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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,
Tavush Province. Darevsky (1957).
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,
Kashatagh Province, Nagorno-Karabakh (290 m).
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
ARAKELYAN AND PARHAM — Geographic Distribution of Turtles in Armenia and the Nagorno-Karabakh Republic 73
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).
74 CHELONIAN CONSERVATION AND BIOLOGY, Volume 7, Number 1 – 2008
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,
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. 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
m). Digital voucher (2004; MVZ 230840).
ARAKELYAN AND PARHAM — Geographic Distribution of Turtles in Armenia and the Nagorno-Karabakh Republic 75
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,
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, 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
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. 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
76 CHELONIAN CONSERVATION AND BIOLOGY, Volume 7, Number 1 – 2008
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
ARAKELYAN AND PARHAM — Geographic Distribution of Turtles in Armenia and the Nagorno-Karabakh Republic 77