Eulipotyphla

Reconstruction of Nesophontes, copyright Michael R. Long.

Belongs within: Pan-Lipotyphla.
Contains: Talpidae, Erinaceomorpha, Soricidae.

The Eulipotyphla, or 'insectivores', are a group of small mammals including shrews (Soricidae), hedgehogs (Erinaceidae), moles (Talpidae) and others. They are supported as a clade by molecular analyses, albeit one that excludes the African families Tenrecidae (tenrecs) and Chrysochloridae (golden moles) previously regarded as close relatives; these latter groups are instead allied to other Africa-derived mammals in a group called the Afrotheria. A number of fossil taxa previously regarded as related to this group are of uncertain placement as a result of this divide. Future analyses may identify them as true eulipotyphlans, as afrotherians, or as independent extinct groups.

Among the more divergent recent eulipotyphlans are the Solenodontidae (solenodons), moderately large (about a foot long excluding the tail) insectivores found only on the islands of Cuba and Hispaniola. Until recently, the Caribbean islands were also home to the now-extinct species of Nesophontes which were smaller than modern solenodons and possibly more superficially shrew-like. The Parapternodontidae from the early Eocene of Wyoming may represent continental relatives of solenodons (Halliday et al. 2017) but were placed closer to Soricidae by Asher et al. (2002).

Characters (from Butler 1988): Mobile snout, with enlarged olfactory organs, enlarged infraorbital canal, and reduced eyes. Palatine displaced from lacrimal by maxillar wing and confined to orbital floor; jugal reduced or absent, lower border of orbit formed from maxilla. Pubic symphysis reduced or lost. Caecum lost.

Eulipotyphla [Centetoidea, Dormaalioidea, Erinaceota, Lipotyphla, Soricoidea, Soricomorpha] HUG17
    |--+--+--+--Talpidae HUG17
    |  |  |  |--Proscalopidae [Arctoryctinae, Proscalopinae] B88
    |  |  |  |    |--Cryptoryctes Reed 1954 V67
    |  |  |  |    |--Oligoscalops Reed 1961 V67
    |  |  |  |    |--Proscalops Matthew 1901 [incl. Arctoryctes Matthew 1907] V67
    |  |  |  |    `--Mesoscalops Reed 1960 V67
    |  |  |  `--Dimylidae B88
    |  |  |       |--Dimylinae V67
    |  |  |       |    |--Dimylus von Meyer 1846 V67
    |  |  |       |    |    `--D. paradoxus D07
    |  |  |       |    |--Exodaenodus Hürzeler 1944 V67
    |  |  |       |    |--Dimyloides Hürzeler 1944 V67
    |  |  |       |    |--Cordylodon von Meyer 1859 V67
    |  |  |       |    |--Pseudocordylodon Hürzeler 1944 V67
    |  |  |       |    `--Metacordylodon Schlosser 1911 V67
    |  |  |       `--Plesiodimylus Gaillard 1897 [Pseudodimylinae] V67
    |  |  `--+--Wyonycteris HUG17
    |  |     |    |--W. chalix HUG17
    |  |     |    `--W. primitivus HUG17
    |  |     `--+--Erinaceomorpha HUG17
    |  |        `--Litocherinae B88
    |  |             |  i. s.: Cedrocherus B88
    |  |             |--Litocherus HUG17
    |  |             |    |--L. lacunatus HUG17
    |  |             |    |--L. notissimus HUG17
    |  |             |    `--L. zygeus HUG17
    |  |             `--+--Litolestes Jepsen 1930 B88, M60
    |  |                |    |--L. ignotus B88
    |  |                |    `--L. notissimus Simpson 1936 M60
    |  |                `--Eolestes B88
    |  `--Soricidae HUG17
    `--+--Solenodontidae [Solenodontinae, Solenodontini] AM02
       |    |--Clinopternodus Clark 1937 [=Clinodon Scott & Jepsen 1936 non Regan 1920] AM02
       |    `--Solenodon Brandt 1833 FS15, M58 [incl. Antillogale Patterson 1962 V67, Atopogale Cabrera 1925 M58]
       |         |  i. s.: S. arredondoi Morgan & Ottenwalder 1993 AC98
       |         |--*S. paradoxus Brandt 1833 M58, FS15, M58 [=Sorex (Solenodon) paradoxus M58]
       |         `--+--S. cubanus Peters 1864 FS15, M58 [=*Atopogale cubana M58; incl. S. poeyanus Barbour 1944 V67]
       |            `--S. marcanoi FS15
       |--Nesophontes Anthony 1916 FS15, M58 [Nesophontidae, Nesophontoidea]
       |    |--*N. edithae Anthony 1916 M58
       |    |--N. hypomicrus Miller 1929 M58
       |    |--N. longirostris Anthony 1919 M58
       |    |--N. major FS15
       |    |--N. micrus Allen 1917 M58
       |    |--N. paramicrus Miller 1929 M58
       |    |--N. submicrus AC98
       |    `--N. zamicrus Miller 1929 M58
       `--Parapternodontidae AM02
            |--Parapternodus Brown & Schankler 1982 HUG17, AM02
            |    `--*P. antiquus Brown & Schankler 1982 AM02
            `--Koniaryctes Robinson & Kron 1998 AM02
                 `--*K. paulus Robinson & Kron 1998 AM02

Eulipotyphla incertae sedis:
  Iconapternodus Tong 1997 AM02
    `--I. qii Tong 1997 AM02
  Ceciliolemur Weigelt 1933 [incl. Microtarsioides Weigelt 1933] S62
  Pontifactor B88
  Deinogalerix Fruedenthal 1973 D07
    `--D. koeingswaldi D07
  Oligoryctes Hough 1956 [Oligoryctidae] AM02
    |--*O. cameronensis Hough 1956 AM02
    `--O. altitalonidus (Clark 1937) [=Apternodus altitalonidus] AM02
  Plesiosoricidae B88
    |--Butselia Quinet & Misonne 1965 B88, V67 [Butselidae]
    |--Pakilestes B88
    |--Plesiosorex Pomel 1848 V67
    |    |--P. germanicus M60
    |    `--P. soricinoides M60
    |--Entomacodon Matthew 1909 V67
    `--Meterix Hall 1929 V67
         `--M. latidens M60

*Type species of generic name indicated

REFERENCES

[AC98] Alcover, J. A., X. Campillo, M. Macias & A. Sans. 1998. Mammal species of the world: additional data on insular mammals. American Museum Novitates 3248: 1–29.

[AM02] Asher, R. J., M. C. McKenna, R. J. Emry, A. R. Tabrum & D. G. Kron. 2002. Morphology and relationships of Apternodus and other extinct, zalambdodont, placental mammals. Bulletin of the American Museum of Natural History 273: 1–117.

[B88] Butler, P. M. 1988. Phylogeny of the insectivores. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 117–141. Clarendon Press: Oxford.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[HUG17] Halliday, T. J. D., P. Upchurch & A. Goswami. 2017. Resolving the relationships of Paleocene placental mammals. Biological Reviews 92 (1): 521–550.

[M58] McDowell, S. B., Jr. 1958. The Greater Antillean insectivores. Bulletin of the American Museum of Natural History 115 (3): 113–214.

[M60] McKenna, M. C. 1960. The Geolabidinae: a new subfamily of early Cenozoic erinaceoid insectivores. University of California Publications in Geological Sciences 37 (2): 131–164.

[S62] Simons, E. L. 1962. A new Eocene primate genus, Cantius, and a revision of some allied European lemuroids. Bulletin of the British Museum (Natural History): Geology 7 (1): 1–36.

[V67] Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135 (5): 217–284.

Blarina

Northern short-tailed shrew Blarina brevicauda, photographed by Jamie McCarthy.

Belongs within: Soricidae.

Blarina, the short-tailed shrews, is a genus found in eastern North America. Members of this genus secrete a neurotoxic venom from submaxillary glands that can cause pain for several days in humans bitten (Nowak 1999). Morphologically, the species are primarily distinguished by size, with the northern short-tailed shrew Blarina brevicauda on average larger than the southern short-tailed shrew B. carolinensis, with Elliot's short-tailed shrew B. hylophaga being intermediate both in size and geographical range. However, substantial overlap in size ranges can occur, and morphological distinction of the species can be difficult (Reilly et al. 2005).

Characters (from Nowak 1999): Head and body length 75-105 mm, tail length 17-30 mm, weight 15-30 g. Upper parts slate-coloured, underparts slightly paler. Body robust; snout pointed; eyes small; ears hidden by fur. 32 teeth, with five unicuspid teeth in upper jaw: the second and third incisors, the canine, a normal premolar and a minute premolar.

<==Blarina Gray 1838 [incl. Anotus Wagner 1855, Mamblarinaus Herrera 1899, Talposorex Pomel 1848] V67
    |--B. brevicauda (Say in James 1823) B75 [=Sorex brevicaudus B75; incl. Galemys micrurus Pomel 1848 G69]
    |    |--B. b. brevicauda GC72
    |    |--B. b. churchi GC72
    |    |--B. b. kirtlandi GC72
    |    |--B. b. minima B75
    |    `--B. b. talpoides GC72
    |--B. carolinensis (Bachman 1837) RM05, B75 [=Sorex carolinensis B75, B. brevicauda carolinensis RM05]
    |    |--B. c. carolinensis RM05
    |    |--B. c. peninsulae RM05
    |    `--B. c. shermani RM05
    |--B. hylophaga [=B. brevicauda hylophaga] RM05
    |    |--B. h. hylophaga RM05
    |    `--B. h. plumbea [=B. brevicauda plumbea] RM05
    `--B. telmalestes GC72

*Type species of generic name indicated

REFERENCES

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[GC72] Genoways, H. H. & J. R. Choate. 1972. A multivariate analysis of systematic relationships among populations of the short-tailed shrew (genus Blarina) in Nebraska. Systematic Zoology 21 (1): 106–116.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

Nowak, R. M. 1999. Walker's Mammals of the World, 6th ed., vol. 1. John Hopkins University Press.

[RM05] Reilly, S. M., R. W. Manning, C. C. Nice & M. R. J. Forstner. 2005. Systematics of isolated populations of short-tailed shrews (Soricidae: Blarina) in Texas. Journal of Mammalogy 86 (5): 887–894.

[V67] Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135 (5): 217–284.

Sorex

Pygmy shrew Sorex minutus, photographed by Pavluvčík.

Belongs within: Soricidae.

Sorex, the long-tailed shrews, is a genus found in most of the temperate and Arctic regions of the Northern Hemisphere, predominately in moist habitats (Nowak 1999). The common shrew Sorex araneus is widespread in northern Europe and northwestern Asia.

Characters (from Nowak 1999): Head and body length 46-100 mm, tail length 25-82 mm, adult weight 2.1-18.0 g. Tail a third to more than half of total length, hairy in the young, naked in old adults. Fur sleek; pelage uni-, bi- or tricoloured, colours varying from tan to black. Body slender; snout long, slender, highly movable, with conspicuous vibrissae. Eyes minute but visible; ears usually projecting slightly above pelage. Five upper unicuspid teeth present, with tips coloured brownish to purplish. Females with three or four pairs of mammae.

<==Sorex Linnaeus 1758 P04 (see below for synonymy)
    |--S. alpinus IT07
    |--S. araneus Linnaeus 1758 KJ08
    |    |--S. a. araneus FK55
    |    `--S. a. tetragonurus Hermann 1780 FK55
    |--S. arcticus B75
    |    |--S. a. arcticus B75
    |    `--S. a. laricorum Jackson 1925 B75
    |--S. arizonae IT07
    |--S. asper IT07
    |--S. bairdii IT07
    |--S. bendirii IT07
    |--S. buchariensis IT07
    |--S. caecutiens IT07
    |--S. cansulus IT07
    |--S. cinereus ANG03
    |    |--S. c. cinereus B75
    |    |--S. c. haydeni Baird 1858 B75
    |    `--S. c. leseurii B75
    |--S. coronatus IT07
    |--S. crassicaudatus T66
    |--S. daphaenodon IT07
    |--S. dispar IT07
    |--S. emarginatus Jackson 1925 MB86
    |--S. excelsus IT07
    |--S. fumeus IT07
    |--S. gaspensis BP87
    |--S. granarius IT07
    |--S. hosonoi Imaizumi 1954 I92
    |    |--S. h. hosonoi I92
    |    `--S. h. shiroumanus Imaizumi 1954 I92
    |--S. isodon IT07
    |--S. jacksoni IT07
    |--S. juncensis BP87
    |--S. kizlovi IT07
    |--S. leucogaster IT07
    |--S. longirostris IT07
    |    |--S. l. longirostris BP87
    |    `--S. l. fischeri BP87
    |--S. lyelli BP87
    |--S. merriami IT07
    |--S. milleri BP87
    |--S. minutissimus I92
    |    |--S. m. minutissimus I92
    |    |--S. m. gracillimus I92
    |    `--S. m. hawkeri Thomas 1906 I92
    |--S. minutus Linnaeus 1766 KJ08
    |--S. mirabilis IT07
    |--S. monticolus IT07
    |--S. nanus IT07
    |--S. oreopolus G69
    |    |--S. o. oreopolus G69
    |    `--S. o. ventralis G69
    |--S. ornatus IT07
    |--S. pacificus IT07
    |--S. palustris ANG03
    |    |--S. p. palustris B75
    |    `--S. p. hygrobadistes Jackson 1926 B75
    |--S. portenkoi IT07
    |--S. preblei IT07
    |--S. pribilofensis BP87
    |--S. pygmaeus T66
    |--S. raddei IT07
    |--S. roboratus IT07
    |--S. sadonis Yoshiyuki & Imaizumi 1986 I92
    |--S. samniticus IT07
    |--S. satunini IT07
    |--S. saussurei G69
    |    |--S. s. saussurei G69
    |    |--S. s. oaxacae G69
    |    `--S. s. veraecrucis G69
    |--S. sclateri BP87
    |--S. shinto IT07
    |--S. sinuosus BP87
    |--S. stizodon BP87
    |--S. sylvaticus CM07
    |--S. tenellus IT07
    |--S. trigonirostris BP87
    |--S. trowbridgii IT07
    |--S. tundrensis IT07
    |--S. ugyunak IT07
    |--S. unguiculatus IT07
    |--S. vagrans IT07
    |--S. veraepacis G69
    |    |--S. v. veraepacis G69
    |    `--S. v. mutabilis G69
    `--S. volnuchini IT07

Inorganic: Sorex minutissimus minilorientalus Okamura 1987 O87
           Sorex minutissimus protominilorientalus Okamura 1987 O87

Sorex Linnaeus 1758 P04 [incl. Atophyrax Merriam 1884 V67, Corsira Gray 1838 V67, Homalurus Schulze 1890 V67, Hydrogale Pomel 1848 non Kaup 1829 V67, Musaraneus Brisson 1762 V67, Neosorex Baird 1858 V67, Otisorex De Kay 1842 V67, Oxyrhin Kaup 1829 V67, Soricidus Altobello 1927 V67]

*Type species of generic name indicated

REFERENCES

[ANG03] Asher, R. J., M. J. Novacek & J. H. Geisler. 2003. Relationships of endemic African mammals and their fossil relatives based on morphological and molecular evidence. Journal of Mammalian Evolution 10 (1–2): 131–194.

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[FK55] Felten, H., & C. König. 1955. Einige Säugetiere aus dem Zentralmassiv, Südfrankreich. Senckenbergiana Biologica 36: 267–269.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[KJ08] Krämer, M. & M. Jentzsch. 2008. Kleinsäuger-Vorkommen aus dem Raum Zeitz—eine vergleichende Studie. Mauritiana 20 (2): 411–427.

[MB86] Matson, J. O. & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.

Nowak, R. M. 1999. Walker's Mammals of the World, 6th ed., vol. 1. John Hopkins University Press.

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.

[P04] Popov, V. V. 2004. Pliocene small mammals (Mammalia, Lipotyphla, Chiroptera, Lagomorpha, Rodentia) from Muselievo (north Bulgaria). Geodiversitas 26 (3): 403–491.

[T66] Tristram, H. B. 1866. Report on the mammals of Palestine. Proceedings of the Zoological Society of London 1866: 84–93.

[V67] Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135 (5): 217–284.

Cryptotis

North American least shrew Cryptotis parva, from The Smithsonian Book of North American Mammals.

Belongs within: Soricidae.

Cryptotis, the small-eared shrews, is a genus of shrews found in southern North America and northern South America. They are mostly found in forests except for the North American least shrew C. parva, which prefers open grassy fields (Nowak 1999). Cryptotis parva is also the only species of the genus found north of Mexico, with its range occupying Mexico and much of the eastern United States.

Characters (from Nowak 1999): Head and body length approximately 55-100 mm, tail length 12-42 mm, adult weight about 4-7 g. Upper parts brownish or blackish, underparts same colour or paler. Snout pointed, eyes minute, ears inconspicuous. 30 teeth, with four upper unicuspids: the second and third incisors, the canine, and a premolar; unicuspid premolar smaller than canine, usually minute.

<==Cryptotis Pomel 1848 [incl. Brachysorex Duvernoy 1842, Soriciscus Coues 1877] V67
    |--C. parva (Say in James 1823) B75 [=Sorex parvus B75; incl. *C. cinereus C57]
    |--C. avius Allen 1923 [=C. avia] C57
    |--C. celatus Goodwin 1956 G69
    |--C. endersi IT07
    |--C. fossor G69
    |--C. frontalis G69
    |--C. goldmani IT07
    |--C. goodwini IT07
    |--C. gracilis IT07
    |--C. hondurensis IT07
    |--C. magna IT07
    |--C. merus Goldman 1912 C57
    |--C. mexicana IT07
    |    |--C. m. mexicana G69
    |    |--C. m. machetes G69
    |    `--C. m. peregrina G69
    |--C. micrura (Tomes 1862) (see below for synonymy) G69
    |--C. montivagus (Anthony 1921) [=Blarina montivaga] C57
    |--C. nigrescens IT07
    |--C. squamipes (Allen 1912) [=Blarina (Cryptotis) squamipes] C57
    |--C. surinamensis (Gmelin 1788) (see below for synonymy) C57
    `--C. thomasi (Merriam 1897) [=Blarina thomasi] C57
         |--C. t. thomasi C57
         |--C. t. equatoris (Thomas 1912) [=Blarina equatoris; incl. B. osgoodi Stone 1916] C57
         |--C. t. medellinius Thomas 1921 C57
         `--C. t. meridensis (Thomas 1898) [=Blarina meridensis] C57

Cryptotis micrura (Tomes 1862) [=Sorex micrurus non Galemys micrurus Pomel 1848 (not preoc. if in dif. gen.), Blarina tropicalis Merriam 1895, Corsira tropicalis Gray 1843 (n. n.)] G69

Cryptotis surinamensis (Gmelin 1788) [=Sorex surinamensis; incl. Blarina pyrrhonota Jentinck 1910, Cryptotis pyrrhonota] C57

*Type species of generic name indicated

REFERENCES

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[C57] Cabrera, A. 1957. Catalogo de los mamiferos de America del Sur. I (Metatheria—Unguiculata—Carnivora). Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Instituto Nacional de Investigacion de Las Ciencias Naturales, Ciencias Zoológicas 4 (1): 1–307.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

Nowak, R. M. 1999. Walker's Mammals of the World, 6th ed., vol. 1. John Hopkins University Press.

[V67] Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135 (5): 217–284.

Talpidae

Shrew mole Uropsilus sp., from here.

Belongs within: Eulipotyphla.
Contains: Talpini.

The Talpidae, moles, are burrowing lipotyphlans found in Eurasia and North America. Moles have a unique digging mode, where the humerus rotates around its long axis. This means that the hands move laterally alongside the body, rather than underneath it as in other animals. As well as the familiar subterranean species, the family also includes the aquatic desmans (Desmaninae) and the shrew-like Uropsilus. Species of Uropsilus, found in eastern Asia, are not strongly specialised for burrowing and have been suggested to occupy a basal position in mole phylogeny. Another contender for this position is the American shrew mole Neurotrichus gibbsii, found in the Pacific northwest of North America, which is also the smallest living member of the family. Talpids are first known in the fossil record by the Late Eocene Eotalpa from the UK; they are not known from North America until the Miocene (Butler 1988).

<==Talpidae [Myaladae, Orycteri, Talpomorpha]
    |--Oreotalpa florissantensis MJ11, HUG17
    |--Eotalpa Sige et al. 1977 MJ11, SM93
    |    `--E. belgica OB13
    `--+--Neurotrichus Günther 1880 FS15, V67
       |    `--N. gibbsii FS15
       `--+--Scaptonyx Milne-Edwards 1872 FS15, P04 [Scaptonychini]
          |    |--S. dolichochir A05
          |    |--S. edwardsi A05
          |    `--S. fusicaudus FS15
          `--+--+--Scapanulus Thomas 1912 FS15, V67
             |  |    `--S. oweni FS15
             |  `--+--Parascalops True 1894 FS15, V67
             |     |    `--P. breweri FS15
             |     `--Scalopinae [Scalopini, Scapanei] MJ11
             |          |--Scapanus Pomel 1848 [=Scapaneus Winge 1917] V67
             |          |    |--S. latimanus FS15
             |          |    `--+--S. orarius FS15
             |          |       `--S. townsendii FS15
             |          `--Scalopus Geoffroy Saint Hilaire 1803 FS15, V67 (see below for synonymy)
             |               `--S. aquaticus FS15
             |                    |--S. a. aquaticus B75
             |                    |--S. a. caryi F68
             |                    |--S. a. machrinoides Jackson 1914 B75
             |                    `--S. a. machrinus B75
             `--Talpinae MJ11
                  |--Urotrichini HO01
                  |    |--Dymecodon True 1886 V67
                  |    |    `--D. pilirostris FS15
                  |    |--Myxomygale Filhol 1890 HO01, V67
                  |    |--Urotrichus Temminck 1841 FS15, V67
                  |    |    |--U. pilirostris IT07
                  |    |    `--U. talpoides FS15
                  |    |--Paratalpa Lavocat 1951 MJ11, V67
                  |    |--Domninoides Green 1956 V67
                  |    |--Mydecodon Wilson 1960 V67
                  |    |--Scalopoides Wilson 1960 V67
                  |    `--Proscapanus Gaillard 1899 [=Proscapaneus Winge 1917] V67
                  |         `--P. sansaniensis A05
                  `--+--Talpini FS15
                     `--+--Desmaninae [Myogalina] B88
                        |    |--Mygalea Schreuder 1940 V67
                        |    |--Mygalinea Schreuder 1940 V67
                        |    |--Gaillardia Matthew 1932 [incl. Hydroscapheus Shotwell 1956; Gaillardiinae] V67
                        |    |--Geomana Brunner 1957 V67
                        |    |--Desmagale Kretzoi 1954 V67
                        |    |--Lemoynea G88
                        |    |--Desmana Gueldenstaedt 1777 FS15, P04 (see below for synonymy)
                        |    |    `--D. moschata FS15
                        |    `--Galemys Kaup 1829 [=Galomys Agassiz 1846; incl. Mygalina Geoffroy Saint Hilaire 1835] V67
                        |         `--G. pyrenaicus FS15
                        `--+--Condylura Illiger 1811 FS15, P04 (see below for synonymy)
                           |    `--C. cristata (Linnaeus 1758) B75
                           `--Uropsilinae HO01
                                |--Asthenoscaptor Hutchinson 1974 HO01
                                |--Mygatalpa Schreuder 1940 HO01, V67
                                |--Desmanella HO01
                                |--Mystipterus Hall 1930 HO01, V67
                                |--Theratiskos Hoek Ostende 2001 HO01
                                |    |--*T. mechteldae Hoek Ostende 2001 HO01
                                |    `--T. rutgeri Hoek Ostende 2001 HO01
                                `--Uropsilus Milne Edwards 1872 FS15, V67 (see below for synonymy)
                                     |--U. andersoni FS15
                                     |--U. gracilis FS15
                                     |--U. investigator FS15
                                     |--U. moschatus HUG17
                                     `--U. soricipes FS15

Talpidae incertae sedis:
  Nesoscaptor uchidai IT07
  Camphotherium Filhol 1884 V67 [=Comphotherium V67, Gomphotherium non Burmeister 1837 M66]
  Galeospalax Pomel 1848 V67
  Hyporyssus Pomel 1848 V67
  Scaptogale Trouessart 1879 [incl. Echinogale Pomel 1848 non Wagner 1841] V67
  Suleimania Hoek Ostende 2001 [Suleimaninae] HO01
    `--*S. ruemkae Hoek Ostende 2001 HO01
  Veratalpa Ameghino 1905 A05
    `--*V. lugdunensiana Ameghino 1905 A05
  Desmanodon HO01

Condylura Illiger 1811 FS15, P04 [incl. Astromycter Harris 1825 V67, Rhinaster Wagler 1830 V67, Talpasorex Schinz 1821 V67; Condylurini]

Desmana Gueldenstaedt 1777 FS15, P04 [=Desman Lacépède 1799 V67, Desmanus Rafinesque 1815 V67; incl. Caprios Wagler 1830 V67, Mygale Cuvier 1800 V67, Myogale Brandt 1836 V67, Myogalea Fischer 1829 V67, Palaeospalax Owen 1846 V67]

Scalopus Geoffroy Saint Hilaire 1803 FS15, V67 [=Scalops Illiger 1811 V67; incl. Hesperoscalops Hibbard 1941 V67, Talpasorex Lesson 1827 non Schinz 1821 V67]

Uropsilus Milne Edwards 1872 FS15, V67 [incl. Nasillus Thomas 1911 V67, Rhynchonax Thomas 1912 V67; Uropsilinae MJ11]

*Type species of generic name indicated

REFERENCES

[A05] Ameghino, F. 1905. La perforation astragalienne sur quelques mammifères du Miocène moyen de France. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 41–58.

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[B88] Butler, P. M. 1988. Phylogeny of the insectivores. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 117–141. Clarendon Press: Oxford.

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[F68] Fisher, D. R. 1968. A study of faunal resemblance using numerical taxonomy and factor analysis. Systematic Zoology 18 (4): 48–63.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[HUG17] Halliday, T. J. D., P. Upchurch & A. Goswami. 2017. Resolving the relationships of Paleocene placental mammals. Biological Reviews 92 (1): 521–550.

[HO01] Hoek Ostende, L. W. van den. 2001. Insectivore faunas from the Lower Miocene of Anatolia—part 5: Talpidae. Zoologica Scripta 122: 1–45.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[M66] McKenna, M. C. 1966. Synopsis of Whitneyan and Arikareean camelid phylogeny. American Museum Novitates 2253: 1–11.

[MJ11] Meredith, R. W., J. E. Janečka, J. Gatesy, O. A. Ryder, C. A. Fisher, E. C. Teeling, A. Goodbla, E. Eizirik, T. L. L. Simão, T. Stadler, D. L. Rabosky, R. L. Honeycutt, J. J. Flynn, C. M. Ingram, C. Steiner, T. L. Williams, T. J. Robinson, A. Burk-Herrick, M. Westerman, N. A. Ayoub, M. S. Springer & W. J. Murphy. 2011. Impacts of the Cretaceous terrestrial revolution and KPg extinction on mammal diversification. Science 334: 521–524.

[OB13] O'Leary, M. A., J. I. Bloch, J. J. Flynn, T. J. Gaudin, A. Giallombardo, N. P. Giannini, S. L. Goldberg, B. P. Kraatz, Z.-X. Luo, J. Meng, X. Ni, M. J. Novacek, F. A. Perini, Z. S. Randall, G. W. Rougier, E. J. Sargis, M. T. Silcox, N. B. Simmons, M. Spaulding, P. M. Velazco, M. Weksler, J. R. Wible & A. L. Cirranello. 2013. The placental mammal ancestor and the post-K-Pg radiation of placentals. Science 339: 662–667.

[P04] Popov, V. V. 2004. Pliocene small mammals (Mammalia, Lipotyphla, Chiroptera, Lagomorpha, Rodentia) from Muselievo (north Bulgaria). Geodiversitas 26 (3): 403–491.

[SM93] Stucky, R. K., & M. C. McKenna. 1993. Mammalia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 739–771. Chapman & Hall: London.

[V67] Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135 (5): 217–284.

Last updated: 9 February 2022.

Soricidae

Water shrew, Neomys fodiens, photographed by J. van der Kooij.

Belongs within: Lipotyphla.
Contains: Sorex, Blarina, Cryptotis, Soriculus, Myosorex, Crocidurinae.

The Soricidae are the shrews, the largest modern 'family' of lipotyphlans. The earliest definite Soricidae are recorded from the middle Eocene of North America, but with confirmed examples from Eurasia shortly thereafter and potential Eurasian species even earlier (Rose 2006). Crown group shrews are characterised by the possession of a double temporo-mandibular joint with neomorphic second lower joint surface and of a deep fossa accomodating the internal temporal muscle opening into the medial side of the coronoid process of the mandible, and the absence of the zygomatic arch, but these characters are not yet present in the stem subfamily Heterosoricinae (Butler 1988, Rose 2006).

The majority of crown-group shrews are divided between the subfamilies Soricinae and Crocidurinae, distinguished by features of the mandibular condyle and of P₄ (Butler 1978). Many species of Soricinae also have the teeth pigmented red by the deposition of iron in a superficial aprismatic layer over the white enamel (Rose 2006). A third subfamily, Myosoricinae, has also been recognised for a group of African genera including the mouse shrews Myosorex. A further extinct subfamily, the Limnoecinae, is known from the Middle Miocene to Middle Pliocence of North America (Van Valen 1967). Within the Soricinae, the living tribes Soricini and Neomyini are each found in both Eurasia and the Americas. The Blarinini are present in the Recent fauna only in the Americas, but are also known from the Pliocene and Pleistocene of Eurasia. The fourth tribe, the extinct Allosoricini, are known from from the Middle Miocence to the late Pliocene of Europe (Van Valen 1967). The water shrews of the genus Neomys hunt aquatic prey, and are found in temperate Eurasia.

See also: Meet the shrews (Taxon of the Week: Soricidae).

Characters (from Rose 2006): Mandible with long, pointed angular process; auditory bulla and zygomatic arch absent. Enlarged anterior incisors, lower incisor procumbent and often with serrated or multilobed crown; remaining antemolar teeth reduced; P₄ primitively unicuspid and triangular; P⁴/M₁ shear developed; dilambdodont molars. Entoconid usually large and separate from hypolophid.

<==Soricidae
    |  i. s.: Arctisorex polaris TH03
    |         Mafia Reumer 1984 P04
    |           `--M. csarnotensis Reumer 1984 P04
    |         Ruwenzorisorex suncoides IT07
    |         Limnoecinae V67
    |           |--Angustidens Repenning 1967 V67
    |           `--Limnoecus Stirton 1930 V67
    |         Podihik Deraniyagala 1958 V67
    |--Heterosoricinae B88
    |    |--Domnina Cope 1873 [incl. Miothen Cope 1873, Protosorex Scott 1895] V67
    |    |    |--*D. gradata Cope 1873 M60
    |    |    |--‘*Miothen’ crassigenus Cope 1873 M60
    |    |    |--‘Protosorex’ crassus M60
    |    |    |--‘Miothen’ gracile Cope 1873 M60
    |    |    `--D. thompsoni Simpson 1941 M58
    |    |--Paradomina Hutchison 1966 V67
    |    |--Ingentisorex Hutchison 1966 V67
    |    `--Trimylus Roger 1885 [incl. Heterosorex Gaillard 1915] V67
    `--+--Soricinae B78
       |    |--Soricini V67
       |    |    |--Sorex MB86
       |    |    |--Crocidosorex Lavocat 1951 [incl. Oligosorex Kretzoi 1959] V67
       |    |    |--Antesorex Repenning 1967 V67
       |    |    |--Drepanosorex Kretzoi 1941 V67
       |    |    |--Microsorex Baird 1877 V67
       |    |    |    `--M. hoyi (Baird 1858) B75 [=Sorex hoyi B75; incl. M. winnemana BP87]
       |    |    |--Alluvisorex Hutchison 1966 V67
       |    |    |--Blarinella Thomas 1911 V67
       |    |    |    |--B. quadraticauda IT07
       |    |    |    `--B. wardi IT07
       |    |    |--Petenyia Kormos 1934 V67
       |    |    `--Zelceina Sulimski 1962 V67
       |    |--Blarinini V67
       |    |    |  i. s.: ‘Sorex’ dehneli Kowalski 1956 V67
       |    |    |--Blarina RM05
       |    |    |--Adeloblarina Repenning 1967 V67
       |    |    |--Cryptotis V67
       |    |    |--Paracryptotis Hibbard 1950 V67
       |    |    |--Shikamainosorex Hasegawa 1957 V67
       |    |    |--Blarinoides Sulimski 1959 V67
       |    |    |    `--B. mariae Sulimski 1959 P04
       |    |    `--Peisorex Kowalski & Li 963 V67
       |    |--Neomyini [Amblycoptinae, Anourosoricinae, Crossopinae, Hydrosoridae, Nectogalinae, Soriculi] V67
       |    |    |--Neomys Kaup 1829 (see below for synonymy) V67
       |    |    |    |--N. anomalus OC03
       |    |    |    |--N. fodiens (Pennant 1771) FK55
       |    |    |    |    |--N. f. fodiens CM07
       |    |    |    |    `--N. f. bicolor CM07
       |    |    |    `--N. schelkovnikovi IT07
       |    |    |--Nectogale Milne-Edwards 1870 V67
       |    |    |    `--N. elegans IT07
       |    |    |--Anourosorex Milne-Edwards 1870 [=Anurosorex Anderson 1875; incl. Pygmura Anderson 1875] V67
       |    |    |    `--A. squamipes M58
       |    |    |--Soriculus V67
       |    |    |--Petenyiella Kretzoi 1956 [incl. Allopachyura Kormos 1934] V67
       |    |    |--Episoriculus Ellerman & Morrison-Scott 1951 V67
       |    |    |--Asoriculus Kretzoi 1959 V67
       |    |    |    `--A. gibberodon (Petényi 1864) (see below for synonymy) P04
       |    |    |--Chodsigoa Kashchenko 1907 V67
       |    |    |--Beremendia Kormos 1934 V67
       |    |    |--Chimarrogale Anderson 1877 [incl. Crossogale Thomas 1921] V67
       |    |    |    |--C. hantu IT07
       |    |    |    |--C. himalayica IT07
       |    |    |    |--C. leander M58
       |    |    |    |--C. phaeura IT07
       |    |    |    |--C. platycephala IT07
       |    |    |    |--C. styani IT07
       |    |    |    `--C. sumatrana IT07
       |    |    |--Amblycoptus Kormos 1926 V67
       |    |    |--Hesperosorex Hibbard 1957 V67
       |    |    |--Notiosorex Baird 1877 V67
       |    |    |    |--N. crawfordi (Coues 1877) MB86
       |    |    |    |    |--N. c. crawfordi MB86
       |    |    |    |    `--N. c. evotis (Coues 1877) MB86
       |    |    |    |--N. gigas G69
       |    |    |    `--N. (Xenosorex) phillipsii Schaldach 1966 G69
       |    |    |--Deinsdorfia Heller 1963 V67
       |    |    |    `--D. kordosi Reumer 1984 P04
       |    |    `--Megasorex Hibbard 1950 V67
       |    |         `--M. gigas IT07
       |    `--Allosoricinae V67
       |         |  i. s.: ‘Sorex’ gracilidens Viret & Zapfe 1951 V67
       |         `--Allosorex Fejfar 1966 V67
       `--+--Myosorex MJ11
          `--Crocidurinae B78

Asoriculus gibberodon (Pétenyi 1864) [=Crocidura gibberodon, Episoriculus gibberodon, Soriculus gibberodon] P04

Neomys Kaup 1829 [incl. Amphisorex Duvernoy 1835, Crossopus Wagler 1832, Hydrogale Kaup 1829, Hydrosorex Duvernoy 1835, Leucorrhynchus Kaup 1829, Myosictis Pomel 1854, Pinalia Gray 1838] V67

*Type species of generic name indicated

REFERENCES

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[B78] Butler, P. M. 1978. Insectivora and Chiroptera. In: Maglio, V. J. & H. B. S. Cooke (eds) Evolution of African Mammals pp. 56–68. Harvard University Press: Cambridge (Massachusetts).

[B88] Butler, P. M. 1988. Phylogeny of the insectivores. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods, vol. 2. Mammals pp. 117–141. Clarendon Press: Oxford.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[FK55] Felten, H., & C. König. 1955. Einige Säugetiere aus dem Zentralmassiv, Südfrankreich. Senckenbergiana Biologica 36: 267–269.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[MB86] Matson, J. O. & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.

[M58] McDowell, S. B., Jr. 1958. The Greater Antillean insectivores. Bulletin of the American Museum of Natural History 115 (3): 113–214.

[M60] McKenna, M. C. 1960. The Geolabidinae: a new subfamily of early Cenozoic erinaceoid insectivores. University of California Publications in Geological Sciences 37 (2): 131–164.

[MJ11] Meredith, R. W., J. E. Janečka, J. Gatesy, O. A. Ryder, C. A. Fisher, E. C. Teeling, A. Goodbla, E. Eizirik, T. L. L. Simão, T. Stadler, D. L. Rabosky, R. L. Honeycutt, J. J. Flynn, C. M. Ingram, C. Steiner, T. L. Williams, T. J. Robinson, A. Burk-Herrick, M. Westerman, N. A. Ayoub, M. S. Springer & W. J. Murphy. 2011. Impacts of the Cretaceous terrestrial revolution and KPg extinction on mammal diversification. Science 334: 521–524.

[OC03] Olson, P. D., T. H. Cribb, V. V. Tkach, R. A. Bray & D. T. J. Littlewood. 2003. Phylogeny and classification of the Digenea (Platyhelminthes: Trematoda). International Journal for Parasitology 33: 733–755.

[P04] Popov, V. V. 2004. Pliocene small mammals (Mammalia, Lipotyphla, Chiroptera, Lagomorpha, Rodentia) from Muselievo (north Bulgaria). Geodiversitas 26 (3): 403–491.

[RM05] Reilly, S. M., R. W. Manning, C. C. Nice & M. R. J. Forstner. 2005. Systematics of isolated populations of short-tailed shrews (Soricidae: Blarina) in Texas. Journal of Mammalogy 86 (5): 887–894.

Rose, K. D. 2006. The Beginning of the Age of Mammals. JHU Press.

[TH03] Tedford, R. H., & C. R. Harington. 2003. An Arctic mammal fauna from the Early Pliocene of North America. Nature 425: 388–390.

[V67] Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135 (5): 217–284.