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CompCytogen 7(4): 293–303, doi: 10.3897/CompCytogen.v7i4.6159
Bibliography of studies on hybrid zones of the common shrew chromosome races distributed in Russia
Rena S. Nadjafova 1
1 A.N. Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences 33 Leninsky pr., 119071 Moscow, Russia

Corresponding author: Rena S. Nadjafova (renan8@yahoo.com)

Academic editor: Jan Zima

received 27 August 2013 | accepted 29 October 2013 | Published 21 November 2013


(C) 2013 Rena S. Nadjafova. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


For reference, use of the paginated PDF or printed version of this article is recommended.

Citation: Nadjafova RS (2013) Bibliography of studies on hybrid zones of the common shrew chromosome races distributed in Russia. Comparative Cytogenetics 7(4): 293–303. doi: 10.3897/CompCytogen.v7i4.6159

Abstract

The common shrew, Sorex araneus Linnaeus, 1758, has become a model species for cytogenetic and evolutionary studies after discovery of extraordinary Robertsonian polymorphism at the within-species level. Development of differential staining techniques (Q-, R-and G-banding) made it possible to identify the chromosomal arms and their combination in racial karyotypes. Entering into contact with each other, the chromosomal races might form hybrid zones which represent a great interest for understanding of the process of speciation. Until recently all known hybrid zones of S. araneus were localized in Western Europe and only one was identified in Siberia (Russia) between Novosibirsk and Tomsk races (Aniskin and Lukianova 1989, Searle and Wójcik 1998, Polyakov et al. 2011). However, a rapidly growing number of reports on discovery of interracial hybrid zones of Sorex araneus in the European part of Russia and neighboring territories appeared lately. The aim of the present work is to compile the bibliography of all studies covering this topic regardless of the original language and the publishing source which hopefully could make research data more accessible to international scientists. It could also be a productive way to save current history of Sorex araneus researches in full context of the ISACC (International Sorex araneus Cytogenetics Committee) heritage (Searle et al. 2007, Zima 2008).

Keywords

Chromosome races, Hybrid zones, Robertsonian variation, Sorex araneus

Introduction

The common shrew, Sorex araneus Linnaeus, 1758, displays exceptional variability of karyotype derived from intraspecific chromosome rearrangements of the Robertsonian type. Metacentric pairs of Sorex araneus are formed by fusion of originally acrocentric chromosomes at their centromeres in different combinations of arms. As a result, the chromosomes number (2n) varies from 20 to 33, the odd number is due to the presence of karyotype of the Robertsonian heterozygote with one metacentric and two acrocentrics, instead of two homozygous metacentrics or four acrocentrics. At the same time the fundamental number of chromosome arms (FN) remains unchanged and is equal to 40. As far as this process takes place within populations, we could talk about Robertsonian polymorphism which occurs in the vast range of Sorex araneus species.

After the pioneer analysis in Western Europe in the 1950s and 1960s, the studies of Robertsonian polymorphism in Sorex araneus populations started in Russia, widening the area of cytogenetic investigations to include European and Asian parts of the former USSR (Orlov 1974). The observed variations in chromosome arm lengths led to conclusion that Robertsonian fusions might involve different arms in different populations, which resulted in widely varying non-homologous metacentrics (Orlov and Kozlovsky 1969, Ford and Hamerton 1970, Hausser et al. 1985).

Introduction of new methods of chromosome identification (Q-, R- and G-banding) improved the karyotype definition and increased the interest in the common shrew chromosome evolution. The International Sorex araneus Cytogenetics Committee, ISACC was founded at Oxford University in 1987 and until recently international meetings were held every 3 years. The results of its activity were summarized in 2007 by Searle et al. Based on chromosome specific G-banding patterns, Searle et al. (1991) established the standard nomenclature for chromosomes of Sorex araneus. Later rules for differentiation of the intrapopulation variants (polymorphism) from the interpopulation ones (polytypy) as well as from individual karyotype forms were developed (Hausser et al. 1994). Chromosome identification made it possible to describe the chromosomal races of Sorex araneus (Halkka et al. 1974, 1987). Results of karyological studies over the full species range were successively summarized first by Zima et al. (1996) and then by Wójcik et al. (2003). In Russia G-banded chromosomes of the common shrew were first described for a Siberian (Novosibirsk) population by Král and Radjabli in 1974. Results of further studies of high resolution G-banding and chromosome painting of race Novosibirsk represented the species in the international “Atlas of Mammalian Chromosomes” (2006) and in comprehensive comparative studies of Sorex (Biltueva et al. 2011).This race was also used for DAPI karyotyping of the common shrew (Minina et al. 2007).

Currently, no less than 72 chromosomal races are recognized in total (White et al. 2010). The number of Russian chromosomal races has already reached 25 (Orlov et al. 1996, 2007, Bulatova et al. 2000, Shchipanov et al. 2009, Pavlova 2010). Only four of these races are common for Russia and some neighboring areas. They include the following: 1) the Neroosa race which spreads over the southern regions of Russia and Ukraine; 2) the West Dvina race which can be found in Russia – Belarus neighboring regions; 3) the Goldap race which inhabits the Baltic coast area of Poland and Kaliningrad region of western Russia; 4) the Ilomantsi race which occurs in the bordering areas of north-western Russia (Karelia) and Finland (Orlov et al. 1996, 2007, Bulatova et al. 2000, Shchipanov et al. 2009, Borisov et al. 2009a). As anticipated, regular studies of distribution of different races resulted in discoveries of interracial zones of contact in Russia (Shchipanov et al. 2009, Orlov et al. 2012, Pavlova 2013, Shchipanov and Pavlova 2013) and neighboring territories (Borisov et al. 2010, 2013). Due to ISACC activity, research that involves detection of the hybrid zones, as well as discovery and description of the chromosome races continues on a regular basis.

The first case of Sorex araneus interracial hybridization in Russia was presented by Aniskin and Lukianova (1989) for Tomsk and Novosibirsk races in Western Siberia. This hybrid zone is characterized by the high number of the chromosome arm combinations and remains one of the most complex and best studied Sorex araneus hybrid zones (Searle and Wójcik 1998, Polyakov et al. 2011). The hybrids here form a complex meiotic configuration, a long chain of 9 monobrachially homologous acrocentrics and metacentrics. Presumably, chromosome incompatibility proved by meiosis data may induce infertility in hybrids which, in turn, could contribute to promotion of the selection for assortative mating (Searle and Wójcik 1998). Given that racial karyotypes of Sorex araneus as a rule differ by 1–5 variable metacentrics, the hybrids should produce rings or chains of different numbers and length in meiosis. Thus, the simplest heterozygotes form the chain of three, CIII, or ring of four, RIV. The most complex heterozygote was registered in Moscow and Seliger races hybrids in European Russia, and represents the chain of eleven, CXI (Bulatova et al. 2007). As far as the meiotic complications may lead to reduced hybrid reproductive fitness, the incompatibility is to be considered as the first stage in reproductive isolation. There are indications that the Robertsonian rearrangements do not interrupt the existent gene flow in hybrid zones and could not promote speciation in Sorex araneus. Instead, races might be merely remnants of past allopatric differentiation followed by the loss of secondary contact (Horn et al. 2012, Polly et al. 2013), presenting in particular astonishing racial ‘patchwork’.

As has been shown in a variety of recent studies, the number and diversity of the chromosome rearrangements along with the relative variety of hybrid zone types represent a great opportunity both for understanding of the aftereffects and possible connections of chromosome mutations with the morphological, ecological and genetic differentiation in wild populations of common shrews (see Bibliographic list). It seems quite appropriate to recall the forecast made the British cytogeneticists CE Ford and JL Hamerton in 1970 (p. 235): “… shrews displayed multiple patterns of chromosome variation predicting the problems essential for the interpretation of species evolution. Information about hybrid meiosis would be of outstanding value and studies of pregnant females and their embryos from polymorphic populations could give important information about the breeding system and relative fertility. At a more modest level there remain many parts of Europe from which simple identification of the karyotype in samples from the local population could at least help to fill in the still rather fragmentary distribution map of Races A and B and might reveal further unsuspected chromosome variation”. Till now only the second part of this task has been mostly accomplished, while our knowledge of the influence of chromosome rearrangements on cells, specimen and species is still too fragmentary.

The first tribute to the bibliography on the Sorex araneus cytogenetic model was paid by Prof. Jan Zima at the 8th ISACC meeting (2008). To support his idea, we compiled the bibliographical list which includes majority if not all of currently available papers devoted to interracial hybrid zones of Sorex araneus in Russia. The Bibliographic list presented here includes 43 full papers published in national and international scientific editions within the last 40 years. As it shown by the published data, hybrid karyotypes and true hybrid zones were reported for at least 14 out of 25 chromosome races (which are indexed below) of the common shrew that inhabit Russia. This index includes the names of the races and their standard abbreviations, karyotypic diagnosis and F1 hybrids meiotic formula followed by the reference number of the relevant papers from our Bibliographic list.

Bibliographic list *

*Papers from the Bibliographic list referred to in the Introduction and not included in the final References are marked with asterisks.


1. *Aniskin VM, Lukianova IV (1989) A new chromosome race and hybridization zone analysis of two Sorex araneus (Insectivora, Soricidae) karyoforms. Doklady Academii Nauk SSSR 309: 1260–1262. [English Translation (1990): Doklady Biological Sciences (Proceedings of the Academy of Sciences of the USSR) 309: 826–829.]
2. Bannikova AA, Bulatova NS, Kramerov DA (2006) Molecular variability in the common shrew Sorex araneus L. from European Russia and Siberia inferred from the length polymorphism of DNA regions flanked by short interspersed elements (Inter-SINE PCR) and the relationships between the Moscow and Seliger chromosome races. Genetika 42: 737–747. [English Translation: Russian Journal of Genetics 42: 595–604.] doi: 10.1134/S1022795406060020
3. Borisov YM, Cherepanova EV, Orlov VN (2010) A wide hybrid zone of chromosomal races of the common shrew, Sorex araneus Linnaeus, 1758 (Mammalia), between the Dnieper and Berezina rivers (Belarus). Comparative Cytogenetics 4: 195-201. doi: 10.3897/compcytogen.v4i2.43
4. Borisov YM, Kovaleva AA, Irkhin SY, Orlov VN (2009) Zones of contact and joint occurrence of three chromosomal races of the common shrew Sorex araneus L. (Mammalia) in the southern Valdai Hills. Doklady Academii Nauk 428: 275–277. [English Translation: Doklady Biological Sciences 428: 437–439.]
5. *Borisov YM, Kovaleva AA, Springer AM, Cherepanova EV, Kashtal’ian AP, Orlov VN (2009a) Hybrid origin of karyotypic variation in the common shrew, Sorex araneus (Mammalia), from the Dnieper river basin. Doklady Academii Nauk 429: 561–564. [English translation: Doklady Biological Sciences 429: 531–534.]
6. Borisov YM, Kozlovsky AI, Balakirev AE, Demidova TB, Irchin SYu, Maligin VM, Okulova NM, Potapov SG, Shchipanov AN, Orlov VN (2008) Zones of contact of the chromosome races of the common shrew Sorex araneus L. (Insectivora, Mammalia) at the extremes of the Veps stage of the Valdai ice sheet. Siberian Journal of Ecology 15: 763-771. [In Russian]
7. Borisov YM, Kryshchuk IA, Cherepanova EV, Gajduchenko HS, Orlov VN (2013) Chromosomal polymorphism of populations of the common shrew, Sorex araneus L., in Belarus. Acta Theriologica. doi: 10.1007/s13364-013-0160-y
8. *Bulatova NSh, Jones RM, White TA, Shchipanov NA, Pavlova SV, Searle JB (2011) Natural hybridization between extremely divergent chromosomal races of the common shrew (Sorex araneus, Soricidae, Soricomorpha): hybrid zone in European Russia. Journal of Evolutionary Biology 24: 573-586. doi: 10.1111/j.1420-9101.2010.02191.x
9. Bulatova N, Pavlova S (2007) The chromosome race in the epicenter of hybrid zones. The Herald of Vavilov Society for geneticists and breeding scientists 11: 432–435. http://www.bionet.nsc.ru/vogis/pict_pdf/2007/t11_2/vogis_11_2_cont.pdf
10. *Bulatova N, Shchipanov N, Searle JB (2007) The Seliger – Moscow hybrid zone between chromosome races of common shrews – an initial description. Russian Journal of Theriology 6: 111-116.
11. *Bulatova N, Searle JB, Bystrakova N, Nadjafova R, Shchipanov N, Orlov V (2000) The diversity of chromosome races in Sorex araneus from European Russia. Acta Theriologica 45: 33-46.
12. Frisman LV, Borodin PM, Frisman EY (2010) The evolutionary dynamics of hybrid zones of mammals. Regional Problems 13: 56-61. [In Russian]
13. Grigor’eva OO, Shestak AG, Potapov SG, Borisov YM, Irkhin SY, Korablev NP, Orlov VN (2011) The microsatellite polymorphism and gene flow in the contact zone of four common shrew (Sorex araneus L., Mammalia) chromosome races. Izvestiya Academii Nauk 5: 501–510. [English translation: Biology Bulletin 38: 425–433.]
14. Grigor’eva OO, Shestak AG, Sycheva VB, Potapov SG, Borisov YM, Orlov VN (2011) Isolation effect of narrow hybrid zone of Sorex araneus chromosome races. Doklady Academii Nauk 436: 830–833. [English translation: Doklady Biochemistry and Biophysics 436: 41–43.]
15. *Horn A, Basset P, Yannic G, Banaszek A, Borodin PM, Bulatova NS, Jadwiszczak K, Jones RM, Polyakov AV, Ratkiewicz M, Searle JB, Shchipanov NA, Zima J, Hausser J (2012) Chromosomal rearrangements do not seem to affect the gene flow in hybrid zones between karyotypic races of the common shrew (Sorex araneus). Evolution 66: 882-889. doi: 10.1111/j.1558-5646.2011.01478.x
16. Karamysheva TV, Belonogova NM, Rodionova MI, Rubtsov NB, Polyakov AV, Searle JB, Borodin PM (2007) Temporal and spatial distribution of Rad51 protein in spermatocytes of the common shrew Sorex araneus L. (Soricidae, Eulipotyphla). Russian Journal of Theriology 6: 15-19.
17. Matveevsky SN, Pavlova SV, Acaeva MM, Kolomiets OL (2012) Synaptonemal complex analysis of interracial hybrids between the Moscow and Neroosa chromosomal races of the common shrew Sorex araneus showing regular formation of a complex meiotic configuration (ring-of-four). Comparative Cytogenetics 6: 301-314. doi: 10.3897/CompCytogen.v6i3.3701
18. Orlov VN, Borisov YM (2007) Chromosome races of the common shrew Sorex araneus Linnaeus, 1758 (Mammalia: Insectivora) from the south part of Valdai Heights (Russia). Comparative Cytogenetics 1: 101-106.
19. *Orlov VN, Borisov YM, Cherepanova EV, Grigor’eva OO, Shestak AG, Sycheva VB (2012) Narrow hybrid zone between Moscow and Western Dvina chromosomal races and specific features of population isolation in common shrew Sorex araneus (Mammalia). Genetika 48: 80–88. [English translation: Russian Journal of Genetics 48: 70–78.] doi: 10.1134/S1022795412010152
20. Orlov VN, Borisov YM, Cherepanova EV, Milishnikov AN (2013) Assortative mating in the hybrid zones of the common shrew (Sorex araneus, Mammalia) chromosome race West Dvina. Doklady Academii Nauk 451: 110–113. [English translation: Doklady Biological Sciences 451: 217–220.]
21. Orlov VN, Borisov YM, Irkhin SY, Kovaleva AA (2010) Characteristics of the contact zone of three chromosome races of the common shrew Sorex araneus L. (Mammalia) as indices of interpopulation competition. Ekologiya 6: 459–463. [English translation: Russian Journal of Ecology 41: 519–523.]
22. *Orlov VN, Kozlovsky AI, Okulova NM, Balakirev AE (2007) Postglacial recolonisation of European Russia by the common shrew Sorex araneus. Russian Journal of Theriology 6: 97-104.
23. Orlov VN, Sycheva VB, Cherepanova EV, Borisov YM (2013) Craniometric differences between karyotypic races of the common shrew Sorex araneus (Mammalia) as a result of limited hybridization. Genetika 49: 479–490. [English translation: Russian Journal of Genetics 49: 417-427. doi: 10.7868/S0016675813040103]
24. *Pavlova SV (2013) Cytogenetic analysis of a hybrid zone between the Moscow and Neroosa chromosomal races of the common shrew (Sorex araneus) differing by a single WART-like chromosome rearrangement. Tsitologiya 55: 271-274.
25. Pavlova SV, Bulatova NSh (2010) Identification of a novel WART-like chromosome rearrangement in complex heterozygotes in an interracial hybrid zone of the common shrew Sorex araneus L. Genetika 46: 1269–1271. [English translation: Russian Journal of Genetics 46: 1125-1126. doi: 10.1134/S1022795410090292]
26. Pavlova SV, Bulatova NSh, Shchipanov NA (2007) Cytogenetic control of a hybrid zone between two Sorex araneus chromosome races before breeding season. Genetika 43: 1619–1626. [English translation: Russian Journal of Genetics 43: 1357–1363.]
27. Pavlova SV, Kolomiets OL, Bulatova NSh, Searle JB (2008) Demonstration of a WART in a hybrid zone of the common shrew (Sorex araneus Linnaeus, 1758). Comparative Cytogenetics 2: 115–120. www.zin.ru/journals/compcyt
28. Pavlova SV, Bystrakova NV, Bulatova NS, Nadjafova RS, Polyakov AV (2006) Materials for cadastre of the chromosome races of the common shrew Sorex araneus L. Biogeography 13: 42-59. [In Russian]
29. *Polly PD, Polyakov AV, Ilyashenko VB, Onischenko SS, White TA, Shchipanov NA, Bulatova NS, Pavlova SV, Borodin PM, Searle JB (2013) Phenotypic variation across chromosomal hybrid zones of the common shrew (Sorex araneus) indicates reduced gene flow. PLoS ONE 8(7): e67455. doi: 10.1371/journal.pone.0067455
30. Polyakov AV (2008) Hybrid zones between the chromosome races of the common shrew in West Siberia. Siberian Journal of Ecology 15: 773-777. [In Russian]
31. *Polyakov AV, Borodin PM, White TA, Jones RM, Searle JB (2011) Natural hybridization between extremely divergent chromosomal races of the common shrew (Sorex araneus, Soricidae, Soricomorpha): Hybrid zone in Siberia. Journal of Evolutionary Biology 24: 1393-1402. doi: 10.1111/j.1420-9101.2011.02266.x
32. Polyakov AV, Ilyashenko VB, Onischenko SS, Panov V, Borodin P (2009) AFLP diversity between the Novosibirsk and Tomsk chromosome races of the common shrew (Sorex araneus). Comparative Cytogenetics 3: 85-89. doi: 10.3897/compcytogen.v3i2.14
33. Polyakov AV, Ladygina TYu, Bochkarev MN, Rodionova MI, Borodin PM, Panov VV (2001) Chromosomal evolution of the common shrew Sorex araneus L. from the Southern Urals and Siberia in the postglacial period. Genetika 37: 448–455. [English translation: Russian Journal of Genetics 37: 351–357.]
34. Polyakov AV, Onischenko SS, Iliashenko VB, Searle JB, Borodin PM (2002) Morphometric difference between the Novosibirsk and Tomsk chromosome races of the common shrew (Sorex araneus) in a zone of parapatry. Acta Theriologica 47: 381-387. doi: 10.1007/BF03192464
35. Polyakov AV, Volobouev VT, Aniskin VM, Zima J, Searle JB, Borodin PM (2003) Altitudinal partitioning of two chromosome races of the common shrew (Sorex araneus) in West Siberia. Mammalia 67: 201-207. doi: 10.1515/mamm.2003.67.2.201
36. Polyakov AV, Volobouev VT, Borodin PM, Searle JB (1996) Karyotypic races of the common shrew (Sorex araneus) with exceptionally large ranges: the Novosibirsk and Tomsk races of Siberia. Hereditas 125: 109-115. doi: 10.1111/j.1601-5223.1996.00109.x
37. Polyakov AV, Zima J, Banaszek A, Searle JB, Borodin PM (2000) New chromosome races of the common shrew Sorex araneus from eastern Siberia. Acta Theriologica 45, Supplement 1: 11–17.
38. *Searle JB, Wójcik JM (1998) Chromosomal evolution: the case of Sorex araneus. In: Wójcik JM, Wolsan M (Eds) Evolution of shrews. Mammal Research Institute, Polish Academy of Sciences, Białowieža, 219-268.
39. Shchipanov NA, Bulatova NSh, Demidova TB, Bobretsov AV (2008) Chromosomal races of the common shrew (Sorex araneus L.) inhabiting northeastern European Russia: do physical obstacles restrict their distribution? Doklady Academii Nauk 422: 714–717. [English translation: Doklady Biological Sciences 422: 348–351.]
40. *Shchipanov NA, Bulatova NSh, Pavlova SV, Shchipanov AN (2009) The common shrew (Sorex araneus) as a model species in ecological and evolutionary studies. Zoologicheskii Zhurnal 88: 975-989. [In Russian]
41. Shchipanov NA, Bulatova NSh, Pavlova SV (2008) Distribution of two chromosome races of the common shrew (Sorex araneus L.) in the hybrid zone: can a change of the dispersal mode maintain independent gene frequencies? Genetika 44: 734–745. [English translation: Russian Journal of Genetics 44: 635–645.]
42. Shchipanov NA, Pavlova SV (2007) Hybridization of the common shrew (Sorex araneus L.) chromosomal races Moscow and Seliger: The probability of crossing and survival of hybrids. Doklady Academii Nauk of Russia 417: 847–849. [English translation: Doklady Biological Sciences 417: 487–489.]
43. *Shchipanov NA, Pavlova SV (2013) Contact zones and ranges of chromosomal races of the common shrew, Sorex araneus, in northeastern European Russia. Folia Zoologica 62: 24-35.
Index

  1. Kirillov (Kr)

    gm, hi, kq, no, pr

    • Manturovo (F1: gm/mn/no/go, hi, kq, pr; RIV): 22, 43

    • Petchora (F1: gm/gi/hi/hn/no/mo, kq, pr; RVI): 39, 43

  2. Manturovo (Ma)

    go, hi, kq, mn, pr

    • Kirillov (F1: gm/mn/no/go, hi, kq, pr; RIV): 22, 43

    • Petchora (F1: gi/hi/hn/mn/mo/go, kq, pr; RVI): 43

    • Sok (F1: go, kq, hi/ip/pr/mr/mn/hn; RVI): 43

  3. Moscow (Mo)

    gm, hi, kr, no, pq

    • Neroosa (F1: gm/go/no/mn, hi, kr, pq; RIV): 17, 24

    • Seliger (F1: g/gm/mq/pq/pr/kr/ik/hi/hn/no/o; CXI): 1, 6, 8, 10, 15, 21, 23, 25, 26, 27, 29, 40, 41, 42

    • West Dvina (F1: gm, hi/ip/pq/qr/kr/hk, no; RVI): 6, 18, 20, 21, 23, 40

  4. Neroosa (Ne)

    go, hi, kr, mn, pq

    • Moscow (F1: gm/go/no/mn, hi, kr, pq; RIV): 17, 24

  5. Novosibirsk (No)

    go, hn, ik, mp, qr

    • Tomsk (F1: o/go/gk/ik/hi/hn/mn/mp/p, qr; CIX): 1, 11, 15, 16, 28, 29, 30, 31, 32, 34, 35, 36, 38

    • Serov (F1: go, hn, ik/ip/mp/km, qr; RIV): 28, 33

  6. Petchora (Pt)

    gi, hn, kq, mo, pr

    • Kirillov (F1: gi/hi/hn/no/mo/gm, kq, pr; RVI): 39, 43

    • Serov (F1: gi/go/mo/km/kq/qr/pr/ip, hn; RVIII): 43

    • Sok (F1: gi/go/mo/mr/pr/ip, hn, kq; RVI): 43

  7. Seliger (Sl)

    g, hn, ik, mq, o, pr

    • Moscow (F1: g/gm/mq/pq/pr/kr/ik/hi/hn/no/o; CXI): 2, 6, 8, 10, 15, 21, 23, 25, 26, 27, 29, 40, 41, 42

    • West Dvina (F1: g/gm/mq/qr/pr/ip/ik/hk/hn/no/o; CXI): 20

  8. Serov (Se)

    go, hn, ip, km, qr

    • Novosibirsk (F1: go, hn, ik/ip/mp/km, qr; RIV): 28, 33

    • Petchora (F1: gi/go/mo/km/kq/qr/pr/ip, hn; RVIII): 43

    • Sok (F1: go, hn, ip, km/mr/qr/kq; RIV): 43

    • Yuryuzan (F1: go, hn, ip, km/mq/qr/kr; RIV): 40, 43

  9. Sok (So)

    go, hn, ip, kq, mr

    • Manturovo (F1: go, kq, hi/ip/pr/mr/mn/hn; RVI): 43

    • Petchora (F1: gi/go/mo/mr/pr/ip, hn, kq; RVI): 43

    • Serov (F1: go, hn, ip, km/mr/qr/kq; RIV): 43

  10. Strelka (Sr)

    go, hi, k, m, n, p, q, r

    • Tomsk (F1: k/gk/go/o, hi, q/r, m, n, p; CIV): 28, 37

  11. Tomsk (To)

    gk, hi, mn, o, p, qr

    • Novosibirsk (F1: o/go/gk/ik/hi/hn/mn/mp/p, qr; CIX): 1, 11, 15, 16, 28, 29, 30, 31, 32, 34, 35, 36, 38

    • Strelka (F1: k/gk/go/o, hi, q/r, m, n, p; CIV): 28, 37

  12. West Dvina (Wd)

    gm, hk, ip, no, qr

    • Moscow (F1: gm, hi/ip/pq/qr/kr/hk, no; RVI): 6, 19, 20, 21, 23, 40

    • Seliger (F1: g/gm/mq/qr/pr/ip/ik/hk/hn/no/o; CXI): 20

  13. Yuryuzan (Yu)

    go, hn, ip, kr, mq

    • Serov (F1: go, hn, ip, km/mq/qr/kr; RIV): 40, 43

Figure 1.

Schematic view of geographic distribution (slash) of hybrid zones between chromosome races of Sorex araneus in Russia. Standard abbreviations are used for the racial names (see Index).

Acknowledgements

Research grant received from the Russian Foundation of Fundamental Investigations (12-04-01283) is acknowledged. Kind advice and help of Drs. Nina Bulatova and Svetlana Pavlova in collecting of the Russian sources are greatly appreciated.

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