Evolution of the common shrew Sorex araneus: chromosomal and molecular aspects

• Autorzy: Wojcik J. M. , Ratkiewicz M. , Searle J.B.
• Języki publikacji: EN

Abstrakty

EN

We review data on the chromosomal variation in the common shrew Sorex araneus Linnaeus, 1758 in the context of recent molecular findings. The article considers all aspects of chromosomal variation in this species: within-population polymorphism, karyotypic races, hybrid zones between karyotypic races, chromosomal evolution, and speciation. The recent molecular data provide vital information on different evolutionary processes such as inbreeding, genetic drift, population expansion, and selective forces. In particular, the molecular data challenge traditional models for the fixation of chro­mosomal variants, provide new insights into the manner of spread of such variants once they are formed and allow in-depth analysis of gene exchange between karyotypic races.

Słowa kluczowe

EN

chromosome variation; history; shrew; common shrew; Sorex araneus; evolution; molecular marker; gene flow; hybrid zone

• Wydawca: -
• Czasopismo: Acta Theriologica. Supplement
• Rocznik: 2002
• Tom: 47
• Numer: 1
• Opis fizyczny: p.139-167,fig.,ref.

Twórcy

autor

Wojcik J. M.

• Polish Academy of Sciences, 17-230 Bialowieza, Poland

autor

Ratkiewicz M.

autor

Searle J.B.

Bibliografia

• Aniskin V. M. and Lukianova I. V. 1989. [A new chromosome race and the analysis of hybridization zone of two karyoforms of Sorex araneus (Insectivora, Soricidae)]. Doklady Akademii Nauk SSSR 309: 1260-1262. [In Russian]
• Baker R. J., Qumsiyeh M. B. and Hoop C. S. 1987. Role of chromosome banding patterns in under­standing mammalian evolution. [In: Current mammalogy. Vol. 1. H. Genoways, ed]. Plenum, New York: 67-96.
• Balloux F., Brünner H., Lugon-Moulin N., Hausser J. and Goudet J. 2000. Microsatellites can be misleading: an empirical and simulation study. Evolution 54: 1414-1422.
• Banaszek A. 1994. The structure of the contact zone between the chromosomal races Druzno and L^gucki Mlyn in the common shrew (Sorex araneus) in north-eastern Poland: preliminary results. Folia Zoologica 43, Suppl. 1: 11-19.
• Banaszek A., Ratkiewicz M., Fedyk S., Szalaj K. A. and Ch^tnicki W. 1996. The chromosomes and isoenzymes in marginal populations of the common shrew (Sorex araneus) in the Vistula delta. Zeitschrift für Sáugetierkunde 61: 65-72.
• Barton N. H. and Bengtsson B. O. 1986. The barrier to genetic exchange between hybridizing populations. Heredity 56: 357-376.
• Barton N. H., Halliday R. B. and Hewitt G. M. 1983. Rare electrophoretic variants in a hybrid zone. Heredity 50: 139-146.
• Barton N. H. and Gale K. S. 1993. Genetic analysis of hybrid zones. [In: Hybrid zones and evolutionary process. R. G. Harrison, ed]. Oxford University Press, New York: 13-45.
• Barton N. H. and Hewitt G. M. 1985. Analysis of hybrid zones. Annual Review of Ecology and Systematics 16: 113-148.
• Bengtsson B. O. and Bodmer W. F. 1976. The fitness of human translocation carriers. Annals of Human Genetics 40: 253-257.
• Bengtsson B. O. and Frykman I. 1990. Karyotype evolution: evidence from the common shrew (Sorex araneus L.). Journal of Evolutionary Biology 3: 85-101.
• Bilton D. T., Mirol P. M., Mascheretti S., Fredga K., Zima J. and Searle J. B. 1998. Mediterranean Europe as an area of endemism for small mammals rather than a source for northwards postglacial colonization. Proceedings of the Royal Society of London, B 265: 1219-1226.
• Brünner H., Fumagalli L. and Hausser J. 1994. A comparison of two contact zones between chromo­somal races of Sorex araneus in the western Alps: karyology and genetics. Folia Zoologica 43, Suppl. 1: 113.
• Brünner H. and Hausser J. 1996. Genetic and karyotypic structure of a hybrid zone between the chromosomal races Cordon and Valais in the common shrew, Sorex araneus. Hereditas 125: 147-158.
• Brünner H., Lugon-Moulin N., Balloux F., Fumagalli L. and Hausser J. 2002. A taxonomical revaluation of the Valais chromosome race of the common shrew Sorex araneus (Insectivora: Soricidae). Acta Theriologica 47 (in press).
• Bulatova N. Sh., Nadjafova R. S. and Krapivko T. P. 2002. Intraspecific phylogenetic relationships in Sorex araneus L.: The southern Baltic subgroup of chromosome races. Russian Journal of Genetics. 38: 64-69Bulatova N., Searle J. B., Bystrakova N., Nadjafova R., Shchipanov N. and Orlov V. 2000. The diversity of chromosome races in Sorex araneus from European Russia. Acta Theriologica 45, Suppl. 1: 33-46.
• Camin J. H. and Sokal R. R. 1965. A method for deducing branching sequences in phylogeny. Evolution 19: 311-326.
• Catzeflis F. 1984. Systématique biochemique, taxonomie et phylogénie des Musaraignes d'Europe (Soricidae, Mammalia). PhD thesis, Universite de Lausanne, Lausanne: 1-164.
• Coyone J. A. and Orr H. A. 1998. The evolutionary genetics of speciation. Philosophical Transactions: Biological Sciences, Royal Society, London B 353: 287-305.
• Croin-Michielsen N. 1966. Intraspecific and interspecific competition in the shrews Sorex araneus L. and Sorex minutus L. Archives Néerlandaises de Zoologie 17: 73-174.
• Demesure B., Comps B. and Petit R. J. 1996. Choloroplast DNA phylogeography of the common beech (Fagus sylvatica L.) in Europe. Evolution 50: 2515-2520.
• Dobzhansky T. 1940. Speciation as a stage in evolutionary divergence. The American Naturalist 74: 312-321.
• Farris J. S., Kluge A. G. and Eckardt M. J. 1970. A numerical approach to phylogenetic systematics. Systematic Zoology 19: 172-191.
• Fedyk S. 1980. Chromosome polymorphism in a population of Sorex araneus L. at Białowieża. Folia Biologica (Kraków) 28: 83-120.
• Fedyk S. 1982. Further studies on chromosome polymorphism of the common shrew. Acta Theriologica 27: 149-153.
• Fedyk S. 1995. [Geographic chromosomal differentiation and hybrid zones between chromosome races of Sorex araneus in north-eastern Poland]. Dissertationes Universitatis Varsoviensis 439, Białystok: 1-125. [In Polish]
• Fedyk S., Banaszek A., Chętnicki W., Cichomska A. and Szałaj K. A. 2000. Reassessment of the range of the Drnholec race: studies on meiosis in Sorex araneus hybrids. Acta Theriologica 45, Suppl. 1: 59-67.
• Fedyk S., Chętnicki W. and Banaszek A. 1991. Genetic differentiation of Polish populations of Sorex araneus L. III. Interchromosomal recombination in a hybrid zone. Evolution 45: 1384-1392.
• Ford P. J. and Graham C. F. 1964. The chromosome number in the common shrew, Sorex araneus. L. Bulletin of the Mammal Society of the British Isles 22: 10-11.
• Fredga K. 1996. The chromosome races of Sorex araneus in Scandinavia. Hereditas 125: 123-135.
• Frykman I. and Bengtsson B. O. 1984. Genetic differentiation in Sorex. III. Electrophoretic analysis of a hybrid zone between two karyotypic races in Sorex araneus. Hereditas 100: 259- 270.
• Frykman I., Simonsen V. and Bengtsson B. O. 1983. Genetic differentiation in Sorex. I. Electrophoretic analysis of the karyotypic races of Sorex araneus in Sweden. Hereditas 99: 279-292.
• Garagna S., Zuccoti M., Searle J. B., Redi C. A. and Wilkinson P. J. 1989. Spermatogenesis in hetero­zygotes for Robertsonian chromosomal rearrangements from natural populations of the common shrew, Sorex araneus. Journal of Reproduction and Fertility 87: 431-438.
• Gyllensten U. and Wilson A. C. 1987. Interspecific mitochondrial DNA transfer and the colonization of Scandinavia by mice. Genetical Research 49: 25-29.
• Haldane J. B. S. 1922. Sex ratio and unisexual sterility in hybrid animals. Journal of Genetics 12: 101-109.
• Halkka L., Söderlund V., Skarén U. and Heikkilä J. 1987. Chromosomal polymorphism and racial evolution of Sorex araneus L. in Finland. Hereditas 106: 257-275.
• Halkka L., Kaikusalo A. and Vakula N. 1994. Revision of Sorex araneus L. chromosome nomenclature, and rece N new to Finland. Annales Zoologici Fennici 31: 283-288.
• Harrison R. G. 1990. Hybrid zones: windows on the evolutionary process. Oxford Surveys of Evolu­tionary Biology 7: 69-128.
• Hatfield T., Barton N. and Searle J. B. 1992. A model of a hybrid zone between two chromosomal races of the common shrew (Sorex araneus). Evolution 46: 1129-1145.
• Hauffe H. C. and Searle J. B. 1993. Extreme karyotypic variation in a Mus musculus domesticus hybrid zone: the tobacco mouse story revisited. Evolution 47: 1374-1395.
• Hausser J., Bosshard F., Taberlet P. and Wójcik J. M. 1991. Relationships between chromosome races and species of Sorex of the araneus group in the western Alps. Mémoires de la Société Vaudoise des Sciences Naturelles 19: 79-95.
• Hausser J., Dannelid E. and Catzeflis F. 1986. Distribution of two karyotypic races of Sorex araneus (Insectivora, Soricidae) in Switzerland and the post-glacial recolonization of the Valais: First results. Zeitschrift für zoologische Systematik und Evolutionsforschung 24: 307-314.
• Hausser J., Fedyk S., Fredga K., Searle J. B., Volobouev V., Wójcik J. M. and Zima J. 1994. Definition and nomenclature of chromosome races of Sorex araneus. Folia Zoologica 43, Suppl. 1: 1-9.
• Hausser J., Fumagalli L. and Taberlet P. 1998. Mitochondrial DNA evolution in shrews. [In: Evolution of shrews. J. M. Wójcik and M. Wolsan, eds]. Mammal Research Institute PAS, Białowieża: 295-308.
• Hedrick P. W. 1981. The establishment of chromosomal variants. Evolution 35: 322-332.
• Heikkilä J. 1989. Population genetics of north temperate shrews (Soricidae). A review. Annales Zoologici Fennici 26: 459-467.
• Hewitt G. M. 1996. Some genetic consequences of ice ages, and their role in divergence and speciation. Biological Journal of the Linnean Society 58: 247-276.
• Howard D. J. 1993. Reinforcement: The origin, dynamics, and fate of an evolutionary hypothesis. [In: Hybrid zones and the evolutionary process. R. G. Harrison, ed]. Oxford University Press, New York: 46-69.
• Ivanitskaya E. Yu. 1994. Comparative cytogenetics and systematics of Sorex: a cladistic approach. [In: Advances in the Biology of Shrews. J. F. Merritt, G. L. Kirkland Jr and R. K. Rose, eds]. Special Publication of Carnegie Museum of Natural History No. 18: 313-323.
• Jiggins C. D. and Mallet J. 2000. Bimodal hybrid zones and speciation. Trends in Ecology and Evolution 15: 250-255.
• Kozlovsky A., Orlov V., Okulova N., Kovalskaya J. and Searle J. B. 2000. Chromosome studies on common shrews from northern and central parts of European Russia. Acta Theriologica 45, Suppl. 1: 27-31.
• Kral B. and Radjabli S. I. 1974. Banding patterns and Robertsonian fusion in the western Siberian population of Sorex araneus (Insectivora, Soricidae). Zoologické Listy 23: 217-227.
• Kreitman M. 1991. Detecting section at the level of DNA. [In: Evolution at the molecular level. R. K. Selander, A. G. Clark and T. S. Whittam, eds]. Sinauer, Sunderland, Massachusetts: 204-221.
• Lande R. 1979. Effective deme sizes during long-term evolution estimated from rates of chromosomal rearrangement. Evolution 33: 234-251.
• Larkin D. M., Serov O. L., Borodin P. M., Zhdanova N. S. and Searle J. B. 2000. Comparative genome mapping in mammals: the shrew map. Acta Theriologica 45, Suppl. 1: 131-141.
• Lugon-Moulin N., Balloux F. and Hausser J. 2000. Genetic differentiation of common shrew Sorex araneus populations among different alpine valleys revealed by microsatellites. Acta Theriologica 45, Suppl. 1: 103-117.
• Lugon-Moulin N., Brünner H., Balloux F., Hausser J. and Goudet J. 1999. Do riverine barriers, history or introgression shape the genetic structuring of a common shrew (Sorex araneus) population? Heredity 83: 155-161.
• Lugon-Moulin N., Wyttenbach A., Brünner H., Goudet J. and Hausser J. 1996. Study of gene flow through a hybrid zone in the common shrew (Sorex araneus) using microsatellites. Hereditas 125: 159-168.
• Lukâcovâ L., Pialek J. and Zima J. 1994. A hybrid zone between the Ulm and Drnholec karyotypic races of Sorex araneus in the Czech Republic. Folia Zoologica 43, Suppl. 1: 37-42.
• Marjoram P. and Donnelly P. 1994. Pairwise comparisons of mitochondrial DNA sequences in sub­divided populations and implications for early human evolution. Genetics 136: 673-683.
• Mercer S. J. 1991. Chromosomal variation of the common shrew Sorex araneus L. in Britain. PhD thesis, University of Oxford, Oxford.Mercer S. J., Searle J. B. and Wallace B. M. N. 1991. Meiotic studies of karyotypically homozygous and heterozygous male common shrews. Mémoires de la Société Vaudoise des Sciences Naturelles 19: 33-43.
• Meylan A. and Hausser J. 1973. Les chromosomes des Sorex du groupe araneus-arcticus (Mammalia, Insectivora). Zeitschrift für Säugetierkunde 38: 143-158.
• Mishta A. V., Searle J. B. and Wójcik J. M. 2000. Karyotypic variation of the common shrew Sorex araneus in Belarus, Estonia, Latvia, Lithuania, and Ukraine. Acta Theriologica 45, Suppl. 1: 47-58.
• Nachman M. W., Boyer S. R., Searle J. B. and Aquadro C. F. 1994. Mitochondrial DNA variation and the evolution of Robertsonian chromosomal races of house mice, Mus domesticus. Genetics 136: 1105-1120.
• Narain Y. and Fredga K. 1996. A hybrid zone between the Hällefors and Uppsala chromosome races of Sorex araneus in central Sweden. Hereditas 125: 137-145.
• Narain Y. and Fredga K. 1997. Meiosis and fertility in common shrews, Sorex araneus, from a chromosomal hybrid zone in central Sweden. Cytogenetics and Cell Genetics 78: 253-259.
• Narain Y. and Fredga K. 1998. Spermatogenesis in common shrews, Sorex araneus, from a hybrid with extensive Robertsonian polymorphism. Cytogenetics and Cell Genetics 80: 158-164.
• Neet C. R. and Hausser J. 1991. Biochemical analysis and determination of living individuals of the Alpine chromosome races and species of the Sorex araneus group. Mémoires de la Société Vaudoise des Sciences Naturelles 19: 97-107.
• Polyakov A. V., Borodin P. M., Lukâcovâ L., Searle J. B. and Zima J. 1997a. The hypothetical Old-Northern chromosome race of Sorex araneus found in the Ural Mts. Annales Zoologici Fennici 34: 139-142.
• Polyakov A. V., Chadova N. B., Rodionova M. I., Panov V. V., Dobrotvorsky A. K., Searle J. B. and Borodin P. M. 1997b. Novosibirsk revisited 24 years on: chromosome polymorphism in the Novosibirsk population of the common shrew Sorex araneus L. Heredity 78: 172-177.
• Polyakov A. V., Panov V. V., Ladygina T. Yu., Bochkarev M. N., Rodionova M. I. and Borodin P. M. 2001. Chromosomal evolution of the common shrew Sorex araneus L. from the Southern Urals and Siberia in the postglacial period. Russian Journal of Genetics 37: 351-357.
• Polyakov A. V., Zima J., Banaszek A., Searle J. B. and Borodin P. M. 2000a. New chromosome races of the common shrew Sorex araneus from Eastern Siberia. Acta Theriologica 45, Suppl. 1: 11-18.
• Polyakov A. V., Zima J., Searle J. B., Borodin P. M. and Ladygina T. 2000b. Chromosome races of the common shrew Sorex araneus in the Ural Mts: a link between Siberia and Scandinavia? Acta Theriologica 45, Suppl. 1: 19-26.
• Prager E. M., Sage R. D., Gyllensten U., Thomas W. K., Hübner R., Jones C. S., Noble L., Searle J. B. and Wilson A. C. 1993. Mitochondrial DNA sequence diversity and the colonization of Scandinavia by house mice from East Holstein. Biological Journal of the Linnean Society 50: 85-122.
• Rafiński J. and Babik W. 2000. Genetic differentiation among northern and southern populations of the moor frog Rana arvalis Nilsson in central Europe. Heredity 84: 610-618.
• Ratkiewicz M. 1997. [Genetic consequences of karyotype divergence in Sorex araneus Linnaeus, 1758, revealed by isoenzymes and mitochondrial DNA]. PhD thesis, Warsaw University, Białystok Branch, Białystok: 1-85. [In Polish]
• Ratkiewicz M., Fedyk S., Banaszek A., Chętnicki W., Szałaj K., Gielly L. and Taberlet P. 2002. The evolutionary history of the two karyotypic groups of the common shrew, Sorex araneus, in Poland. Heredity 88: 235-242.
• Ratkiewicz M., Supruniuk J., Fedyk S., Banaszek A., Chętnicki W. and Szałaj K. A. 2000. Genetic differentiation and gene flow between the Drnholec and Łęgucki Młyn chromosome races of the common shrew Sorex araneus in northern Poland. Acta Theriologica 45, Suppl. 1: 79-91.
• Ratkiewicz M., Wójcik J. M., Lukâcovâ L. and Macholan M. 1996. Nomenclature of electrophoretically studied loci and allozymes in the common shrew (Sorex araneus L.). [In: Abstract of the ISAAC Fourth International Meeting "Evolution in the Sorex araneus group. Cytogenetic and molecular aspects". Norr Malma and Uppsala, Sweden, August 22-27, 1996]. Uppsala University, Uppsala.
• Riginos C. and Nachman M. W. 1999. The origin of a Robertsonian chromosomal translocation in house mice inferred from linked microsatellite markers. Molecular Biology and Evolution 16: 1763-1773.
• Rogers A. R. and Harpending H. 1992. Population growth makes waves in the distribution of pairwise genetic differences. Molecular Biology and Evolution 9: 552-569.
• Searle J. B. 1983. Robertsonian chromosomal variation in the common shrew Sorex araneus L. PhD thesis, University of Aberdeen, Aberdeen.
• Searle J. B. 1984. Three new karyotypic races of the common shrew Sorex araneus (Mammalia: Insectivora) and a phylogeny. Systematic Zoology 33: 184-194.
• Searle J. B. 1985. Isoenzyme variation in the common shrew (Sorex araneus) in Britain, in relation to karyotype. Heredity 55: 175-180.
• Searle J. B. 1986a. Meiotic studies of Robertsonian heterozygotes form natural populations of the common shrew, Sorex araneus L. Cytogenetics and Cell Genetics 41: 154-162.
• Searle J. B. 1986b. Factors responsible for a karyotypic polymorphism in the common shrew, Sorex araneus. Proceedings of the Royal Society London, B 229: 277-298.
• Searle J. B. 1986c. A trimeric esterase in the common shrew (Sorex araneus). Journal of Heredity 77: 121-122.
• Searle J. B. 1988 Selection and Robertsonian variation in nature: The case of the common shrew. [In: The cytogenetics of mammalian autosomal rearrangements. A. Daniel, ed]. Alan R. Liss Inc., New York: 507-531.
• Searle J. B. 1993. Chromosomal hybrid zones in eutherian mammals. [In: Hybrid zones and evolu­tionary process. R. G. Harrison, ed]. Oxford University Press, New York: 309-353.
• Searle J. B., Fedyk S., Fredga K., Hausser J. and Volobouev V. T. 1991. Nomenclature for the chromosomes of the common shrew (Sorex araneus). Mémoires de la Société Vaudoise des Sciences Naturelles 19: 13-22.
• Searle J. B., Hübner R., Wallace B. M. N. and Garagna S. 1990. Robertsonian variation in wild mice and shrews. Chromosomes Today 10: 253-263.
• Searle J. B. and Wilkinson P. J. 1987. Karyotypic variation in the common shrew (Sorex araneus) in Britain - a "Celtic Fringe". Heredity 59: 345-351.
• Searle J. B. and Wójcik J. M. 1998. Chromosomal evolution: the case of Sorex araneus. [In: Evolution of shrews. J. M. Wójcik and M. Wolsan, eds]. Mammal Research Institute PAS, Białowieża: 219-268.
• Stewart J. R. and Lister A. M. 2001. Cryptic northern refugia and the origins of the modern biota. Trends in Ecology and Evolution 16: 608-613.
• Swofford D. L. 1985. PAUP: Phylogenetic Analysis Using Parsimony, Version 2.4. Illinois Natural History Survey, Champaign.
• Szałaj K. A., Fedyk S., Banaszek A., Chętnicki W. and Ratkiewicz M. 1996. A hybrid zone between two chromosome races of the common shrew, Sorex araneus L., in eastern Poland: preliminary results. Hereditas 125: 169-176.
• Taberlet P., Fumagalli L. and Hausser J. 1994. Chromosomal versus mitochondrial DNA evolution: Tracking the evolutionary history of the southwestern European populations of the Sorex araneus group (Mammalia, Insectivora). Evolution 48: 623-636.
• Taberlet P., Fumagalli L., Wust-Saucy A. and Cosson J. 1998. Comparative phylogeography and postglacial colonization routes in Europe. Molecular Ecology 8: 1923-1934.
• Tegelström H. 1987. Genetic variability in mitochondrial DNA in a regional population of the great tit (Parus major). Biochemical Genetics 25: 95-110.
• Volobouev V. T. 1983. Les types du polymorphisme chromosomique et leur rôle evolutif chez les mammiferes (Insectivora, Rodentia et Carnivora). PhD thesis, University of Pierre and Marie Curie, Paris.
• Volobouev V. T. 1989. Phylogenetic relationships of the Sorex araneus-arcticus species complex (Insectivora, Soricidae) based on high-resolution chromosome analysis. Journal of Heredity 80: 284-290.
• Volobouev V. T. and Catzeflis F. 1989. Mechanisms of chromosomal evolution in three European species of the Sorex araneus-arcticus group (Insectivora: Soricidae). Zeitschrift für Zoologische Systematik und Evolutionsforschung 27: 252-262.
• Wallace B. M. N. and Searle J. B. 1990. Synaptonemal complex studies of the common shrew (Sorex araneus). Comparison of Robertsonian heterozygotes and homozygotes by light microscopy. Heredity 65: 359-367.
• Wallace B. M. N. and Searle J. B. 1994. Oogenesis in homozygotes and heterozygotes for Robertsonian chromosomal rearrangements from natural populations of the common shrew, Sorex araneus. Journal of Reproduction and Fertility 100: 231-237.
• White M. J. D. 1978. Modes of speciation. Freeman, San Francisco: 1-455.
• Wójcik J. M. 1991. Chromosomal polymorphism in the common shrew Sorex araneus and its adaptive singnificance. Mémoires de la Société Vaudoise des Sciences Naturelles 19: 51-62.
• Wójcik J. M. 1993. Chromosome races of the common shrew Sorex araneus in Poland: a model of karyotype evolution. Acta Theriologica 38: 315-338.
• Wójcik J. M. and Searle J. B. 1988. The chromosome complement of Sorex granarius - the ancestral karyotype of the common shrew (Sorex araneus)? Heredity 61: 225-229.
• Wójcik J. M. and Wójcik A. M. 1994. Protein variation in the common shrew (Sorex araneus L.) in Poland, in relation to karyotype. Folia Zoologica 43, Suppl. 1: 53-61.
• Wójcik J. M., Wójcik A. M. and Zalewska H. 1996. Chromosome and allozyme variation of the common shrew, Sorex araneus, in different habitats. Hereditas 125: 183-189.
• Wyttenbach A., Borodin P. and Hausser J. 1998. Meiotic drive favours Robertsonian metacentric chromosomes in the common shrew (Sorex araneus, Insectivora, Mammalia). Cytogenetics and Cell Genetics 83: 199-206.
• Wyttenbach A., Goudet J., Cornuet J.-M. and Hausser J. 1999a. Microsatellite variation reveals low genetic subdivision in a chromosome race of Sorex araneus (Mammalia, Insectivora). Journal of Heredity 90: 323-327.
• Wyttenbach A. and Hausser J. 1996. The fixation of metacentric chromosomes during the chromo­somal evolution in the common shrew (Sorex araneus, Insectivora). Hereditas 125: 209-217.
• Wyttenbach A., Narain Y. and Fredga K. 1999b. Genetic structuring and gene flow in a hybrid zone between two chromosome races of the common shrew (Sorex araneus, Insectivora) revealed by microsatellites. Heredity 82: 79-88.
• Zima J. 1991. Is the trend toward low 2Na numbers inescapable for Sorex araneus populations? Mémoires de la Société Vaudoise des Sciences Naturelles 19: 63-71.
• Zima J., Fedyk S., Fredga K., Hausser J., Mishta A., Searle J. B., Volobouev V. T. and Wójcik J. M. 1996. The list of the chromosome races of the common shrew (Sorex araneus). Hereditas 125: 97-107.
• Zima J., Lukâcovâ L. and Macholân M. 1998. Chromosomal evolution in shrews. [In: Evolution of shrews. J. M. Wójcik and M. Wolsan, eds]. Mammal Research Institute PAS, Białowieża: 175-218.
• Zima J., Macholân M., Filippucci M. G., Reiter A., Andreas M., Lipa M. and Krystufek B. 1994. Karyotypic and biochemical status of certain marginal populations of Sorex araneus. Folia Zoologica 43, Suppl. 1: 43-51.
• Zima J., Wójcik J. M. and Horâkovâ M. 1988. The number of karyotypic variants in the common shrew (Sorex araneus). Acta Theriologica 33: 467-475.