23. Road between Shamlugh (village) and Akhtala (village), Lori Province. (E. Yavruyan, pers. obs., 1998).

  24. “Arax valley.” (Zugmayer, 1906).

  25. Vicinity of Yerevan city, Yerevan Province. (Chantre, 1883; Dal, 1954).

  26. “Yerevan.” Museum specimens (ZISP 9015a, b, c). (S. Pipoyan, pers. obs., <1970).

  27. Vicinity of Yerevan, near Voghdjaberd (village), Kotayk Province (A. Malkhasyan, pers. obs., 2004).

  28. Vicinity of Aragyugh (village), Kotayak Province (1500 m). Digital voucher (2004; MVZ 230838).

  29. Vicinity of Aralikh (Griboedov) (village). Armavir Province. Nikolskii (1913)

  30. Vicinity of New Shavarut (Hushakert) (village), Armavir Province. Dal (1954).

  31. Vicinity of Armavir (town), Armavir Province (S. Pipoyan, pers. obs., 2004).

  32. Vicinity of Vedi (town), Ararat Province. Museum specimen (MZIA169/1926, 500/1927). Chernov (1939).

  33. Gorovan sand, Ararat Province (911 m). Digital voucher (2004; MVZ 230839). Tuniev and Unanyan (1986).

  34. Vicinity of Davalu (Ararat) (town), Ararat Province. Chernov (1939).

  35. Vicinity of Armash (village), Ararat Province. Museum specimen (MZIA 1930). Chernov (1939).

  36. Near Khor Virap (village), Ararat Province (837 m) (pers. obs., 2000).

  37. Near Urtsadzor (village), Vedi River, Ararat Province (1062 m) (pers. obs., 2004).

  38. Vicinity of Aragats Station (Aragats village), Aragatsotn Province. Dal (1954).

  39. Vicinity of Rind (village), Vayots Dzor Province. (pers. obs., 2004)

  40. Vicinity of Nyuvadi (Nrnadzor) (village), Syunik Province. Museum specimen (ZISP 15943). Dal (1954) (pers. obs., 2004).

  41. Meghri (town), Syunik Province (721 m). Museum specimen (ZIN 23026). Chikhvadze and Barkadze (1991), Taskavak et al. (2004), Parham et al. (2006).

  42. Agarak (town), Syunik Province (663 m) (pers. obs., 2004).

  43. Two km east of Meghri (town), Syunik Province (546 m). Digital voucher (2004; MVZ 230840).

  44. Eleven km east of Meghri (town), Syunik Province (515 m). Museum specimen (ZIN 23025). Parham et al. (2006)

  45. Bussakiar (village), Kashatagh Province, Nagorno-Karabakh. Pieh et al. (2002).

  46. Vicinity of Stepanakert (capital), Askeran Province, Nagorno-Karabakh. Alekperov (1978).

  47. Vicinity of Aghdam (town), Aghdam Province, Nagorno-Karabakh. Alekperov (1978).

  48. Vicinity of Zangelan (Kovsakan) (town), Kashatagh Province, Nagorno-Karabakh. Alekperov (1978).

  49. Vicinity of Martuni (town), Martuni Province, Nagorno-Karabakh. Alekperov (1978).

  50. Near Lachin (Berdzor) (town), Kashatagh Province, Nagorno-Karabakh. S. Pipoyan (pers. comm.)

  51. Vicinity of Aghbend (village), 8 km from the border of Armenia on the west bank of the Arax River, Kashatagh Province, Nagorno-Karabakh (360 m). Digital voucher (2004; MVZ 230841).

  52. Seven km southeast of Tzobi/Sobi (village), Tzobajur River valley, Basutajur, Kashatagh Province, Nagorno-Karabakh (357 m). Digital voucher (2004; MVZ 230842). (pers. obs., 2006).

  53. Four km northwest of Chopadara (village), Tzobajur River valley, Basutajur, Kashatagh Province, Nagorno-Karabakh (432 m). Digital voucher (2004; MVZ 250843).

  54. Five km east of Krmen (village), west bank of the Hakari (Akera) River, at the confluence with the Arax River, Kashatagh Province, Nagorno-Karabakh (301 m). Digital voucher (2004; MVZ 250844).

  55. Nine km north of Varanda (town), Martuni Province, Nagorno-Karabakh (322 m). Digital voucher (2004; MVZ 250845).

  56. Seven km west of Martuni (town), Martuni Province, Nagorno-Karabakh (364 m) (pers. obs., 2003).

  57. Two km southwest of Amaras Monastery, Martuni Province, Nagorno-Karabakh (729 m) (pers. obs., 2006).

  58. Four km north of Martakert (town), Martakert Province, Nagorno-Karabakh (556 m) (pers. obs., 2006).

DISCUSSION

The geographic distributions of Armenian turtles are defined by elevation. The average altitude of Armenia is 1850 m, which is higher than any turtle in Armenia is known to occur. Mauremys caspica was recorded at elevations of up to 1800 m in Armenia; whereas, T. graeca is almost always found below 1500 m, and E. orbicularis is found even lower (below 1000 m). Because most of the country is uninhabitable for turtles, populations are restricted to the lowest portions of 2 distinct drainages: the Kura River drainage in the northeast and the Arax River valley in the south and west. Although disjunct in Armenia, these 3 main turtle areas are not allopatric, because they are connected outside of the territory studied here.

From a phylogeographic perspective, the 3 main turtle areas form a transect that is in need of further study. For example, the Arax populations of T. graeca have been referred to a new taxon Testudo graeca armeniaca Chkhikvadze and Bakradze, 1991. The populations in northeast Armenia are very close to the type locality of Testudo graeca ibera (Pallas 1814; near Tbilisi, Georgia, by subsequent designation of Bour 1987 in accordance with Eichwald 1831) and might be referred to this taxon. A recent genetic study (Parham et al. 2006) confirmed that tortoises from each of the type localities are in separate mitochondrial clades. Fritz et al. (2007) suggested that these clades could be considered subspecies; whereas, Parham et al. (2006) made no such designation. Part of the problem is that (as pointed out by Parham et al. 2006) tortoises from nearby the type locality of T. g. armeniaca (Meghri, by original designation) can exhibit dramatically different morphologies, despite being in the same mitochondrial clade. Clearly more fine-grained studies that integrate morphology with mitochondrial DNA and other genetic markers are needed, especially where different morphotypes and genetic clades come into contact. In this respect, the Nagorno-Karabakh populations of T. graeca are especially interesting because they occupy the area intervening the Kura and Arax drainages where 2 different taxa/clades occur. European pond turtles may exhibit similar genetic differentiation within Armenia. The Emys populations in northeast Armenia (in the Kura drainage) are referred to E. o. iberica Eichwald, 1831 (Fritz 1994); whereas, the populations of the Arax are taxonomically unassigned pending further study. The details of Arax drainage endemism in turtles is an important area for future study aimed at better defining taxonomic units with the Caucasus.

The historic data are not sufficient to track recent changes in abundance or distribution. However, we did record the apparent extirpation of T. graeca from the close vicinity of Yerevan (likely because of urbanization). 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 tracking additional extirpations, as well as for future studies aimed at the conservation status, ecology, or systematics of Caucasus turtles.

Acknowledgments

This research was made possible by a Linnaeus Fund Research Grant. We thank the Armenian biologists that shared their data on the distribution of Armenian turtles: Prof Felix Danielyan, Dr Aram Agasyan, Dr Samvel Pipoyan, Dr Eduard Yavruyan, Alexander Malkhasyan, and Andranik Virabyan. Ted Papenfuss (U.C. Berkeley) also provided invaluable assistance. Special thanks to Jarmo Perälä for kindly providing essential information on turtles from the Zoological Institute of St. Petersburg, Russia. We also thank Uwe Fritz for reviewing an earlier draft of the manuscript. This is University of California Museum of Paleontology contribution no. 1922.

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Copyright: 2008
Figure 1.
Figure 1.

Top: Map of Armenia and Nagorno-Karabakh Republic (Artsakh) and surrounding countries showing the 3 areas of relatively low elevation where turtles occur (A–C): A) Western Armenia, Arax River drainage; B) Southern Armenia and Nagorno-Karabakh, Arax River Drainage; C) Northern Armenia, Kura River drainage; Y) Yerevan. Bottom: Political map of Armenia and Nagorno-Karabakh Republic (Artsakh) showing localities mentioned in the text.

Figure 2.
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).

Figure 3.
Figure 3.

Distribution of Emys orbicularis in Armenia and Nagorno-Karabakh Republic (Artsakh).

Figure 4.
Figure 4.

Distribution of Mauremys caspica in Armenia and Nagorno-Karabakh Republic (Artsakh).

Figure 5.
Figure 5.

Distribution of Testudo graeca in Armenia and Nagorno-Karabakh Republic (Artsakh).

Figure 6.
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 Province, central Armenia (T. graeca locality no. 28, photo by MA).

Received: 03 Jul 2006
Accepted: 07 Apr 2008
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