the geographic distribution of turtles in armenia and the nagorno-karabakh republic (artsakh)

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The Geographic Distribution of Turtles in Armenia and the Nagorno-KarabakhRepublic (Artsakh)Author(s): Marine Arakelyan and James F. ParhamSource: Chelonian Conservation and Biology, 7(1):70-77. 2008.Published By: Chelonian Research FoundationDOI: http://dx.doi.org/10.2744/CCB-0664.1URL: http://www.bioone.org/doi/full/10.2744/CCB-0664.1

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Chelonian Conservation and Biology, 2008, 7(1): 70–77� 2008 Chelonian Research Foundation

The Geographic Distribution of Turtles in Armenia and the Nagorno-KarabakhRepublic (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 geographicdistribution in some Caucasian regions are poorly collated, hard to access, or entirely lacking. Wepresent a comprehensive review of turtle localities in such a region, Armenia and Nagorno-Karabakh Republic (Artsakh), based on literature records, personal observations, and museumspecimens. The compiled data for the 3 native species (Emys orbicularis, Mauremys caspica, andTestudo graeca) show that their distributions in this region are restricted to 3 areas of lowelevation. 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 areasvary, with E. orbicularis being the most restricted by elevation. We note that the 3 regions hereform part of an important transect from the interior Arax drainage (in the Ararat region ofwestern Armenia) to near its confluence with the Kura drainage (in southern Armenia andNagorno-Karabakh) to the Kura drainage sensu stricto (in northeast Armenia). Future studies ofturtles in these areas will play an important role in understanding the historical biogeography ofCaucasian turtles.

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

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.

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, Darev-

sky’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 nonvouch-

ered 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. 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).

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

ARAKELYAN AND PARHAM — Geographic Distribution of Turtles in Armenia and the Nagorno-Karabakh Republic 71

1. Airum railway station, Debed River, Tavush Province.

Darevsky (1957).

2. Above Uzuntale (Aygehovit) (village), Agshtev River,


3. Byurakan (village), Amberd River, Aragatsotn Prov-

ince. 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 Prov-

ince (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-Kar-

abakh (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,


Digital voucher (2004; MVZ 230836).

34. Mamark (village), Vorotan River, Kashatagh Prov-

ince, 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).

!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 AmarasMonastery, 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 abundantaquatic vegetation (E. orbicularis locality no. 4, photo by MA); G) Dorsal view Mauremys caspica from Ararat Valley (M. caspicalocality no. 23, photo by MA); H) Ventral view of same individuals in B; I) Subadult M. caspica from Aknakhbyur Village, HudrutProvince, Nagorno-Karabakh (M. caspica locality no. 36, photo by R. Sindaco); J–L) Typical habitats of M. caspica in Armenia andNagorno-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 TzobiVillage, Nagorno-Karabakh (M. caspica locality no. 30, photo by MA); M) Testudo graeca from vicinity of Martakert, MartakertProvince, Nagorno-Karabakh (T. graeca locality no. 58 photo by R. Sindaco); N) T. graeca from the vicinity of Meghri (T. graecalocality 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) Typicalhabitat of T. graeca in the vicinity of Meghri (T. graeca locality no. 43, photo by K. Milto); R) Typical habitat of T. graeca in theGorovan sands (T. graeca locality no. 33, photo by MA).

72 CHELONIAN CONSERVATION AND BIOLOGY, Volume 7, Number 1 – 2008

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).

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

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

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

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

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, KotayakProvince, central Armenia (T. graeca locality no. 28, photo byMA).


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. graecainhabits 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 spec-

imen (MZIA 504/1932). Chernov (1939).

2. Paravakar (village), Tavush Province. Museum spec-

imen (MZIA 505/1934). Chernov (1939) (A. Virab-

yan, pers. obs., 2004).

3. Between the towns of Kuibishev (Dilijan) and Idjevan,


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.


10. Vicinity of Nerkin Karmir (village), Tavush Province

(A. Virabyan, pers. obs., 2004).

11. Vicinity of Aigedzor (village), Tavush Province (A.


12. Vicinity of Chinari (village), Tavush Province (A.


13. Vicinity of Choratan (village), Tavush Province (A.


14. Vicinity of Mosesgegh (village), Tavush Province (A.


15. Vicinity of Tsachkavan (village), Tavush Province (A.


16. Vicinity of Varagavan (village), Tavush Province (A.


17. Vicinity of Airum (village), Tavush Province (A.


18. Vicinity of Koghb (village), Tavush Province (A.


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. Muse-

um 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), Aragat-

sotn 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



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, Na-

gorno-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,


Digital voucher (2004; MVZ 230841).

52. Seven km southeast of Tzobi/Sobi (village), Tzobajur

River valley, Basutajur, Kashatagh Province, Nagor-

no-Karabakh (357 m). Digital voucher (2004; MVZ

230842). (pers. obs., 2006).

53. Four km northwest of Chopadara (village), Tzobajur

River valley, Basutajur, Kashatagh Province, Nagor-

no-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


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. graecais almost always found below 1500 m, and E. orbicularisis 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 armeniacaChkhikvadze 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. graecain 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 Perala 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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Received: 3 July 2006

Revised and Accepted: 7 April 2008


the geographic distribution of turtles in armenia and the nagorno-karabakh republic (artsakh)

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