Äîêóìåíò âçÿò èç êýøà ïîèñêîâîé ìàøèíû. Àäðåñ îðèãèíàëüíîãî äîêóìåíòà : http://zmmu.msu.ru/rjt/articles/ther12_2%20083_090%20Lissovsky.pdf
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Ïîèñêîâûå ñëîâà: molecular cloud
Russian J. Theriol. 12(2): 8390

¿ RUSSIAN JOURNAL OF THERIOLOGY, 2013

Morphological and genetic variation of narrow-headed voles Lasiopodomys gregalis from South-East Transbaikalia
Andrey A. Lissovsky*, Ekaterina V. Obolenskaya & Tatyana V. Petrova
ABSTRACT. The sample of 115 skulls and 28 sequences of cytochrome b gene of narrow-headed voles Lasiopodomys gregalis, mainly from South-East Transbaikalia was studied. Geographic variation of craniometric features was not found in the studied sample. Such morphological variation contradicts deep mitochondrial divergence between voles from the South and North of the region under discussion. KEY WORDS: narrow-headed voles, Lasiopodomys gregalis, South-East Transbaikalia.
Andrey A. Lissovsky [andlis@zmmu.msu.ru] and Ekaterina V. Obolenskaya [obolenskaya@zmmu.msu.ru], Zoological Museum of Moscow State University, Bolshaya Nikitskaya 6, Moscow 125009, Russia; Tatyana V. Petrova [p.tashka@inbox.ru], Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, Saint Petersburg 199034, Russia.

Ìîðôîëîãè÷åñêàÿ è ãåíåòè÷åñêàÿ èçìåí÷èâîñòü óçêî÷åðåïíûõ ïîëåâîê Lasiopodomys gregalis Þãî-Âîñòî÷íîãî Çàáàéêàëüÿ
À.À. Ëèñîâñêèé, Å.Â. Îáîëåíñêàÿ, Ò.Â. Ïåòðîâà
ÐÅÇÞÌÅ. Èçó÷åíà âûáîðêà èç 115 ÷åðåïîâ è 28 ïîñëåäîâàòåëüíîñòåé ãåíà öèòîõðîìà b óçêî÷åðåïíîé ïîëåâêè Lasiopodomys gregalis, ãëàâíûì îáðàçîì, èç Þãî-Âîñòî÷íîãî Çàáàéêàëüÿ. Ïîêàçàíî, ÷òî ãåîãðàôè÷åñêàÿ èçìåí÷èâîñòü êðàíèîìåòðè÷åñêèõ ïðèçíàêîâ â èçó÷åííîé âûáîðêå íå âûðàæåíà. Ýòî íå ñîîòâåòñòâóåò ãëóáîêîé ìèòîõîíäðèàëüíîé äèâåðãåíöèè ïîëåâîê þãà è ñåâåðà ðåãèîíà. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: óçêî÷åðåïíûå ïîëåâêè, Lasiopodomys gregalis, Þãî-Âîñòî÷íîå Çàáàéêàëüå.

Introduction
Morphological variation of narrow-headed vole Lasiopodomys gregalis (Pallas, 1779) was studied in detail (Dupal, 2000; Golenishchev & Petrovskaya, 2002; Dupal & Abramov, 2010). According to these studies, morphologically homogeneous population inhabits the major part of the distribution range of the species. Some studies found clinal variation of craniometrical features. Several geographical samples were found to be slightly different from the major part of the species; these samples were different depending on the features and geographic area analyzed. Nobody found, however, morphological peculiarity of narrow-headed voles from South-East Transbaikalia. Meanwhile pilot genetic study found considerable mitochondrial difference between voles from SouthEast Transbaikalia and other distribution range (Abramson et al., 2006). The aim of this study was to examine mitochondrial and morphological variation of narrow-headed voles from South-East Transbaikalia and to evaluate concordance between mitochondrial and morphological data.

Materials and methods
Narrow-headed vole is a species that was formerly included in the genus Microtus Schrank, 1798 within the separate subgenus Stenocranius Kastschenko, 1901 (Gromov & Polyakov, 1977). Molecular data suggest that this taxon is close to species of the g enus Lasiopodomys Lataste, 1887. Furthermore, this latter genus is separate from the Microtus clade (Abramson et al., 2009; Bannikova et al., 2010). Thus, following Abramson & Lissovsky (2012), we use the name Lasiopodomys gregalis. Specimens used in the morphometric study were taken from the collection of the Zoological Museum of Moscow State University. The sample contained 115 intact skulls of L. gregalis (Appendix 1). Fifteen measurements were taken (with an accuracy of 0.1 mm) from each skull as follows: condylobasal length, minimal distance between maxillary toothrows, diastemal length, alveolar length of maxillary toothrow, zygomatic breadth, maximal width between lateral edges of auditory bullae, skull height at maxillary toothrows level, orbital length and width, width of rostrum base, orbital constriction, length of auditory bul-


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Figure 1. Maximum parsimony network of L. gregalis cytochrome b haplotypes from Transbaikalia. Haplotype labels refer to Appendix 2. The size of circles is proportional to the haplotype frequency.

la, width of auditory bulla, alveolar length of mandibular toothrow, distance between the base of incisor and the apex of mandibular articular process. All calculations were performed on logarithmic measurements. We used the skulls of voles of different ages. All skulls were divided into three age categories corresponding to obviously juvenile (1), obviously adult with developed crests on the skull (3), and all others (2). In order to exclude age bias from morphometric analysis, we used an orthogonal projection of initial data along the vector of age variation (Burnaby, 1966). The vector of age variation was calculated as the first eigenvector of the between-group covariance matrix computed with MANOVA, in which the variable containing age gradations was used as a grouping variable. We used only the first and the third age classes during calculations of the covariance matrix in order to avoid an error induced by inaccuracy in determining the second age class. There were five samples included in this analysis. The samples for hierarchical cluster analysis included only specimens collected in the same locality (45 samples total, Appendix 1). Only samples with n>3 were used in cluster analysis (16 samples). Cluster analysis was performed on the basis of a matrix of Mahalanobis distances using the unweighted pair group method with arithmetic mean. The bias induced by using samples of different sizes was corrected (Marcus, 1993). The following approach was applied as an ordination method: first, the eigenvectors of the within-group

covariance matrix (with 16 large geographic samples as groups) of the dataset with reduced age was calculated. Secondly, the initial data matrix was multiplied with the matrix of the eigenvectors. Thus the initial data was rotated into the space of intergroup variation without distortion of the initial space. Craniometric data were processed using standard modules of STATISTICA 8.0 (StatSoft 2007) and several custom algorithms written by the first author using Statistica Visual Basic programming language. A fragment of mitochondrial gene cytochrome b from 28 voles from 11 geographical localities (Appendix 2) was used for the study of genetic variation. The gene region (891 bp) was amplified by polymerase chain reaction (PCR) using primer combination L14728 (Lebedev et al., 2007) / H15985 (Ohdachi et al., 2001). PCR entailed 30 thermal cycles as follows: 30 s denaturation at 94œC, 45 s annealing at 55œC and 1 min 20 s extension at 72œC. All PCR experiments included negative controls. PCR products were visualized on 1.5% agarose gel and then purified using Omnix DNA purification kit. Both strands were directly sequenced using the ABI PRISM BigDye Terminator v. 3.1. A maximum parsimony haplotype network was constructed with Network 4.6 (Bandelt et al., 1999) using the median joining method. Field material was gathered in Zabaikalskiy Kray (former Chitinskaya Region and Aginskiy Buryat Autonomous District) within Krasnokamenskiy, Kalgan-


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Figure 2. Distribution of L. gregalis in South-East Transbaikalia. Localities where specimens were studied genetically are shown.

skiy, Aginskiy, Alexandrozavodskiy and Sretenskiy districts (2005, 2007, 2011), Russia. The map of distribution of narrow-headed voles in South-East Transbaikalia was constructed on the basis of localities of the specimens from the following collections: Zoological Museum of Moscow State University; Zoological Institute of the Russian Academy of Sciences, Saint-Petersburg; Institute of Systematics and Ecology of Animals, Siberian Branch of the Russian Academy of Sciences; Zoological Museum of Irkutsk State University.

of total variance respectively. Specimens from different parts of the range form the single cloud within this space (Fig. 3). Distribution of specimens from two regions of Transbaikalia, outlined above, completely overlaps. Cluster analysis of craniometric features does not separate voles from these two regions of Transbaikalia also (Fig. 4).

Discussion
Our results do not show concordance between the deep mitochondrial divergence of Transbaikalian narrow-headed voles and their cranial morphology. While the extent of mitochondrial differences between haplogroups A and B is comparable to the divergence between subgenera Alexandromys and Pallasiinus (Bannikova et al., 2010), craniometric data does not display any geographic variation. This discordance can be explained in two different ways. The first way means that voles from northern and southern parts of South-East Transbaikalia (corresponding to haplogroups A and B) belong to two sister taxa separated long time ago from each other. In this case, these taxa are sibling species, since they have no morphological distinction. Another possible explanation is mitochondrial artifact conservation of ancient mito-

Results
All haplotypes of Transbaikalian narrow-headed voles constituted two groups, divided by 98 substitutions (Fig. 1). Such difference roughly corresponds to p-distance of 11%. Geographic distribution of the two haplogroups representatives is shown in Fig. 2. Representatives of the haplogroup A distributed to the South from Shilka River and in steppe along Onon River. Voles bearing haplotypes of the haplogroup B inhabit western Transbaikalia and steppe to the North of Shilka and Ingoda Rivers. The first two axes of the space of maximized intersample morphological differences explain 41 and 28%


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Figure 3. Distribution of L. gregalis specimens in the space of maximized differences between geographical samples. The specimens, studied genetically, are outlined with circle.

Figure 4. Dendrogram for hierarchical cluster analysis of craniometric features of L. gregalis samples. Sample labels refer to Appendix 1. Samples from the territory of South-East Transbaikalia lying to the south of Ingoda and Shilka Rivers are marked by asterisk; to the north, by the number sign.


Narrow-headed voles of South-East Transbaikalia chondrial lineage within South-East Transbaikalia. It is possible for example as a result of preferred exchange of males between southern and northern populations together with limited exchange of females. Selection of one of these two hypotheses will be possible after analysis of nuclear genes. As was hypothesized by Obolenskaya (2010), Shilka and Onon Rivers can be a barrier to narrow-headed voles dispersion, similarly to Northern Ochotona hyperborea and Manchurian O. mantchurica pikas (Lissovsky et al., 2008) and striped hamsters Cricetulus b. barabensis and C. b. pseudogriseus (Lebedev & Lissovsky, 2008). Later, the importance of Shilka as a strong zoogeographical barrier was questioned after a detailed study of striped hamsters distribution (Korablyov et al., 2010). The latter authors found Unda River, instead of Shilka, as a border between Cricetulus b. barabensis and C. b. pseudogriseus distribution. Our data on distribution of narrow-headed voles from two haplogroups, presented herein, allow hypothesizing Shilka River as possible barrier of voles dispersion. However since both banks of Onon River are inhabited by voles with A haplotypes, this river does not play a zoogeographical role for the species under discussion. Probably, forests along the right bank of Ingoda River limit dispersion of voles in this region. ACKNOWLEDGEMENTS. We thank Pilnikov A.E., Popov V.P., Agapov V.A., Gredyushko A.N., Kirilyuk V.I. for the help with organizing of our field work; Kruskop S.V., Dudov S.V., Lisovskaya V.A., Vlasak A. for the help in the field. Kartavtseva I.V., Sheremetyeva I.N., Voyta L.L., and Lebedev V.S. donated tissues for genetic analysis. Abramson N.I. gave the opportunity of molecular analysis. The work was partly financially supported by RFBR (## 07-04-10059; 11-04-10066; 12-04-01310) and RAS Presidium Program of Fundamental Researches The living nature: contemporary conditions and problems of development.

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References
Abramson N.I., Kostygov A.Yu. & Gambaryan N.G. 2006. Phylogeography of narrow-headed vole (Microtus gregalis, Cricetidae, Rodentia) inferred from the variation of mitochondrial cyt b and a number of nuclear genes // Hystrix, Italian Journal of Mammalogy (n.s.) supp. P.155 156. Abramson N.I., Lebedev V.S., Tesakov A.S. & Bannikova A.A. 2009. [Supraspecies relationships in the subfamily Arvicolinae (Rodentia, Cricetidae): An unexpected result of nuclear gene analysis] // Molekulyarnaya Biologiya. Vol.43. No.5. P.834846 [in Russian]. Abramson N.I. & Lissovsky A.A. 2012. Subfamily Arvicolinae // Pavlinov I.Ya. & Lissovsky A.A. (eds). The Mammals of Russia: A taxonomic and Geographic Reference (Archive of the Zoological Museum of MSU. Vol. 52). Moscow: KMK Scientific Press. P.220276. Bandelt H.J., Forster P. & Röhl A. 1999. Median-joining networks for inferring intraspecific phylogenies // Molecular Biology and Evolution. Vol.16. P.3748.

Bannikova A.A., Lebedev V.S., Lissovsky A.A., Matrosova V., Abramson N.I., Obolenskaya E.V. & Tesakov A.S. 2010. Molecular phylogeny and evolution of the Asian lineage of vole genus Microtus (Arvicolinae, Rodentia) inferred from mitochondrial cytb sequence // Biological Journal of the Linnean Society. Vol.99. P.595613. Burnaby T.P. 1966. Growth-invariant discriminant functions and generalized distances // Biometrics. Vol.22. P.96 110. Dupal T.A. 2000. [Geographical variation and subspecies systematics of narrow-headed vole Microtus (Stenocranius) gregalis (Rodentia, Cricetidae)] // Zoologicheskii Zhurnal. Vol.79. No.7. P.851858 [in Russian, with English summary]. Dupal T.A. & Abramov S.A. 2010. [Intrapopulational morphological variation of narrow-headed vole (Microtus gregalis, Rodentia, Arvicolidae)] // Zoologicheskii Zhurnal. Vol.89. No.7. P.850861 [in Russian]. Golenishchev F.N. & Petrovskaya N.A. 2002. [Geographic variation of narrow-headed vole Microtus (Stenocranius) gregalis Pall., 1779] // Theriologicheskie Issledovania. Vol.1. P.1734 [in Russian]. Gromov I.M. & Polyakov I.Ya. 1977. [Fauna of the USSR, Voles (Microtinae): Mammals]. Leningrad: Izdatelstvo Nauka. Vol.3. No.8. 504 p. [in Russian]. Korablyov V.P., Pavlenko M.V., Kiriliuk V.I. & Bazhenov Yu.A. 2010. [Distribution of different chromosomal forms of the striped-back hamsters of the superspecies Cricetulus barabensis in Transbaikalia] // Prirodoohrannoye Sotrudnichestvo v Globalnykh Ekologicheskikh Regionakh: Rossiya-Kitai-Mongoliya: Sbornik Nauchnyh Materialov. No.1. Chita: Express-Izdatelstvo. P.131133 [in Russian]. Lebedev V.S., Bannikova A.A., Tesakov A.S. & Abramson N.I. 2007. Molecular phylogeny of the genus Alticola (Cricetidae, Rodentia) as inferred from the sequence of the cytochrome b gene // Zoologica Scripta. Vol.36. No.6. P.547563. Lebedev V.S. & Lissovsky A.A. 2008. [The geographical variability of cranial characteristics and systematics of striped-back hamsters (Cricetulus barabensis Cricetinae, Cricetidae, Rodentia)] // Zoologicheskii Zhurnal. Vol.87. No.3. P.361374 [in Russian]. Lissovsky A.A., Yang Q. & Pilnikov A. 2008. Taxonomy and distribution of the pikas (Ochotona, Lagomorpha) of alpina-hyperborea group in North-East China and adjacent territories // Russian Journal of Theriology. Vol.7. No.1. P.516. Marcus L.F. 1993. Some aspects of multivariate statistics for morphometrics // Marcus L.F., Bello E. & García-Valdecasas A. (eds.). Contributions to Morphometrics. Madrid: C.S.I.C. P.95-130. Obolenskaya E.V. 2010. [Zoogeographical peculiarities of South-East Transbaikalia (by example of small mammals)] // Vestnik MGU. Seriya Geographiya. No.5. P.60 66 [in Russian]. Ohdachi S., Dokuchaev N. E., Hasegawa M. & Masuda R. 2001. Intraspecific phylogeny and geographical variation of six species of northeastern Asiatic Sorex shrews based on the mitochondrial cytochrome b sequences // Molecular Ecology. Vol.10. P.21992213.


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Appendix 1. The list of specimens used in morphometrical analysis. Information is in the following order: the name of the sample in quotes (only for the samples, used in cluster analysis): locality and geographical coordinates: the specimen museum ID-s.
'Kyker': Russia, Zabaikalskiy Kray, Tungokochenskiy District, Kyker Village; 53.16œ N, 115.81œ E: S-23258, S-23259, S-23260, S-23261, S-23262, S-23264, S-23265, S-23266, S-23256. 'Matakan': Russia, Zabaikalskiy Kray, Sretenskiy District, Matakan River; 52.26œ N, 117.64œ E: S-182034, S182033, S-182031. 'Zyulzikan': Russia, Zabaikalskiy Kray, Nerchinskiy District, Zyulzikan Village; 52.67œ N, 116.28œ E: S23315, S-23254, S-23253. '': Russia, Zabaikalskiy Kray, Nerchinskiy District, vicinities of Nerchinsk; 51.95œ N, 116.56œ E: S-25672. '': Russia, Zabaikalskiy Kray, Nerchinskiy District, vicinities of Chita; 52.04 N, 113.53 E: S-25438. '': Russia, Zabaikalskiy Kray, Nerchinskiy District, Kalinino Village; 52.02œ N, 116.79œ E: S-25667. 'BorunSheviya': Russia, Zabaikalskiy Kray, Aginskiy District, valley of Borun-Sheviya; 51.43œ N, 114.95œ E: S-90776, S-90777, S-90773. '': Russia, Zabaikalskiy Kray, Borzinskiy District, 42 km from Borzya to Solovevsk; 50.1œ N, 116.14œ E: S-90758, S-90757, S-90765, S-90761. 'DalaiNor': China, Manchuria Province, Dalai-Nor Lake, mouth of Urshun River; 48.96œ N, 117.73œ E: S90754, S-90755. 'Kozlovo': Russia, Zabaikalskiy Kray, Kalganskiy District, Kozlovo Village; 51.21œ N, 118.93œ E: S-178592, S-178593, S-178594, S-178595, S-178596, S-178590, S-178591. 'Kuytun': Russia, Zabaikalskiy Kray, Krasnokamenskiy District, 7 km S from Kuytun Village; 50.12œ N, 118.67œ E: S-180440, S-180441, S-180443, S-180442, S-180444, S-180445, S-180446, S-180447. 'ZunAraltuy': Russia, Zabaikalskiy Kray, Borzinskiy District, Zun-Araltuy Lake; 50.05œ N, 117.37œ E: S90747, S-42816, S-42813, S-42810, S-42800, S-42806. '': Russia, Zabaikalskiy Kray, Aleksandrovozavodskiy District, valley of Gazimur River; 50.81œ N, 117.71œ E: S-90779. '': Russia, Zabaikalskiy Kray, Aginskiy District, Tsagan-Nor Lake; 50.7œ N, 115.22œ E: S-90774. '': Russia, Zabaikalskiy Kray, Borzinskiy District, Tsagan-Oluy Village; 50.46œ N, 117.15œ E: S-90762, S90763. '': Russia, Zabaikalskiy Kray, Byrkinskiy District, valley of Urulunguy River; 50.12œ N, 116.1œ E: S-90748. '': Zabaikalskiy Kray, Borzinskiy District, Nikolay-Torom Lake; 49.86œ N, 117.7œ E: S-90768. '': Russia, Zabaikalskiy Kray, Borzinskiy District, Chumrostuy place, 12 km from Kharanor station; 50.212œ N, 116.8œ E: S-90760, S-90767. '': Zabaikalskiy Kray, Borzinskiy District, Macievskaya station, valley of Bugutur; 49.68œ N, 117.47œ E: S45297, S-45298. 'Borzya': Russia, Zabaikalskiy Kray, Borzinskiy District, Borzya railway station; 50.38œ N, 116.51œ E: S25454. '': Russia, Zabaikalskiy Kray, Ononskiy District, vicinities of Kulusutay; 50.23œ N, 115.67œ E: S-25465. '': Russia, Zabaikalskiy Kray, Zabaikalskiy District, vicinities of Matsievskaya station; 49.72œ N, 117.25œ E: S-182077. '': Russia, Zabaikalskiy Kray, Chitinskiy District, Tasey Lake, vicinities of source Kholoya River; 52.3œ N, 113.14œ E: S-37834. 'Schuchya': Russia, Yamalo-Nenetskiy Autonomous Area, Yamal Peninsula, middle course of Schuchya River; 67.37œ N, 68.71œ E: S-111820, S-111822, S-111805, S-111826, S-111825, S-111814, S-32106, S-32138. '': Russia, Yamalo-Nenetskiy Autonomous Area, Yamal peninsula, Schuchya River, mouth of Kheyakhe River; 67.07œ N, 68.2œ E: S-96937. '': Russia, Yamalo-Nenetskiy Autonomous Area, Yamal peninsula, Seyakha River; 69.921œ N, 71.785œ E: S-100705. 'Irkutsk': Russia, Irkutsk region, Irkutskiy District, vicinities of Granovshina Village; 52.47œ N, 104.28œ E: S25444, S-25445, S-25446. '': Russia, Irkutsk region, Alarskiy District, Alar Village; 53.07œ N, 102.54œ E: S-140559, S-140560. 'Naushki': Russia, Buryatia Republic, Kyakhtinskiy District, vicinities of Naushki Village; 50.38œ N, 106.27œ E: S-179179, S-179182, S-179177. '': Mongolia, Khentii Aymak, left bank of Onon river; 48.93œ N, 110.04œ E: S-107110. '': Mongolia, Khurkha River, tributary of Onon; 48.58œ N, 110.61œ E: S-39942, S-39858. 'BuirNor': Mongolia, Dornod Aymak, east bank of Buir-Nuur Lake; 47.89œ N, 117.87œ E: S-119843, S39850, S-39854.


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'': Mongolia, Khentii Aymak, Khentey Range, Eren-Daba-Nuruu pass; 48.83œ N, 111.67œ E: S-137828. '': Mongolia, Khentii Aymak, Kerulen River, vicinities of Idermeg; 47.55œ N, 111.22œ E: S-128156. 'MungenMort': Mongolia, Khentey Range, vicinities of Mungen-Mort; 47.84 N, 107.85 E: S-110443, S110444, S-110445. 'Munhu': Mongolia, Sukhe-Batorskiy Aymak, Munkhu-Khany Somon; 46.95 N, 112.02 E: S-39941, S39946. 'Shamar': Mongolia, Selenge Aymak, vicinities of Shamar; 50.1œ N, 106.2œ E: S-149509, S-149510, S149511, S-149512, S-149513, S-149515, S-149516, S-149517, S-149518, S-149519, S-127966, S-127965, S137829. '': Mongolia, Sukhe-Batorskiy Aymak, 20 km E from Politin-Khuduk frontier post, Shiliyn-Bogd-Ula; 45.47œ N, 114.57œ E: S-137830, S-137831. '': Mongolia, Sukhe-Batorskiy Aymak, 80 km NE from Erden-Tsagan; 46.4œ N, 115.9œ E: S-111950. '': Mongolia, Dornod Aymak, frontier post Tereg, S from Khermiyn-Ula; 46.86œ N, 117.67œ E: S-137827. '': Mongolia, Kerulen River, 13 km E from Chaybalsan; 48.1œ N, 114.66œ E: S-41866. '': Mongolia, Urgyn-Gol River 12 km SW from Yugodzer; 45.87œ N, 115.22œ E: S-39844. '': Mongolia, Central of Khangay, upper Chulutuin River; 47.66œ N, 97.68œ E: S-39846. '': Mongolia, Dornod Aymak, 80 km SE Bayan Tumen; 47.35œ N, 114.62œ E: S-39855. '': Mongolia, Khentii Aymak, Bain-Gol River, tributary of Khurkha River; 48.43œ N, 110.37œ E: S-39862.


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A.A. Lissovsky, E.V. Obolenskaya & T.V. Petrova

Appendix 2. The list of specimens used in genetic analysis.
Locality Buryatia Republic, Kabanskiy District, Istomino Village Buryatia Republic, Kabanskiy District, Istomino Village Buryatia Republic, Kabanskiy District, Istomino Village Buryatia Republic, Kabanskiy District, Istomino Village Buryatia Republic, Kyakhtinskiy District, vicinities of Naushki Village, 50,367œ N, 106,25œ E Buryatia Republic, Kyakhtinskiy District, vicinities of Naushki Village, 50,367œ N, 106,25œ E Buryatia Republic, Ulan-Udinskiy District, Oshurkovo vVillage, 51,933œ N, 107,433œ E Zabaikalskiy Kray, Sretenskiy District, Matakan River, 52,383œ N, 117,9œ E Zabaikalskiy Kray, Sretenskiy District, Matakan River, 52,383œ N, 117,9œ E Zabaikalskiy Kray, Chitinskiy District, vicinities of Ugdan Village Zabaikalskiy Kray, Aginskiy Buryatskiy District, Kharganashi River, 50,669œ N, 114,667œ E Zabaikalskiy Kray, Aginskiy Buryatskiy District, Kharganashi River, 50,669œ N, 114,667œ E Zabaikalskiy Kray, Aginskiy Buryatskiy District, Kharganashi River, 50,669œ N, 114,667œ E Zabaikalskiy Kray, Aginskiy Buryatskiy District, Kharganashi River, 50,669œ N, 114,667œ E Zabaikalskiy Kray, Aleksandrovo-Zavodskiy District, Aleksandrovskiy Zavod, 55,861œ N, 117,931œ E Zabaikalskiy Kray, Baleyskiy District, vicinities of Baley Village Zabaikalskiy Kray, Baleyskiy District, vicinities of Baley Village Zabaikalskiy Kray, Kalganskiy District, vicinities of Kozlovo Village 51,35œ N, 118,217œ E Zabaikalskiy Kray, Kalganskiy District, vicinities of Kozlovo Village 51,35œ N, 118,217œ E Zabaikalskiy Kray, Kalganskiy District, vicinities of Kozlovo village 51,35œ N, 118,217œ E Zabaikalskiy Kray, Kalganskiy District, vicinities of Kozlovo Village 51,35œ N, 118,217œ E Zabaikalskiy Kray, Kalganskiy District, vicinities of Kozlovo Village 51,35œ N, 118,217œ E Zabaikalskiy Kray, Krasnokamenskiy District, vicinities of Kuytun Village, 50,2œ N, 118,083œ E Zabaikalskiy Kray, Krasnokamenskiy District, vicinities of Kuytun Village, 50,2œ N, 118,083œ E Zabaikalskiy Kray, Krasnokamenskiy District, vicinities of Kuytun Village, 50,2œ N, 118,083œ E Zabaikalskiy Kray, Krasnokamenskiy District, vicinities of Kuytun Village, 50,2œ N, 118,083œ E Zabaikalskiy Kray, Krasnokamenskiy District, vicinities of Kuytun Village, 50,2œ N, 118,083œ E Zabaikalskiy Kray, Ononskiy District, vicinities of Ust'-Borzya Village, 50,0œ N, 115,0œ E Haplotype h1 h1 h2 h3 h4 h5 h6 h7 h8 h9 h10 h11 h12 h10 h13 h14 h14 h14 h14 h14 h14 h14 h14 h14 h14 h14 h15 h16 Museum ID BPI-2029 BPI-2032 BPI-2033 BPI-2035 S-179177 S-179178 BPI-1989 S-182032 S-182034 coll_8 S-191190 S-191191 S-191192 S-191193 S-191194 ZIN-96277 ZIN-96283 S-178590 S-178593 S-178591 S-178592 S-178594 S-180440 S-180441 S-180442 S-180443 S-180446 S-180439 GenBank ID KF751101 KF751102 KF751103 KF751104 KF751098 KF751099 KF751100 KF751078 KF751079 KF751077 KF751088 KF751089 KF751090 KF751091 KF751087 KF751085 KF751086 KF751096 KF751092 KF751093 KF751094 KF751095 KF751080 KF751081 KF751082 KF751083 KF751084 KF751097