Does Alexandromys Evoronensis Inhabit the Northeastern Part of Verkhnezeiskaya Plain? I
Total Page：16
File Type：pdf, Size：1020Kb
ISSN 1062-3590, Biology Bulletin, 2017, Vol. 44, No. 9, pp. 1049–1055. © Pleiades Publishing, Inc., 2017. Original Russian Text © I.N. Sheremetyeva, I.V. Kartavtseva, T.V. Vasiljeva, 2017, published in Zoologicheskii Zhurnal, 2017, Vol. 96, No. 4, pp. 477–484. Does Alexandromys evoronensis Inhabit the Northeastern Part of Verkhnezeiskaya Plain? I. N. Sheremetyeva*, I. V. Kartavtseva, and T. V. Vasiljeva Institute of Biology and Soil Science, Far East Branch, Russian Academy of Sciences, Vladivostok, 690022 Russia *e-mail: [email protected] Received October 16, 2015 Abstract⎯Gray voles of the “maximowiczii” group of the genus Alexandromys were found in the Argi River of the Amur region (northeastern part of Verkhnezeiskaya Plain). Analysis of mtDNA showed its similarity to A. evoronensis. The karyotype was 2n = 36, and NFa = 51–52. The karyotypes of these individuals did not contain the marker chromosome typical for Alexandromys maximowiczii. Keywords: Alexandromys, Verkhnezeiskaya Plain, inhabit, karyotypes, control region of mtDNA DOI: 10.1134/S1062359017090126 Alexandromys maximowiczii (Schrenck 1858) is (Verkhnebureinskaya Depression), which, according a wide-ranging species of voles of eastern Asia, which to the results of molecular genetic analysis, were close is characterized by a complex chromosomal polymor- to A. evoronensis (Sheremetyeva et al., 2016). It should phism (2n = 36–44, NFa = 50–60). Five chromo- be noted that the karyotype of the vole from the vicin- somal forms with geographical confinement have been ity of Chegdomyn (2n = 37, NF = 55) did not fit the distinguished. In the northern peripheral area in range of variability of the karyotype known for A. evo- opposite parts of the species range, two forms were ronensis (2n = 38–40, NF = 53–56) (Koval’skaya and 1 described in intermountain isolates. Later, as a result Sokolov, 1980; Golenishchev and Radzhabli, 1981; of the study of their karyotypes and hybridization Meyer et al., 1996; Kartavtseva et al., 2007) but was experiments, these forms acquired the species status: similar in the number and morphology of chromo- A. evoronensis (Kowalskaia et Sokolov 1980) (2n = 38– somes to the karyotype of A. maximowiczii from Khen- 40, NFa = 51–54) and A. mujanensis (Orlov et Kowal- tei in Mongolia (Orlov et al., 1978). The similarity in skaia 1978) (2n = 38, NFa = 46–50). A. evoronensis is the number and morphology of chromosomes (with- one of the sibling species of gray voles of the “maxi- out differential staining) for these species cannot be mowiczii” group, which was described 35 years ago regarded as evidence of their identity. However, near the source of the Devyatka River on the shore of Koval’skaya and Sokolov (1980) assumed that, Evoron Lake, Solnechnyi district, Khabarovsk krai according to the types of mutations known, the diploid only on the basis of the results of karyological analysis number in A. evoronensis theoretically may vary from (Koval’skaya and Sokolov, 1980). Until recently, this 36 to 42. However, extreme variants of chromosome species was reliably known only from the Evoron- numbers have not yet been found. These authors also Chukchagirskaya Lowland, Khabarovsk krai (Fig. 1, assumed that the distribution of A. evoronensis is not points 1–3): 1, in the vicinities of Evoron Lake (Gole- limited to the Evoron-Chukchagirskaya Lowland and nishchev and Radzhabli, 1981; Meyer et al., 1996; that this species may be found in some lacustrine basins Shenbrot and Krasnov, 2005); 2, in the environs of the in the eastern part of the Baikal–Amur Railroad. village of Kharpichan (51°19′ N, 136°35′ E) (Kartavt- Within the framework of the study of the variability seva et al., 2007; Sheremetyeva et al., 2010); and 3, in of marginal and isolated populations of species of gray the environs of the village named after Polina voles of the “maximowiczii” group, in July 2015 we Osipenko, Khabarovsk krai, near the confluence of made a second attempt to catch a representative of this Amgun and Nimelen rivers (Sheremetyeva et al., group (A. maximowiczii) in the northeastern part of the 2010). It was believed that the penetration of A. evo- Verkhnezeiskaya Plain. The first attempt was made ronensis to the west is hampered by the Bureya, Dusse- one year ago (Kartavtseva et al., 2015), when we Alin, and Yam-Alin ranges (Meyer et al., 1996). How- caught only a pair of root voles in the vicinity of the vil- ever, in July 2014, a pair of gray voles was caught in the lage of Verkhnezeysk. At this time, attempts to catch the Urgal River valley near the village of Chegdomyn (Fig. 1, animals were made in the Argi River valley (Fig. 1), point 4), Verkhnebureinskii district, Khabarovsk krai Amur region, near the place where, according to 1049 1050 SHEREMETYEVA et al. (a)(a) – A. evoronensis Yam-Alin Range Stanovoi Range – New find Tugur Уда Zeya Dusse-Alin Range Argi 3 Dzhagdy Range Nimelen Ezol Range 2 Amur Selemdzha Nora 1 Soktakhan RangeDep Amgun 4 Zeya Turana Range Bureinskii Range Bureya Amur China ((b)b) a ey ZeyaZ ЗеяЗ ея l i ne ArgiArg han a c 4 ZeyaZey channel 3 1 2 Fig. 1. (a) Sites of finds and (b) scheme of location of the biotopes in which the catches of A. evoronensis were performed. Sapaev (Sapaev, 1973; Sapaev and Voronov, 1976), completely covered with water. In total, 15 gray voles A. maximowiczii was previously caught. To set traps, were caught in 80 traps. Also we caught two gray red- we selected four biotopes: (1) the left bank of the Zeya backed voles, five wood mice, one pika, and one shrew Channel with a broadleaf forest site (54°40′17″ N, of the genus Sorex. The biological diversity in the catches 129°6′36″ E); (2) the left bank of the Zeya channel was maximum in biotope 1, and the number of gray voles overgrown with willows, at the confluence into the was maximum (eight individuals) in biotope 3. Argi River (54°40′6″ N, 129°6′39″ E); (3) an island on The species diagnostics of gray voles was performed the Argi River, overgrown willow and horsetail using karyological analysis and sequencing of the (54°40′31″ N, 129°5′49″ E); and (4) a high bank of the mtDNA control region. For ten individuals, species Argi River with a larch forest, overgrown with rosehip identification was performed in vivo, and the individ- (54°40′40″ N, 129°6′1″ E). It should be noted that, uals themselves were left for further breeding. Chro- during a heavy flood in 2013, biotopes 2 and 3 were mosome preparations were obtained in the field from BIOLOGY BULLETIN Vol. 44 No. 9 2017 DOES ALEXANDROMYS EVORONENSIS INHABIT 1051 (a)(a) XY ((b)b) XY Fig. 2. (a) Karyotype and (b) C-banding of chromosomes of the vole (male no. 3859) from the Argi River basin (Amur region). the femoral bone marrow cells (Grafodatskii and and the live specimens are kept in the vivarium of the Radzhabli, 1988). Staining for structural heterochro- Institute of Biology and Soil Science, Far East matin was performed according to Sumner (1972) Branch, Russian Academy of Sciences (Vladivostok). without pretreatment of preparations with hydrochlo- ric acid. G-bands on chromosomes were identified The analysis of the chromosomal preparations pre- using trypsin (Seabright, 1971). Chromosome images pared for one male (no. 3950) from the natural popu- were obtained with an AxioImager M1 microscope. lation and one female (no. 4017), which was obtained as a result of vivarium breeding, showed that the DNA was isolated by salt extraction (Aljanabi and karyotypes of these individuals contained 36 chromo- Martinez, 1997) from tissues fixed with alcohol. The somes (Fig. 2a). The number of autosomal arms of the procedure of the reaction mixture preparation, the male (no. 3950) and the female (no. 4017) was 51 and scheme of PCR reaction for obtaining the fragment, 52, respectively. Thus, this population is characterized and the preparation of samples for sequencing were by polymorphism, which requires a more detailed described previously (Sheremetyeva et al., 2015). The analysis. The Y chromosome is acrocentric and con- nucleotide sequence was determined with an ABI sists of heterochromatin only. The Y chromosome was Prizm 3130 automatic sequencer (Applied Biosystems, determined by the structural heterochromatin staining United States) on the basis of the Institute of Biology method (Fig. 2b). The X chromosome is a medium- and Soil Science, Far East Branch, Russian Academy size submetacentric (it was determined by G-banding) of Sciences (Vladivostok). The editing and alignment and has a slight C-block in the pericentromeric region. of the sequences obtained was performed using the Pericentromeric C-blocks were identified in three BioEdit 7.0.9.0 software (Hall, 1999). Constructing pairs of meta-submetacentric chromosomes (one pair phylogenetic trees and the calculation of genetic dis- of medium-size chromosomes and two pairs of tances performed using MEGA 6.0 program (Tamura smaller chromosomes) as well as in the first pair and et al., 2011). In constructing phylogenetic trees, the the two last pairs of acrocentric chromosomes. Other mtDNA control region sequences of seven species of acrocentric chromosome pairs may contain small eastern Asian voles (A. fortis, A. sachalinensis, A. gro- amounts of heterochromatin. In addition, the short movi, A. mujanensis, A. middendorffii, A. evoronensis, arms of one of the subtelocentric chromosomes con- and A. maximowiczii), obtained previously (Haring sist solely of heterochromatin. When comparing the et al., 2011; Sheremetyeva et al., 2015), as well as the newly described karyotype of the individuals from the mtDNA control region sequences of the voles that Argi River basin (2n = 36, NF = 54) (Verkhnezeiskaya were caught in July 2015 in the vicinity of Chegdomyn Plain) with the karyotype of the male (no.