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This article was downloaded by: [University of York] On: 15 March 2013, At: 03:32 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Zoology in the Middle East Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tzme20 Chromosomal forms of the Mole Rat, Nannospalax nehringi (Satunin, 1898), from the Van Lake Basin in Eastern Turkey Yüksel Coşkun a , Alaettin Kaya a & Gökhan Yürümez a a Dicle University, Science and Art Faculty, Biology Department, 21280–Diyarbakır, Turkey Version of record first published: 28 Feb 2013. To cite this article: Yüksel Coşkun , Alaettin Kaya & Gökhan Yürümez (2009): Chromosomal forms of the Mole Rat, Nannospalax nehringi (Satunin, 1898), from the Van Lake Basin in Eastern Turkey, Zoology in the Middle East, 48:1, 17-24 To link to this article: http://dx.doi.org/10.1080/09397140.2009.10638362 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms- and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages

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Page 1: Chromosomal forms of the Mole Rat,               Nannospalax nehringi               (Satunin, 1898), from the Van Lake Basin in Eastern Turkey

This article was downloaded by: [University of York]On: 15 March 2013, At: 03:32Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH,UK

Zoology in the Middle EastPublication details, including instructions for authorsand subscription information:http://www.tandfonline.com/loi/tzme20

Chromosomal forms of theMole Rat, Nannospalax nehringi(Satunin, 1898), from the VanLake Basin in Eastern TurkeyYüksel Coşkun a , Alaettin Kaya a & Gökhan Yürümez a

a Dicle University, Science and Art Faculty, BiologyDepartment, 21280–Diyarbakır, TurkeyVersion of record first published: 28 Feb 2013.

To cite this article: Yüksel Coşkun , Alaettin Kaya & Gökhan Yürümez (2009):Chromosomal forms of the Mole Rat, Nannospalax nehringi (Satunin, 1898), from theVan Lake Basin in Eastern Turkey, Zoology in the Middle East, 48:1, 17-24

To link to this article: http://dx.doi.org/10.1080/09397140.2009.10638362

PLEASE SCROLL DOWN FOR ARTICLE

Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions

This article may be used for research, teaching, and private study purposes.Any substantial or systematic reproduction, redistribution, reselling, loan,sub-licensing, systematic supply, or distribution in any form to anyone isexpressly forbidden.

The publisher does not give any warranty express or implied or make anyrepresentation that the contents will be complete or accurate or up todate. The accuracy of any instructions, formulae, and drug doses should beindependently verified with primary sources. The publisher shall not be liablefor any loss, actions, claims, proceedings, demand, or costs or damages

Page 2: Chromosomal forms of the Mole Rat,               Nannospalax nehringi               (Satunin, 1898), from the Van Lake Basin in Eastern Turkey

whatsoever or howsoever caused arising directly or indirectly in connectionwith or arising out of the use of this material.

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Page 3: Chromosomal forms of the Mole Rat,               Nannospalax nehringi               (Satunin, 1898), from the Van Lake Basin in Eastern Turkey

Chromosomal forms of the Mole Rat, Nannospalax nehringi (Satunin, 1898), from the Van Lake Basin in

Eastern Turkey

(Mammalia: Rodentia)

Yüksel Co�kun, Alaettin Kaya, Gökhan Yürümez

Abstract. Two different chromosomal forms of the Mole Rat, Nannospalax nehringi (Satunin, 1898), have been found in the Van Basin, Eastern Turkey: a population around the city of Van with 2n=48, NF=72 and NFa=68, and a population around the city of Bitlis with 2n=54, NF=74 and NFa=70. Both populations are confined to well-defined distribution areas. While both popu-lations are separated in the south by Mt Süphan, an extinct volcano over 4000 m high, there is no geographic barrier between both populations in the north, and there seems to be a secondary con-tact zone. The Van population constitutes a new chromosomal form of Mole Rat in Turkey, and could be regarded as a sibling species. However, no taxomomic decision is taken until more evi-dence on the evolution of these forms has become available.

Key words: Rodentia, Spalacidae, Nannospalax nehringi, karyotype, Turkey.

Introduction Mole rats of the genus Nannospalax have been widely studied in recent years to clarify spe-cies boundaries and phylogenetic relationships. Approximately 30 karyotypes of Nanno-spalax have been reported from Turkey so far. The diploid number (2n) of chromosomes ranges from 38 to 62 (see e.g. GÜLKAÇ & YÜKSEL 1989, IVANITSKAYA et al. 1997, TEZ et al. 2002), and the fundamental number of chromosomal arms (NF) varies between 66 and 92, while the fundamental number of autosomal arms (NFa) ranges from 62 to 88 (CO�KUN 2004a, b, c, SÖZEN et al. 2000, SÖZEN 2004, CO�KUN et al. 2006, and references therein). Unfortunately, karyological studies of the Nannospalax populations within the territory of Turkey are, on the whole, far from satisfactory for clarifying the distribution pattern and evolution of these chromosomal forms.

The range of Nannospalax nehringi (Satunin, 1898), first described as Spalax nehringi, is confined to the Caucasus and Eastern Turkey (TOPACHEVSKII 1969). It has been regarded as conspecific with Spalax leucodon Nordmann, 1840 (ELLERMAN & MORRISON-SCOTT 1951), but, following GROMOV & BARANOVA (1981) who differentiated Nannospalax from Spalax based on morphological characters, we consider this taxon to be a distinct species.

The first studies on the karyological peculiarities of N. nehringi were carried out by MAT-THEY (1959), who recorded 2n=48 in samples from the Caucasus. Later, NEVO et al. (1995) reported the diploid number of chromosomes of specimens from Erzurum and Kars (Sar�-kam��) as 2n=50. SÖZEN et al. (2000) found 2n=50 and NF=72 in the populations from Kars (Susuz), Erzurum and Ardahan, CO�KUN (2003) 2n=48, NF=68 and NFa=64 from A�r� and

Zoology in the Middle East 48, 2009: 17–24. ISSN 0939-7140 © Kasparek Verlag, Heidelberg

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18 Zoology in the Middle East 48, 2009

Van-Çald�ran, and again CO�KUN (2003) 2n=50, NF=70, NFa=66 from Erzurum and the Kars province.

We have studied the karyology of the Mole Rat in the Van Basin, which is relatively iso-lated and where different climate regions occur. We wanted to determine how the karyologi-cal characteristics have evolved in different, relatively isolated populations.

Material and methods The Mole rat specimens used in this study were collected in the Van Lake basin (Mu�, Bitlis and Van districts) in Eastern Turkey (37°58’–39°40’N; 41°18’–44°30’E). The region is characterised by high mountains (average altitude around 2000 m) and by large undulating plateaus at an alti-tude of 1500–2500 m. The high plateaus are subject to cold which limits arboreal vegetation, so that the region is covered mostly by steppe vegetation. Annual rainfall occurs more or less evenly throughout the year and varies between 400 and 600 mm. However, the rain shadow from large mountains, such as A�r�, has 200–300 mm rainfall, whilst the regions surrounding the large lakes may have 800–1000 mm rainfall.

A total of 21 Mole Rats (12 �, 9 �) was captured at 7 different localities in the Van Lake basin of Eastern Turkey (Fig. 1) during three surveys in 2006. We opened burrow systems and col-lected the animals with hoes when they came up to plug the opening. Voucher specimens were deposited at the Department of Biology, Science and Art Faculty, University of Dicle, Turkey.

Karyotyping was carried out according to standard procedures (see e.g. CO�KUN et al. 2006).

Results According to our karyotype analyses, two different populations of N. nehringi occur in the Van Lake basin in Eastern Turkey. The diagnostic features of the different karyotypes are described below and summarised in Table 1, which provides an overview of all the popula-tions of N. nehringi from Eastern Turkey that have been analysed so far. The approximate geographic area of each chromosomal form is shown in Fig. 1. Van population. The specimens of the Van population were collected near the type locality (Kars-Gaziler-Kaskoparan) of Nannospalax nehringi. The karyotypes of the two specimens from the city of east Mu� (Malazgirt town) and the four specimens from the city of Van (town of Erci� - Bozyaka village) have 2n=48, chromosomal arms NF=72, and autosomal arms NFa=68 (Table 1). Their karyotypes consist of 11 pairs of meta/submetacentric auto-somes and 12 pairs of acrocentric autosomes. The X chromosomes are large and submeta-centric, whereas the Y chromosomes are large and acrocentric (Fig. 2A). The karyotype of this population represents a cytogenetically distinct taxon, with a high percentage of biarmed chromosomes. Hence, the records from Mu� (Malazgirt town) and from Van (Erci�-Bozyaka village) have enlarged the distribution range of the 2n=48 form to the south and west. How-ever, whilst the karyotype of these populations has the same diploid chromosome number (i.e. 2n=48), the proportion of chromosomal and autosomal arms is variable. The karyotype of this population was first defined by CO�KUN (2003) as 2n = 48, NF = 68, NFa = 64, and the karyotypes observed here are different from those.

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Mammalia 19

Fig. 1. Sampling localities and geographic distribution of chromosomal forms of Nannospalax nehringi in eastern Turkey (locality names are given in the sequence province – district – village). Bitlis Population: (1) Bitlis - Hizan - Ta�top village (3 specimens); (2) Bitlis- Tatvan - A�a��kolba�� village (4 specimens); (3) Bitlis – Ahlat - Uludere village (4 specimens); (4) Mu� - Bulan�k - Erentepe village (3 specimens); (5) Mu� - Küm-bet village (1 specimen). – Van Population: (6) Van – Erci� - Bozyaka village (4 specimens); (7) Mu� - Malazgirt (2 specimens). Old records: (8) Bitlis – Tatvan – Kuskunk�ran; (9) Van – 1 km east of Çald�ran; (10) A�r� – Ta�l�çay – Yanalyol; (11) A�r� - Küpk�ran [data from localities 8–11 taken from CO�KUN (2003)]. Bitlis Population. The specimens of the Bitlis population were collected from five localities: three specimens at Ta�top village near Hizan, four specimens at A�a��kolba�� village near Tatvan, four specimens at Uludere village near Ahlat, three specimens at Erentepe village near Bulan�k, and one specimen at Kümbet village near Mu�.

The karyotypes of the 15 specimens from the Bitlis and Mu� (Erentepe and Kümbet vil-lage) populations have 2n=54, chromosomal arms NF=74, and autosomal arms NFa=70 (Table 1). The karyotypes consist of nine pairs of meta/submetacentric autosomes and 17 pairs of acrocentric autosomes. The X chromosomes are large and submetacentric, whereas the Y chromosomes are minute and acrocentric (Fig. 2B). This karyotype form is new to Turkey (Table 1).

This 2n=54 karyotype is different from the 2n=54 karyotypes described from Bingöl (NEVO et al. 1995), Yozgat (YÜKSEL & GÜLKAÇ 2001), Karabük province (SÖZEN 2004) and Kastamonu (SÖZEN et al. 2006b), but has the same chromosome arm numbers (NF and NFa) as the 2n=54 karyotypes of Nannospalax tuncelicus (Co�kun, 1996) from Tunceli, Elaz��, and Bingöl (CO�KUN 2004a) (Table 2).

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20 Zoology in the Middle East 48, 2009

Table 1. Chromosomal records of Nannospalax nehringi from Eastern Turkey. Abbreviations: sm = submetacentric; st = subtelocentric; a = acrocentric; m = metacentric; 2n = diploid chromosome number; NF = fundamental chromosome arm number; NFa = fundamental autosomal arm num-ber; X = X chromosome; Y = Y chromosome.

Autosomes 2n Biarmed:

m, sm, st a NFa NF X Y References

A�r� and Çald�ran 48 9 14 64 68 sm a CO�KUN 2003 Mu� (Malazgirt), Van (Erci�) 48 11 12 68 72 Sm a This study Erzurum, Kars (Sar�kam��) 50 9 15 66 70 sm – NEVO et al. 1995 Erzurum and Kars 50 9 15 66 70 sm a CO�KUN 2003 Erzurum, Kars (Susuz), Ardahan 50 10 14 68 72 sm a SÖZEN et al. 2000 Bingöl 54 - - - – – – NEVO et al. 1995 Tunceli, Elaz��, Bingöl 54 9 17 70 74 sm a CO�KUN 2004a Bitlis, Mu� 54 9 17 70 74 sm a This study

Table 2. Survey of the 2n=54 and 2n=48 chromosomal forms of Nannospalax described from Turkey (abbreviations as in Table 1).

Autosomes 2n Biarmed:

m, sm, st a NFa NF X Y References

Karabük 54 8 18 68 72 sm a SÖZEN 2004 Kastamonu 54 8 18 68 72 sm a SÖZEN et al. 2006b Tunceli, Bingöl, Elaz�� 54 9 17 70 74 sm a CO�KUN 2004a Bolu 54 7 19 66 70 sm - NEVO et al. 1995 Bingöl 54 - - - - sm - NEVO et al. 1995 Yozgat 54 9 17 70 74 sm st YÜKSEL & GÜLKAÇ 2001 Bitlis Mu� 54 9 17 70 74 sm a This study A�r�, Çald�ran 48 9 14 64 68 sm a CO�KUN 2003 Gümü�hane 48 10+1 12 67 71 Sm - SÖZEN et al. 2006a Mu�, Van 48 11 12 68 72 Sm a This study

Discussion The karyotype of the N. nehringi specimens collected from Malazgirt, Mu� province (local-ity 7) and Bozyaka village, Van provionce (locality 6), which are called the Van population, differs in the number of chromosomal arms from the specimens collected in A�r�, Çald�ran and Gümü�hane (CO�KUN 2003, SÖZEN et al. 2006a) (Table 1-2).

The 2n=54 chromosomal form shows a great variability in the number of chromosomal arms (NF and NFa) among the various populations (Table 2). 2n=54 karyotypes have been recorded from Bolu and Bingöl (NEVO et al. 1994, 1995), Yozgat (YÜKSEL & GÜLKAÇ 2001), Karabük province (SÖZEN 2004), Kastamonu (SÖZEN et al. 2006b) and Tokat (SÖZEN et al. 2006a). The chromosome morphology of the Bingöl population studied by NEVO et al. (1995) is actually unknown since the figure given for the 2n=54 karyotype is for the Bolu population. However, the karyotype of the populations from Bitlis (this study) is similar (2n=54, NF=74, NFa=70) to that of the population from Yozgat reported by YÜKSEL &

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Mammalia 21

Fig. 2. Standard karyotypes of Nannospalax nehringi: (A) Van population (�, No: 580), with 2n = 48; (B) Bitlis population (�, No: 592) with 2n = 54; (C) Nannospalax tuncelicus population (�, No: 397), with 2n = 54 (CO�KUN 2004a).

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22 Zoology in the Middle East 48, 2009

Fig. 3. Nannospalax nehringi, male (Photo: Y. CO�KUN, 26.9.2006, Bitlis- Hizan- Ta�top village, No. 576).

GÜLKAÇ (2001). However, these two areas are geographically very distant one from another. The forms from Yozgat and from Bitlis are therefore regarded as different (Tables 1-2).

The karyotype 2n=54, NF=74, NFa=70 of the Bitlis population of N. nehringi (this study) is also similar to that of the Nannospalax tuncelicus populations from Elaz��, Tunceli, and Bingöl (CO�KUN 2004a), but the chromosome morphology is different. The main difference is the presence of only one large biarmed and one large (first pair) acrocentric chromosome pair in N. nehringi, while N. tuncelicus has two pairs biarmed and no large acrocentric pair (Figs 2B-C).

According to these results, the Van and Bitlis populations represent two different chromo-somal forms of the species N. nehringi, even though there are no significant geographic barriers between these two populations (from the Malazgirt and Bulan�k districts). However, each karyotype form occupies a well-defined geographic range, and so some ecological barriers may limit its expansion. No hybrids have been found between these two populations.

NEVO et al. (1995) stated that each chromosomal form should be assigned to a separate biological species and that there are presumably about 30 such species in Turkey. While the chromosomal diversity in Turkish N. nehringi is rather great, it is difficult at this time to decide whether or not a particular karyotype represents a separate species.

ULUTÜRK & CO�KUN (2000) showed that the Bitlis population possesses morphological characters different from other populations. This could indicate that the Bitlis population is a separate subspecies of Nannospalax nehringi (Fig. 3). According to the karyotype analyses in this study, however, the Bitlis population of N. nehringi could even be considered a sepa-rate species.

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Mammalia 23

Acknowledgements. This study was supported by the Scientific and Technical Research Council of Turkey (TÜB�TAK Project No. 105T192). We are grateful to T. PAVLICEK (Haifa, Israel) for his comments on the manuscript.

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CO�KUN, Y. (2004a): A new species of mole rat, Nannospalax munzuri sp. n. and karyotype of Nanno-spalax tuncelicus (Co�kun, 1996) (Rodentia: Spalacidae) in eastern Anatolia. – Zoology in the Mid-dle East 33: 153-162.

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CO�KUN, Y. (2004c): A new chromosomal form of Nannospalax ehrenbergi from Turkey. – Folia Zoologica 53: 351-356.

CO�KUN, Y., S. ULUTÜRK & G. YÜRÜMEZ (2006): Chromosomal diversity in mole-rats of the species Nannospalax ehrenbergi (Rodentia: Spalacidae) from South Anatolia, Turkey. – Mammalian Biol-ogy – Zeitschrift für Säugetierkunde 71: 244-250.

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NEVO, E., M. G. FILIPPUCCI, C. REDI, A. KOROL & A. BEILES (1994): Chromosomal speciation and adaptive radiation of mole rats in Asia Minor correlated with increased ecological stress. – Proceed-ings of the National Academy of Sciences of the United States of America 91: 8160-8164.

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SÖZEN, M., E. ÇOLAK & N. YIIT (2000): Contributions to the karyology and taxonomy of Spalax leucodon nehringi Satunin, 1898, and Spalax leucodon armeniacus Mehely, 1909 (Mammalia: Ro-dentia) in Turkey. – Zeitschrift für Säugetierkunde 65: 309-312.

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ULUTÜRK, S. & Y. CO�KUN (2000): Morphological peculiarities of Mole Rats from Bitlis Province, Turkey. – Proceedings of the XVth National Congress of Biology, Ankara 270-276 [in Turkish with English summary].

YÜKSEL, E & M. D. GÜLKAÇ (2001): The cytogenetical comparison of Spalax (Rodentia: Spalacidae) populations from middle K�z�l�rmak basin, Turkey. – Turkish Journal of Biology 25: 17-24.

Authors’ address: Yüksel Co�kun, Alaettin Kaya and Gökhan Yürümez, Dicle University, Science and Art Faculty, Biology Department, 21280–Diyarbak�r, Turkey. – Email: [email protected].

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