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The 'Common Vole' in Svalbard Identified As Microtus Epiroticus by Chromosome Analysis

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The 'Common Vole' in Svalbard Identified As Microtus Epiroticus by Chromosome Analysis The 'common vole' in Svalbard identified as Microtus epiroticus by chromosome analysis KARL FREDGA, MAARIT JAAROLA. ROLF ANKER IMS, HARALD STEEN AND NlGEL G. YOCCOZ Fredga, K., Jaarola, M., Ims, R. A,, Steen, FI. & Yoccoz. N. G. 1990: The 'common vole' in Svalbard identified as Microtus epiruticur by chromosome analysis. Polar Research 8, 283-2Y0. The chromosomes were studied in six individuals from a population of Microtus from Grumantbyen, Svalbard, and in six Microtur arualis (Pallas 1778) from Lauwersee, Holland. It was shown that the voles ' from Svalbard did not belong, as earlier supposed, to the species M. arualis (2n = 46) but to M. epiroticus (Ondrias, 1966) (2n = 54). We suggest that the Svalbard voles were introduced by man between 1920 and 1960 together with hay on Russian ships from the vicinity of Leningrad, USSR. Karl Fredga and Maarit Jaarola, Department of Genetics, Uppsala University, P.O. Box 7003. S-750 07 Uppsala. Sweden: Rolf Anker Ims and Harald Steen, Department of Biology, Division of Zoology, University of Oslo, P.O. Box 1050 Blindern, N-0316 Oslo 3, Norway; Nigel G. Yoccoz, Deparrment of Biology, Division of Zoology, Uniuersify of Oslo. P.O. Box I050 Blindern, N-0316 Oslo 3, Norway. or: Laboratoire de Biometrie, Unioersitk Claude Bernard Lyon 1. 69622 Villeurhanne Cedex. France; February 1990 (revised July 1990). In 1960 a member of the Finnish zoological the name was changed to M. epiroticus Ondrias expedition to Svalbard found the first vole in 1966 (see Honacki et al. 1982).* It is a sibling Svalbard at Longyearbyen. In 1964 and 1965 voles were abundant in the Longyearbyen village and district and several specimens were collected *Recently. the namc Microtus rossiaemeridionrrlis Ognev 1924 (Nyholm 1966). In 1975 voles were trapped at was applied to this species (Malygin 1983: Malygin & Yatscnko Hotellneset, Longyearbyen, and in 1976 at 1986). Fifteen spccimens of the 'common volc' were collcctcd in the type locality (Aninsk region. Voronej district) of the Fuglefjellet, Grumantbyen, W Adventfjorden, taxon M. arualis rossiaemeridionalis Ognev 1924. All fiftccn by Alendal (1977). The voles collected in the had 2n = 54 and since, according to Malygin & Yatscnko 1960s and 1970s were identified as Microtus (1986). M. arualis (2n = 46) docs not cxist in this area they aruulis (Pallas 1778) on morphological grounds. conclude that thc animals described by Ogiicv also must have had 54 chromosomes. Consequcntly, the sibling spccies with A more recent report describes voles collected at 54 chromosomes should be called M. russiaemeridiorralls Ognev Coles Bay and Barentsburg as M. arualis 1924 and not M. epiroticus Ondrias 1966. Howevcr, we arc not (Bolshakov & Shubnikova 1988). The present willing to accept their conclusion for the following reasons: study is part of a comparative ecological I. We can not kriow which taxon Ogncv described in 1924 investigation aimed at revealing which cir- since the two sibling spccies of 'common voles' can not he distinguished by their external morphology. He did not study cumstances make it possible for a southern vole the characters that distinguish the two (chromosomes. blood species to establish itself in arctic conditions. In proteins or sperm). order to find a relevant reference population it 2. M. arualis 'arualis' and M. arualis 'ohvcurus' occur west was important to find out, if possible, the origin and east of Voroncj, respectivelv. 1s it possiblc to excludc the present occurrence of either of these at the type locality of M. of the ancestors. a. rossiaemeridiormlisis" Only 15 specimens were invcstigatcd. Microtus must have been brought to Svalbard Also. the distribution of thc two sihling spccics may have by man and according to Alendal (1977), the changcd since 1924 ('?)when the holotypc of ro.\iormrridionali.~ voles might have been introduced by whale and was collected. walrus hunters from Holland in the 17th or 18th 3. Wc recommend that this question bc considcred by the Intcrnational Commission on Zoological Nomenclature. century. This conclusion was based on the According to Article 80, Status of casc under consideration (a) identification of the specimens captured in (International Code of Zoological Nomenclature 1985), existing Svalbard as Microtus arvalis. usage is to he maintained until a ruling of the Commission is In 1972 a new Microtus species was described published, Prcscntly, the twoauthoritative taxonomic references (Niethammcr & Krapp 1982; Honacki et al. 1982) usc the name by Meyer et al. It was called M. subarvalis, but M. epiroticus Ondrias 1966. since this name is an absolute homonym of a For thcse reasons we prefer to use the name M. epiroticus fossil species, Microtus subarualis Heller 1933, in the present article. 284 K. Fredga et al. FIR. I Didhution of M. uruulk (solid linc) and M. rpirulicus (\hudowcd) in Europc and western USSR (haacd on Matygin S: Orlov 1974; Kril CI al. 1980: Nicthammcr 8: Krapp 1982: Vorontsov el al. 1984). species of M. arualis, and is found within the the spermatozoa and with regard to several central part of the vast distribution area of M. proteins (Mejer et al. 1972, 1973: Sakiyan et al. arualis (Fig. 1). The new species cannot be 1984). distinguished from M. arvalis by external Since both M. arvalis and M. epiroticus morphology, but the two species have charac- potentially could have been introduced from teristic karyotypes: M. epiroticus has 2n = 54 and Russia with supply ships to some of the permanent M. arvalis has 2n = 46. There is also a difference Russian settlements in Svalbard, it was important between the two species in the size and shape of to study the chromosomes of the Svalbard voles. The ‘common vole’ in Svalbard 285 Furthermore, the comparison of the chromosomes Each locality was first examined for signs of and the mitochondria1 DNA of voles from small rodents (grazing, runways or faeces) in Svalbard with voles from Holland and other parts grassy vegetation. Such signs of Microtus activity of northern Europe might solve the problem of are conspicuous even at low densities. At places origin and perhaps also tell something about the where signs were found multiple-capture live- number of specimens introduced. traps (type: Ugglan) were set. We here show that the Microtus, belonging to Only the area between Bjarndalen, 2 km W a viable population living in natural habitats in Longyearbyen, and Coles Bay (Fig. 2) was found the surroundings of Grumantbyen, Isfjorden, are to be inhabited by Microtus. Signs of Microtus- M. epiroticus and not M. arvalis as believed activity were, however, most pronounced in the earlier (Alendal 1977). grassy slopes under Fuglefjella and the animals seemed to prefer places on peat soil with lush vegetation of grass and herbs, interspersed with patches of boulders. Generally, these preferred Material and methods areas were on stable well-drained ground. During a two-week period in late August 1989, Approximately 100 live-traps were set for three of us visited localities in the Isfjorden area 2‘12 days in the surroundings of Grumantbyen. A where Micrntus either had been captured or ‘seen’ total of 46 individuals were caught, indicating a according to Alendal(1977). Coles Bay and some dense population. The animals were brought alive other localities were also checked (Fig. 2). to the Department of Biology, University of Oslo, where a breeding colony was established. Fifty M. urvalis were live trapped in October 1989 at Lauwersee, NE Holland. All specimens were brought alive to the Department of Biology, University of Oslo, where a breeding colony was established. Six of the voles from Svalbard and six from Holland, 4males and2femalesof eachpopulation, were karyotyped. Chromosome preparations were made from bone marrow by the direct method of Fredga (1987). For G- and C-banding the techniques of Wang & Fedoroff (1972) and Sumner (1972) were used, respectively. Results lOkm M. epiroticus The six voles studied from Svalbard had identical autosomal karyotypes and males were XY, females XX. The chromosome number was 2n = 54 and all chromosomes but the smallest pair of autosomes were telocentric (Fig. 3.4). The X Fig. 2. Map of the Isfjorden area. Circle: Voles caught during chromosome was the largest of the complement the prcsent study. Stars: Voles reported to be present by and the Y the next largest, but close in size to Nyholni (1966). Alendal (1977) or Bolshakov & Shubnikova the largest autosome. This autosome was the only (1988). hut nor found by us examining thc loealitics in August 1989. Trianglcs: Localities with no earlier reports about vole telocentric that could be identified without G- occurrence, but examined by us in August 1989 with negative banding. The rest of the single-armed autosomes results. Squares: Localities previously reported to contain voles decreased continuously in size. A G-banded (Alendal 1977; Bolshakov & Shubnikova 1988) but not karyotype is shown in Fig. 3B. After C-staining, cxarnined by us. 1 = Grurnantbyen, 2 = Longyearbyen, 3 = all autosomes showed centromeric C-bands (Fig. ColesBay, 4 = Kapp Laila.5 = Bjondhavna.Ternpelfjellet, 6 = Alkhornet, 7 = Knpp Lime. 8 = Sassendalen. 9 = Pyramiden, 3C). The distal half of the X stained as a positive 10 = Barentshurg. 11 = Bellsund. C-block and the entire Y appeared dark after 286 K. Fredga et al. Fig. 3. Karyotypes of male Microtus epiroticur from Grumantbyen, Svalbard. A) unbanded, B) G-banded, C) C-banded. Boxed: sex chromosomes from a female with one normal and one deleted X chromosome (Xd). Arrows indicate the position of the centromeres in the sex chromosomes. Preparations from bone marrow. Bar = 10 pm. Same magnification in A. B and C. The ‘common vole’ in Svalbard 287 Fig. 4. Karyotypes of Microtus arualis from Lauwersee, Holland. A) male, unbanded, B) female, G-banded, C) male, C-banded Preparations from bone marrow. Bar = 10 pm, Same magnification in A.
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