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The G-banded karyotype of Sorex satunini was compared with the karyotype of Sorex araneus. Extensive homology was revealed. The major chromosomal rearrangements involved in the evolutionary divergence of these species have been identified as centric fusions and centromeric shifts. From the known palaeontological age of Sorex satunini it is obvious that the vast chromosomal polymorphism of the Sorex araneus group originated during the middle Pleistocene.
G-banding, common shrew, Sorex araneus, Sorex satunini, karyotype, phylogeny
Within the genus Sorex Linnaeus, 1758, the Sorex araneus group includes eight species characterized by the sex chromosome complex XY1Y2 (
The common shrew is widely distributed in Europe and Asia up to as far east as Lake Baikal, and the Caucasian shrew is known to be present in the Caucasus and in the northern parts of Asia Minor (
The common shrew displays phenomenal variability of the autosomal complement (
From the plain between the Kuban and Don rivers we described a new subspecies of the Caucasian shrew Sorex satunini tembotovi Orlov, Balakirev, Borisov, 2010(
In this study the karyotypes of Sorex satunini tembotovi and Sorex araneus (chromosome race Moscow) were examined and compared.
Material and methodsThree females and four males of Sorex satunini were captured in the valley of the Beisoog River (45°40'N, 39°41'E), 90 km N of the Krasnodar city in June 2009. Two shrews of the race Moscow (male and female) were captured in Moscow vicinity.
Mitotic chromosome spreads were prepared in the field conditions from bone marrow and spleen cells using the air-drying technique after fixation with methanol and glacial acetic acid. For G-banding, the slides were treated with trypsin solution according to
The karyotype of Sorex satunini consists of 24–25 chromosomes. The sex chromosomes are a large metacentric X, a small acrocentric Y1, and a medium-sized Y2. Of 11 autosomal pairs, only a single pair of small chromosomes is acrocentric, all other autosomes are biarmed. Such a karyotype has been described for many populations from the North Caucasian and Transcaucasian regions (
The Caucasian shrew has the following chromosome formula: XX / XY1Y2, af, bc, gh, ik, jn, lo, tu, m, p, q, r, tu. The comparison of the G-banded metaphase chromosomes of Sorex satunini and of Sorex araneus (the race Moscow) is presented in Fig. 1. This comparison revealed a considerable homology between individual chromosomal arms. Identical banding patterns and centromeric positions were found in two large biarmed autosomes af and bc, in small metacentric tu, in acrocentric pare m and in the sex chromosomes (Fig. 1a). The acrocentric m is found in the karyotypes of the chromosomal races of Sorex araneus either as an individual acrocentric, or in combination with other acrocentrics.
Seven arms of Sorex araneus, namely g, i, k, j, n, l, and o, were also identified in the complement of Sorex satunini. The difference in G-banding of the arm h between Sorex satunini and other species of the Sorex araneus group was observed (Fig. 1b). An identical banding pattern and a different centromeric position were found in three autosomal pairs: p, q, and r, suggesting occurrence of centromeric shift. The chromosomes p, q, and r were found to be metacentric in the complement of Sorex satunini (Fig. 1c) and acrocentric in Sorex araneus.
The large biarmed chromosome bc was also identified in the complement of Sorex antinorii Bonaparte, 1840, and only in Sorex coronatus Millet, 1828it was substituted by ci (
The G-banded karyotype of Sorex satunini (male) in comparison with the karyotype of Sorex araneus (the race Moscow, male). The chromosomes of the race Moscow are given in the frameworks. Some chromosomes are identical (a), the others are different because of the arms involved in different fusions (b) or because of the centromeric shift (c). Centromere position is indicated by “<”. Bar = 3 µm.
In karyotype of Sorex coronatus there are only two species-specific chromosome rearrangements (Rb fusions ci and mp). In karyotype of Sorex antinorii there are only two species-specific chromosome rearrangements, too (hj and kn). In karyotype of Sorex satunini there are five species-specific chromosome rearrangements (Rb fusion gh, centromeric shifts in the chromosomes p, q, r, and, probably, a paracentric inversionin the chromosomal arm h), i.e. Sorex satunini has more rearranged karyotype than the species of Sorex araneus group in Western Europe.
A number of chromosome rearrangements shared by Sorex araneus, Sorex satunini and Sorex antinorii (centric fusions bc), by Sorex coronatus, Sorex satunini and Sorex antinorii (lo), by Sorex coronatus and Sorex satunini (jn) suggest the existence of a common ancestral species in the Pleistocene of Europe analogous to the modern Sorex araneus.
The known paleontological age points to an early origin of Sorex satunini. At present, the dating of fossils confirmed by the radiocarbohydrate analysis is known only for Sorex satunini. These fossils, morphologically very similar to the recent Sorex satunini were found in the Transcaucasian region (Kudaro caves) in all layers of the middle and late Pleistocene, beginning since 0.36 Myr BP (
We are grateful to V.V. Stacheev, A.E. Balakirev for the assistance during the expedition andE.V. Cherepanova for the comments on the manuscript. The study was supported by the Russian Foundation for Basic Research (grant 12-04-00551) and Program of the Russian Academy of Sciences “Dynamics and conservation of gene pools”.