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Published November 1, 2022 | Version v1
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Dental polymorphisms in Crocidura (Soricomorpha: Soricidae) and evolutionary diversification of crocidurine shrew dentition

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Voyta, Leonid L, Abramov, Alexei V, Lavrenchenko, Leonid A, Nicolas, Violaine, Petrova, Ekaterina A, Kryuchkova, Lyudmila Yu (2022): Dental polymorphisms in Crocidura (Soricomorpha: Soricidae) and evolutionary diversification of crocidurine shrew dentition. Zoological Journal of the Linnean Society 196 (3): 1069-1093, DOI: 10.1093/zoolinnean/zlab103, URL: https://academic.oup.com/zoolinnean/article/196/3/1069/6473014

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urn:lsid:plazi.org:pub:5D30B31CFFF3FF8AFFC2FFB0FFF7DB38

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Figure: 10.5281/zenodo.7318347 (DOI)
Figure: 10.5281/zenodo.7318349 (DOI)
Figure: 10.5281/zenodo.7318351 (DOI)
Figure: 10.5281/zenodo.7318353 (DOI)
Figure: 10.5281/zenodo.7318355 (DOI)
Figure: 10.5281/zenodo.7318357 (DOI)
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References

  • Abramov AV, Jenkins PD, Rozhnov VV, Kalinin AA. 2008. Description of a new species of Crocidura (Soricimorpha: Soricidae) from the island of Phu Quoc, Vietnam. Mammalia 72: 269-272.
  • Adler D, Murdoch D. 2021. Package 'rgl', version 0.104.16. Available at: https://cran.r-project.org/web/packages/rgl/rgl. pdf
  • Almecija S, Tallman M, Sallam HM, Fleagle JG , Hammond AS, Seiffert ER. 2019. Early anthropoid femora reveal divergent adaptive trajectories in catarrhine hindlimb evolution. Nature Communications 10: 4778.
  • Ameen C, Hulme-Beaman A, Evin A, Genmonpre M, Britton K, Cucchi T, Larson G, Dobney K. 2017. A landmark-based approach for assessing the reliability of mandibular tooth crowding as a marker of dog domestication. Journal of Archaeological Science 85: 41-50.
  • Bookstein FL. 1991. Morphometric tools for landmark data: geometry and biology. Cambridge: Cambridge University Press.
  • Brummitt RK. 2002. How to chop up a tree. Taxon 51: 31-41.
  • Bulatova N. 2020. Archives: the law of homologous series in variation (N. I. Vavilov). Comparative Cytogenetics 14: 329-338.
  • Burgin CJ, He K. 2018. Family Soricidae. In: Wilson DE, Russell AM, eds. Handbook of the mammals of the world, Vol. 8. Insectivores, sloths and colugos. Barcelona: Lynx Edicions, 332-551.
  • Butler PM. 1998. Fossil history of shrews in Africa. In: Wojcik JM, Wolsan M, eds. Evolution of shrews. Bialowieza: Mammal Research Institute, Polish Academy of Sciences, 121-132.
  • Butler PM, Greenwood M. 1979. Soricidae (Mammalia) from the Early Pleistocene of Olduvai Gorge, Tanzania. Zoological Journal of the Linnean Society 67: 329-379.
  • Butler PM, Thorpe RS, Greenwood M. 1989. Interspecific relations of African crocidurine shrews (Mammalia: Soricidae) based on multivariate analysis of mandibular date. Zoological Journal of the Linnean Society 96: 373-412.
  • Ceriaco LMP, Marques MP, Jacquet F, Nicolas V, Colyn M, Denys C, Sardinha PC, Bastos-Silveira C. 2015. Description of a new endemic species of shrew (Mammalia, Soricomorpha) from Principe Island (Gulf of Guinea). Mammalia 79: 325-341.
  • Chen S, Qing J, Liu Z, Liu Y, Tang M, Murphy RW, Pu Y, Wang X, Tang K, Guo K, Jiang X, Liu S. 2020. Multilocus phylogeny and cryptic diversity of white-toothed shrews (Mammalia, Eulipotyphla, Crocidura) in China. BMC Evolutionary Biology 20: 29.
  • Churchfield S. 2013. Family Soricidae. Shrews.In: Happold M, Happold DCD, eds. Mammals of Africa: hedgehogs, shrews and bats, vol. IV. London: Bloomsbury, 43-50.
  • Claude J. 2008. Morphometrics with R. New York: Springer Science+Business Media.
  • Cornette R, Baylac M, Souter T, Herrel A. 2013. Does shape co-variation between the skull and the mandible have functional consequences? A 3D approach for a 3D problem. Journal of Anatomy 223: 329-336.
  • Cornette R, Tresset A, Herrel A. 2015a. The shrew tamed by Wolff 's law: do functional constraints shape the skull through muscle and bone covariation? Journal of Morphology 276: 301-309.
  • Cornette R, Tresset A, Houssin C, Pascal M, Herrel A. 2015b. Does bite force provide a competitive advantage in shrews? The case of the greater white-toothed shrew. Biological Journal of the Linnean Society 114: 795-807.
  • Cucchi T, Mohaseb A, Peigne S, Debue K, Orlando L, Mashkour M. 2017. Detecting taxonomic and phylogenetic signals in equid cheek teeth: towards new palaeontological and archaeological proxies. Royal Society Open Science 4: 160997.
  • Dannelid E. 1998. Dental adaptations in shrew. In : Wojcik JM, Wolsan M, eds. Evolution of shrews. Bialowieza: Mammal Research Institute, Polish Academy of Sciences, 157-174.
  • Demos TC, Achmadi AS, Handika H, Maharadatunkamsi, Rowe KC, Esselstyn JA. 2017. A new species of shrew (Soricomorpha: Crocidura) from Java, Indonesia: possible character displacement despite interspecific gene flow. Journal of Mammalogy 98: 183-193.
  • Drake AG. 2011. Dispelling dog dogma: an investigation of heterochrony in dogs using 3D geometric morphometric analysis of skull shape. Evolution & Development 13: 204-213.
  • Drake AG, Klingenberg CP. 2008. The pace of morphological change: historical transformation of skull shape in St Bernard dogs. Proceedings of the Royal Society B: Biological Sciences 275: 71-76.
  • Dubey S, Salamin N, Ohdachi SD, Barriere P, Vogel P. 2007. Molecular phylogenetics of shrews (Mammalia: Soricidae) reveal timing of transcontinental colonizations. Molecular Phylogenetics and Evolution 44: 126-137.
  • Dubey S, Salamin N, Ruedi M, Barriere P, Colyn M, Vogel P. 2008. Biogeographic origin and radiation of the Old World crocidurine shrews (Mammalia: Soricidae) inferred from mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 48: 953-963.
  • Eble GJ. 2000. Contrasting evolutionary flexibility in sister groups: disparity and diversity in Mesozoic atelostomate echinoids. Paleobiology 26: 56-79.
  • Eble GJ. 2002. Developmental morphospaces and evolution. In: Crutchfield JP, Schuster P, eds. Evolution dynamics. Oxford: Oxford University Press, 35-65.
  • Ekdale EG, Racicot RA. 2015. Anatomical evidence for low frequency sensitivity in an archaeocete whale: comparison of the inner ear of Zygorhiza kochii with that of crown Mysticeti. Journal of Anatomy 226: 22-39.
  • Engesser B. 1980. Insectivora und Chiroptera (Mammalia) aus dem Neogen der Turkei. Schweizerische Palaontologische Abhandlungen 102: 4-149.
  • Esselstyn JA, Goodman SM. 2010. New species of shrew (Soricidae: Crocidura) from Sibuyan Island, Philippines. Journal of Mammalogy 91: 1467-1472.
  • Flynn LJ, Jacobs LL, Kimura Y, Taylor LH, Tomida Y. 2020. Siwalik fossil Soricidae: a calibration point for the molecular phylogeny of Suncus. Paludicola 12: 247-258.
  • Furio M, Santos-Cubedo A, Minwer-Barakat R, Agusti J. 2007. Evolutionary history of the African soricid Myosorex (Insectivora, Mammalia) out of Africa. Journal of Vertebrate Paleontology 27: 1018-1032.
  • Geiger M, Evin A, Sanchez-Villagra MR, Gascho D, Mainini C, Zollikofer CPE. 2017. Neomorphosis and heterochrony of skull shape in dog domestication. Scientific Reports 7: 13443.
  • Gould GC. 2001. The phylogenetic resolving power of discrete dental morphology among extant hedgehogs and the implications for their fossil record. American Museum Novitates 3340: 1-52.
  • Hinckley A , Camacho-Sanchez M , Ruedi M , Hawkins MTR, Mullon M, Cornellas A, Tuh Yit Yuh F, Leonard JA. 2021. Evolutionary history of Sundaland shrews (Eulipotyphla: Soricidae: Crocidura) with a focus on Borneo. Zoological Journal of the Linnean Society 194(2): 478-501.
  • Hugueney M, Maridet O. 2011. Early Miocene soricids (Insectivora, Mammalia) from Limagne (Central France): new systematic comparisons, updated biostratigraphic data and evolutionary implications. Geobios 44: 225-236.
  • Hutterer R. 2005a. Order Soricomorpha. In: Wilson DE, Reeder DA, eds. Mammal species of the world: a taxonomical reference, 3rd edn, Vol. 1. Baltimore: Johns Hopkins University Press, 220-311.
  • Hutterer R. 2005b. Homology of unicuspid and tooth nomenclature in shrews. In: Merritt JF, Churcfield S, Hutterer R, Sheftel BI, eds. Advances in the biology of shrews II. New York: International Society of Shrew Biologists, 397-404.
  • Hutterer R, Balete DS , Giarla TC, Heaney LR , Esselstyn JA. 2018. A new genus and species of shrew (Mammalia: Soricidae) from Palawan Island, Philippines. Journal of Mammalogy 99: 518-536.
  • Jackson DA. 1993. Stopping rules in principal components analysis: a comparison of heuristical and statistical approaches. Ecology 74: 2204-2214.
  • Jacquet F, Hutterer R, Nicolas V, Decher J, Colyn M, Couloux A, Denys C. 2013. New status for two African giant forest shrews, Crocidura goliath goliath and C. goliath nimbasilvanus (Mammalia: Soricomorpha), based on molecular and geometic morphometric analyses. African Zoology 48: 13-29.
  • Jarvinen E, Salazar-Ciudad I, Birchmeier W, Taketo MM, Jernvall J, Thesleff I. 2006. Continuous tooth generation in mouse is induced by activated epithelial Wnt/β- catenin signaling. Proceedings of the National Academy of Sciences of the United States of America 103: 18627-18632.
  • Jenkins PD, Abramov AV, Bannikova AA, Rozhnov VV. 2013. Bones and genes: resolution problems in three Vietnamese species of Crocidura (Mammalia, Soricomorpha, Soricidae) and the description of an additional new species. ZooKeys 313: 61-79.
  • Jenkins PD, Abramov AV, Rozhnov VV, Makarova OV. 2007. Description of two new species of white-toothed shrews belonging to the genus Crocidura (Soricomorpha: Soricidae) from Ngoc Linh Mountain, Vietnam. Zootaxa 1589: 57-68.
  • Jenkins PD, Abramov AV, Rozhnov VV, Ollson A. 2010. A new species of Crocidura (Soricomorpha: Soricidae) from southern Vietnam and north-eastern Cambodia. Zootaxa 2345: 60-68.
  • Jenkins PD, Lunde DP, Moncrieff CB. 2009. Descriptions of new species of Crocidura (Soricomorpha: Soricidae) from mainland Southeast Asia, with synopses of previously described species and remarks on biogeography. Bulletin of the American Museum of Natural History 331: 356-405.
  • Kamalakannan M, Sivaperuman C, Kundu S , Gokulakrishnan G, Venkatraman C, Chandra K. 2021. Discovery of a new mammal species (Soricidae:Eulipotyphla) from Narcondam volcanic island, India. Scientific Reports 11: 9416.
  • Kavanagh KD, Evans AR, Jernvall J. 2007. Predicting evolutionary patterns of mammalian teeth from development. Nature 449: 427-432.
  • Klingenberg CP. 2011. MorphoJ: an integrated software package for geometric morphometrics. Molecular Ecology Resources 11: 353-357.
  • Konecny A, Hutterer R, Meheretu Y, Bryja J. 2020. Two new species of Crocidura (Mammalia: Soricidae) from Ethiopia and updates on the Ethiopian shrew fauna. Journal of Vertebrate Biology 69: 20064.
  • Koyasu K, Kawahito K, Hanamura H, Oda S. 2005. Dental anomalies in Suncus murinus. In: Merritt JF, Churcfield S, Hutterer R, Sheftel BI, eds. Advances in the biology of shrews II. New York: International Society of Shrew Biologists, 405-411.
  • Kurachi M, Chau BL, Dang VB, Dorji T, Yamamoto Y, Nyunt MM, Maeda Y, Chhum-Phith L, Namikawa T, Yamagata T. 2007. Population structure of wild musk shrews (Suncus murinus) in Asia based on mitochondrial DNA variation, with research in Cambodia and Bhutan. Biochemical Genetics 45: 165-183.
  • Lavrenchenko LA, Voyta LL, Hutterer R. 2016. Diversity of shrews in Ethiopia, with the description of two new species of Crocidura (Mammalia: Lipotyphla: Soricidae). Zootaxa 4196: 38-60.
  • Lebrun R. 2020. MorphoDig user's guide. Available at: https:// morphomuseum.com/tutorialsMorphoDig
  • Lopatin AV. 2006. Early Paleogene insectivore mammals of Asia and establishment of the major group of Insectivora. Paleontological Journal 40: 205-405.
  • Maddalena T. 1989. Systematics and biogeography of Afrotropical and Palaearctic shrews of the genus Crocidura (Insectivora: Soricidae): an electrophoretic approach. In: Peters G, Hutterer R, eds. Vertebrates in the tropics. Bonn: Museum Alexander Koenig, 297-308.
  • Maddison WP, Maddison DR. 2019. Mesquite: a modular system for evolutionary analysis. Version 3.61. Available at: http://www.mesquiteproject.org
  • Mason MJ. 2016. Structure and function of the mammalian middle ear. I: large middle ears in small desert mammals. Journal of Anatomy 228: 284-299.
  • Masuda Y, Yohro T. 1994. The development of the skull of Suncus murinus. In: Merritt JF, Kirkland GL, Rose RK, eds. Advances in the biology of shrews. Pittsburg: Carnegie Museum of Natural History, 171-180.
  • McLellan LJ. 1994. Evolution and phylogenetic affinities of the African species of Crocidura, Suncus, and Sylvisorex (Insectivora: Soricidae). In: Merritt JF, Kirkland GL, Rose RK, eds. Advances in the biology of shrews. Pittsburg: Carnegie Museum of Natural History, 379-391.
  • Meegaskumbura S, Meegaskumbura M, Pethiyagoda R, Manamendra-archchi K, Schneider CJ. 2007. Crocidura hikmiya, a new shrew (Mammalia: Soricomorpha: Soricidae) from Sri Lanka. Zootaxa 1665: 19-30.
  • Meegaskumbura S, Schneider CJ. 2008. A taxonomic evaluation of the shrew Suncus montanus (Soricidae: Crocidurinae) of Sri Lanka and India. Ceylon Journal of Science (Biological Sciences) 37: 129-136.
  • Ni X, Flynn JJ, Wyss AR. 2012. Imaging the inner ear in fossil mammals: high-resolution CT scanning and 3-D virtual reconstruction. Palaentologia Electronica 15.2.18A: 1-10.
  • Nixon RC. 2003. WinClada ver. 1.00.08. Ithaca: published by the author.
  • Ohdachi S, Kinoshita G, Nasher AK, Yonezawa T, Arai S, Kikichi F, Lin KS, Bawm S. 2017. Re-evaluation of the phylogeny based on mitochondrial cytochrome b gene in the house shrew, Suncus murinus-S. montanus species complex, with special reference to Yemen and Myanmar populations. Journal of Wildlife and Biodiversity 1: 79-87.
  • Ohdachi S, Kinoshita G, Oda SI , Motokawa M, Jogahara T, Arai S, Nguyen ST, Suzuki H, Katakura K, Bawm S, Min MZ, Thwe TL, Gamage CD, Hashim R, Omar H, Maryanto I, Ghadirian T, Ranorosoa MC, Moribe J, Tsuchiya K. 2016. Intraspecific phylogeny of the house shrews, Suncus murinus-S. montanus species complex, based on the mitochondrial cytochrome b gene. Mammal Study 41: 229-238.
  • O'Higgins P, Jones N. 1998. Facial growth in Cercocebus torquatus: an application of three-dimensional geometric morphometric techniques to the study of morphological variation. Journal of Anatomy 193: 251-272.
  • O'Leary MA, Kaufman SG. 2007. MorphoBank 2.5: web application for morphological phylogenetics and taxonomy. Available at: https://morphobank.org/
  • Orliac MJ , Billet G. 2016 . Fallen in a dead ear : intralabyrinthine preservation of stapes in fossil artiodactyls. Palaeovertebrata 40: 1-10.
  • Polly PD. 2007. Phylogeographic differentiation in Sorex araneus: morphology in relation to geography and karyotype. Russian Journal of Theriology 6: 73-84.
  • Polly PD. 2019. Spatial processes and evolutionary models: a critical review. Palaeontology 62: 175-195.
  • Polly PD, Wojcik JM. 2019. Geometric morphometric tests for phenotypic divergence between chromosomal races. In: Searle JB, Zima J, Polly PD, eds. Shrews, chromosomes and speciation. Cambridge: Cambridge University Press, 336-364.
  • Querouil S, Hutterer R , Barriere P , Colyn M , Kerbis Peterhans JC, Verheyen E. 2001. Phylogeny and evolution of African shrews (Mammalia: Soricidae) inferred from 16s rRNA sequences. Molecular Phylogenetics and Evolution 20: 185-195.
  • Renvoise E, Kavanagh KD, Lazzari V, Hakkinen TJ, Rice R, Pantalacci S, Salazar-Ciudad I, Jernvall J. 2017. Mechanical constraint from growing jaw facilitates mammalian dental diversity. Proceedings of the National Academy of Sciences of the United States of America 114: 9403-9408.
  • Repenning CA. 1967. Subfamilies and genera of the Soricidae. Classification, historical zoogeography and temporal correlation of the shrews. United States Survey Professional Paper 565: 1-74.
  • Reumer JWF. 1984. Ruscinian and Early Pleistocene Soricidae (Insectivora, Mammalia) from Tegelen (The Netherlands) and Hungary. Scripta Geologica 73: 1-173.
  • Reumer JWF. 1987. Redefinition of the Soricidae and the Heterosoricidae (Insectivora, Mammalia) with the description of the Crocidosoricinae, a new subfamily of Soricidae. Revue de Paleobiologie 6: 89-192.
  • Reumer JWF. 1989. Speciation and evolution in the Soricidae (Mammalia: Insectivora) in relation with the paleoclimate. Revue Suisse de Zoologie 96: 81-90.
  • Rofes J, Cuenca-Bescos G. 2009. A new genus of red-toothed shrew (Mammalia, Soricidae) from the early Pleistocene of Gran Dolina (Atapuerca, Burgos, Spain), and a phylogenetic approach to the Eurasian Soricinae. Zoological Journal of the Linnean Society 155: 904-925.
  • Ruedi M, Maddalena T, Vogel P, Obara Y. 1993. Systematic and biogeographic relationships of the Japanese white-toothed shrew (Crocidura dsinezumi). Journal of Mammology 74: 535-543.
  • Schlager S. 2017. Morpho and Rvcg - shape analysis in R: R-packages for geometric morphometrics, shape analysis and surface manipulations. In: Zheng G, Li S, Szekely G, eds. Statistical shape and deformation analysis, 1st edn. San Diego: Academic Press, 217-256.
  • Sese C. 1980. Mamiferos del Miocene Medio de Escobosa de Calatanazor. Madrid: Universidad Complutense de Madrid.
  • Sheftel BI. 2018. Piebald shrew Diplomesodon pulchellus. In: Wilson DE, Russell AM, eds. Handbook of the mammals of the world, Vol. 8. Insectivores, sloths and colugos. Barcelona: Lynx Edicions, 472.
  • Storch G, Qiu Z, Zazhigin V. 1998. Fossil history of shrews in Asia. In: Wojcik JM, Wolsan M, eds. Evolution of shrews. Bialowieza: Mammal Research Institute, Polish Academy of Sciences, 93-120.
  • Swofford DL.1998. PAUP:phylogeneticanalysisusingparsimony (and other methods). Sunderland: Sinauer Associates.
  • Thorn E, Kerbis Peterhans J. 2009. Small mammals of Uganda. Bonner Zoologischer Monographen 55: 1-126.
  • Upham NS, Esselstyn JA, Jetz W. 2019. Inferring the mammal tree: species-level sets of phylogenies for questions in ecology, evolution, and conservation. PLoS Biology 17: e3000494.
  • Van Dam JA, Van den Hoek Ostende LW, Reumer JWF. 2011. A new short-snouted shrew from the Miocene of Spain. Geobios 44: 299-307.
  • Voyta LL, Omelko VE, Tiunov MP, Vinokurova MV. 2021. When beremendiin shrews disappeared in East Asia, or how we can estimate fossil redeposition. Historical Biology 33: 2656-2667.
  • Voyta LL, Zazhigin VS, Petrova EA, Krjutchkova LYu. 2020. Shrew dentition (Lipotyphla: Soricidae)-endodontic morphology and its phylogenetic resolving power. Mammal Research 65: 33-48.
  • Wible JR. 2008. On the cranial osteology of the hispaniolan solenodon, Solenodon paradoxus Brandt, 1833 (Mammalia, Lipotyphla, Solenodontidae). Annals of Carnegie Museum 77: 321-402.
  • Wiens JJ. 1995. Polymorphic characters in phylogenetic systematics. Systematic Biology 44: 482-500.
  • Wills M, Briggs DEG, Fortey RA. 1994. Disparity as an evolutionary index: a comparison of Cambrian and Recent arthropods. Paleobiology 20: 93-130.
  • Woodman N, Morgan JJ. 2005. Skeletal morphology of the forefoot in shrews (Mammalia: Soricidae) of the genus Cryptotis, as revealed by digital X-rays. Journal of Morphology 266: 60-73.
  • Wu X, Schepartz LA. 2009. Application of computed tomography in paleoanthropological research. Progress in Natural Science 19: 913-921.
  • Yamagata T, Ohishi K, Faruque MO, Masangkay JS, Ba-Loc C,Vu-Binh D,Mansjoer SS,Ikeda H,Namikawa T. 1995. Genetic variation and geographic distribution on the mitochondrial DNA in local populations of the musk shrew, Suncus murinus. Idengaku zasshi 70: 321-337.
  • Yamanaka A, Yasui K, Sonomura T, Iwai H, Uemura M. 2010. Development of deciduous and permanent dentitions in the upper jaw of the house shrew (Suncus murinus). Archives of Oral Biology 55: 279-287.
  • Yang L, Zhang H, Zhang C, Wu J, Wang Z, Li C, Zhang B. 2020. A new species of the genus Crocidura (Mammalia: Eulipotyphla: Soricidae) from Mount Huang, China. Zoological Systematics 45: 1-14.
  • Young RL, Badyaev AV. 2010. Developmental plasticity links local adaptation and evolutionary diversification in foraging morphology. Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 314: 434-444.
  • Zaitseva AS, Volodin IA, Mason MJ, Frey R, Fritsch G, Ilchenko OG, Volodina EV. 2015. Vocal development during postnatal growth and ear morphology in a shrew that generates seismic vibrations, Diplomesodon pulchellum. Behavioural Processes 118: 130-141.
  • Zamudio KR, Bell RC, Mason NA. 2016. Phenotypes in phylogeography: species' traits, environmental variation, and vertebrate diversification. Proceedings of the National Academy of Sciences of the United States of America 113: 8041-8048.
  • Zhang H, Wu GY, Wu YQ, Yao JF, You S, Wang CC, Cheng F, Chen JP, Tang MX, Li CL, Zhang BW. 2019. A new species of the genus Crocidura from China based on molecular and morphological data (Eulipotyphla: Soricidae). Zoological Systematics 44: 279-293.
  • Ziegler R. 2009. Soricids (Soricidae, Mammalia) from Early Oligocene fissure fillings in South Germany - and a phylogenetic analysis of the Heterosoricinae. Palaeodiversity 2: 321-342.