Myospalacinae Lilljeborg 1866

Wilson, Don E. & Reeder, DeeAnn, 2005, Order Rodentia - Family Spalacidae, Mammal Species of the World: a Taxonomic and Geographic Reference (3 rd Edition), Volume 2, Baltimore: The Johns Hopkins University Press, pp. 907-926 : 907

Myospalacinae Lilljeborg 1866

Myospalacinae Lilljeborg 1866 , Syst. Ofversigt. Gnag. Daggdjuren: 25.

Synonyms: Mesosiphneinae Zheng 1994 ; Myospalacini Lilljeborg 1866 ; Myospalacidae Kretzoi 1961 ; Myotalpinae Miller 1896 ; Prosiphneinae Leroy 1940 ; Siphneinae Gill 1872 ; Siphneidae Zheng 1994 .

Genera: 2 genera with 6 species:

Genus Eospalax G. M. Allen 1938 (3 species)

Genus Myospalax Laxmann 1769 (3 species)

Discussion: All family-group names listed in the synonymy have been used at one time or another, and recently Zheng (1994) preferred to employ Siphneidae . Regardless of preference, Lilljeborg’s (1866) Myospalacinae is the oldest available name, whether used as family or subfamily. Diagnosis, morphological and chromosomal characteristics, distribution, and remarks on habits and habitat are provided by Carleton and Musser (1984). Phylogenetic arrangements of myospalacines were attempted by G. M. Allen (1940) and Leroy (1940) dealing with extant species; by Teilhard de Chardin (1942) studying extant and extinct species; by Kretzoi (1961), who reorganized the group using all named taxa up to 1961; and by Lawrence (1991) and Zheng (1994), who conducted phylogenetic analyses of living and extinct species.

Lawrence (1991) consolidated synapomorphic morphological traits defining Myospalacinae and recognized four species groups: 1) M. psilurus ; 2) M. myospalax , M. epsilanus , M. aspalax , and the extinct M. youngi and M. pseudarmandi (which Zheng, 1994, regarded as an immature youngi ); 3) M. tingi and related species, all extinct; 4) M. fontanieri , M. smithi , and M. rothschildi . Extant species in groups 1 and 2 are traditionally included in the subgenus and genus Myospalax (G. M. Allen, 1940; Corbet, 1978 c; Ellerman, 1941; Kretzoi, 1961, proposed the genus Episiphneus for pseudarmandi and youngi ) and have been arranged in subfamily Myospalacinae by Zheng (1994). Mesosiphneus ( Kretzoi, 1961) , Chardinia , and Youngia ( Zheng, 1994; invalid as no type species was identified) were erected to contain the species that Lawrence recognized in group 3, as well some additional extinct species, and these three genera were placed in the subfamily Mesosiphneinae by Zheng. Species in group 4 are traditionally separated as the subgenus Eospalax of Myospalax (G. M. Allen, 1940; Corbet, 1978 c; Ellerman, 1941, proposed Zokor for the same assemblage), and Zheng listed these species in the subfamily Prosiphneinae , along with the genera Prosiphneus , Myotalpavus , Pliosiphneus , and Allosiphneus .

Zheng’s (1994) classification, reflecting his hypothesis of evolution in the family, recognizes three subfamilies and ten genera, including those already named, all those proposed by Kretzoi (1961), and three new ones. While Zheng’s subfamilial clusters correspond to three of Lawrence’s species groups, and apparently represent different lineages, elevation of each to subfamily level inflates their phylogenetic status relative to other spalacid assemblages. Although we do not employ tribes, we do recognize Myospalax for Lawrence’s groups 1 and 2, the extinct Mesosiphneus for Lawrence’s group 3 (with Chardina and Youngia as synonyms), and Eospalax for her group 4. Zheng’s (1994) Myospalacinae is thus equivalent to Myospalax , his Prosiphneinae to Eospalax and Prosiphneus ( Myotalpavus and Pliosiphneus as synonyms), and his Mesosiphneinae to the extinct species of Mesosiphneus . Whether extant and extinct species are eventually separated into many genera and three subfamilies or some other arrangement, they unquestionably constitute a monophyletic group as indicated by analyses of cranial and dental characters ( Lawrence, 1991; Zheng, 1994).

Phylogenetic position of myospalacines has been differently conveyed in classifications and taxonomic compilations by treating them as a separate family in Muroidea (Pavlinov and Rossilimo, 1987; Reig, 1980; Zheng, 1994), subfamily of Muridae ( Alston, 1876; Carleton and Musser, 1984; Ellerman, 1941; Musser and Carleton, 1993; Thomas, 1896), subfamily of Cricetidae ( Chaline et al., 1977; Corbet, 1978 c; Lindsay, 1994; Pavlinov et al., 1995 a; Pavlinov and Rossilimo, 1998; Simpson, 1945), closely related to arvicolines (G. M. Allen, 1940; Miller, 1896; Milne-Edwards, 1868 -1874), or as a genus or subfamily in Spalacidae ( Miller and Gidley, 1918; Ognev, 1947; Tullberg, 1899). Gambaryan and Gasc (1993) documented the musculoskeletal system and burrowing behavior of Myospalax myospalax , contrasted it with other burrowing rodents, primarily Spalax nehringi , and noted that Myospalax was unrelated to these other burrowers and belonged in its own monotypic family. Phylogenetic analyses of the nuclear gene LCAT ( Michaux and Catzeflis, 2000) and LACT and nuclear vWF in combination ( Michaux et al., 2001 b) nested Myospalax next to Phodopus within a cricetine clade (also Cricetulus and Mesocricetus ), prompting those authors to strongly urge its transfer to Cricetinae as the tribe Myspalacini . Such an arrangement would revitalize Winge’s (1941) classification of zokors, which he included in the "Cricetini." We unsuccessfully tried to locate their voucher and suspect that it was misidentified. Michaux and Catzeflis (2000:283) listed their tissue sample as Myospalax sp. from an unknown locality in Russia, a vast region that could have produced a true hamster or a zokor. The analyses of mitochondrial ( Norris et al., 2004) and nuclear ( Jansa and Weksler, 2004) sequences tellingly unites zokors, based on accurately identified myospalacines, with Spalax , Rhizomys , and Tachyoryctes in a clade distinct from other muroid groups sampled. Furthermore, scrutiny of morphological diagnoses for myospalacines ( Carleton and Musser, 1984; Lawrence, 1991; Ognev, 1947; Tullberg, 1899) and first-hand study of specimens reveal little about extant myospalacines that indicates their close affinity to living hamsters, a conclusion earlier reached by Ognev (1947).

When zokors diverged from the ancestral spalacid stock is so far untraced in the fossil record. Earliest fossils readily identified as myospalacine are about 11 million years old (beginning of late Miocene; Zheng, 1994). Zheng (1993) speculated that zokors derived from middle Miocene Plesiodipus , which some researchers place close to the ancestry of arvicolines (see discussion in Lindsay, 1994). Considering the extreme fossorial adaptations of myospalacines, their evolutionary history must be rooted in some ancestral group existing well before the Miocene, probably in late or middle Oligocene times, a view echoed by Lawrence (1991:279): "myospalacines, fossil and Recent, are a closely related group of species derived from a primitive muroid stock.".