Muridae Illiger 1811

Wilson, Don E. & Reeder, DeeAnn, 2005, Order Rodentia - Family Muridae, Mammal Species of the World: a Taxonomic and Geographic Reference (3 rd Edition), Volume 2, Baltimore: The Johns Hopkins University Press, pp. 1189-1531 : 1189

Muridae Illiger 1811

Muridae Illiger 1811 , Abh. K. Akad. Wiss. Berlin for 1804-11: 46.

Genera: 150 genera with 730 species in 5 subfamilies:

Subfamily Deomyinae Thomas 1888

Genus Acomys I. Geoffroy 1838 (19 species)

Genus Deomys Thomas 1888 (1 species)

Genus Lophuromys Peters 1874 (21 species)

Genus Uranomys Dollman 1909 (1 species)

Subfamily Gerbillinae Gray 1825

Genus Ammodillus Thomas 1904 (1 species)

Genus Brachiones Thomas 1925 (1 species)

Genus Desmodilliscus Wettstein 1916 (1 species)

Genus Desmodillus Thomas and Schwann 1904 (1 species)

Genus Dipodillus Lataste 1881 (13 species)

Genus Gerbilliscus Thomas 1897 (11 species)

Genus Gerbillurus Shortridge 1942 (4 species)

Genus Gerbillus Desmarest 1804 (38 species)

Genus Meriones Illiger 1811 (17 species)

Genus Microdillus Thomas 1910 (1 species)

Genus Pachyuromys Lataste 1880 (1 species)

Genus Psammomys Cretzschmar 1828 (2 species)

Genus Rhombomys Wagner 1841 (1 species)

Genus Sekeetamys Ellerman 1947 (1 species)

Genus Tatera Lataste 1882 (1 species)

Genus Taterillus Thomas 1910 (9 species)

Subfamily Leimacomyinae

Genus Leimacomys Matschie 1893 (1 species)

Subfamily Murinae Illiger 1811

Genus Abditomys Musser 1982 (1 species)

Genus Abeomelomys Menzies 1990 (1 species)

Genus Aethomys Thomas 1915 (9 species)

Genus Anisomys Thomas 1903 (1 species)

Genus Anonymomys Musser 1981 (1 species)

Genus Apodemus Kaup 1829 (20 species)

Genus Apomys Mearns 1905 (9 species)

Genus Archboldomys Musser 1982 (2 species)

Genus Arvicanthis Lesson 1842 (7 species)

Genus Bandicota Gray 1873 (3 species)

Genus Batomys Thomas 1895 (4 species)

Genus Berylmys Ellerman 1947 (4 species)

Genus Bullimus Mearns 1905 (3 species)

Genus Bunomys Thomas 1910 (6 species)

Genus Carpomys Thomas 1895 (2 species)

Genus Chiromyscus Thomas 1925 (1 species)

Genus Chiropodomys Peters 1869 (6 species)

Genus Chiruromys Thomas 1888 (3 species)

Genus Chrotomys Thomas 1895 (4 species)

Genus Coccymys Menzies 1990 (2 species)

Genus Colomys Thomas and Wroughton 1907 (1 species)

Genus Conilurus Ogilby 1838 (2 species)

Genus Coryphomys Schaub 1937 (1 species)

Genus Crateromys Thomas 1895 (4 species)

Genus Cremnomys Wroughton 1912 (2 species)

Genus Crossomys Thomas 1907 (1 species)

Genus Crunomys Thomas 1897 (4 species)

Genus Dacnomys Thomas 1916 (1 species)

Genus Dasymys Peters 1875 (9 species)

Genus Dephomys Thomas 1926 (2 species)

Genus Desmomys Thomas 1910 (2 species)

Genus Diomys Thomas 1917 (1 species)

Genus Diplothrix Thomas 1916 (1 species)

Genus Echiothrix Gray 1867 (2 species)

Genus Eropeplus Miller and Hollister 1921 (1 species)

Genus Golunda Gray 1837 (1 species)

Genus Grammomys Thomas 1915 (12 species)

Genus Hadromys Thomas 1911 (2 species)

Genus Haeromys Thomas 1911 (3 species)

Genus Hapalomys Blyth 1859 (2 species)

Genus Heimyscus Misonne 1969 (1 species)

Genus Hybomys Thomas 1910 (6 species)

Genus Hydromys E. Geoffroy 1804 (5 species)

Genus Hylomyscus Thomas 1926 (8 species)

Genus Hyomys Thomas 1903 (2 species)

Genus Kadarsanomys Musser 1981 (1 species)

Genus Komodomys Musser and Boeadi 1980 (1 species)

Genus Lamottemys F. Petter 1986 (1 species)

Genus Leggadina Thomas 1910 (2 species)

Genus Lemniscomys Trouessart 1881 (11 species)

Genus Lenomys Thomas 1898 (1 species)

Genus Lenothrix Miller 1903 (1 species)

Genus Leopoldamys Ellerman 1947 (6 species)

Genus Leporillus Thomas 1906 (2 species)

Genus Leptomys Thomas 1897 (3 species)

Genus Limnomys Mearns 1905 (2 species)

Genus Lorentzimys Jentink 1911 (1 species)

Genus Macruromys Stein 1933 (2 species)

Genus Madromys Sody 1941 (1 species)

Genus Malacomys Milne-Edwards 1877 (3 species)

Genus Mallomys Thomas 1898 (4 species)

Genus Malpaisomys Hutterer, Lopez-Martinez, and Michaux 1988 (1 species)

Genus Mammelomys Menzies 1996 (2 species)

Genus Margaretamys Musser 1981 (3 species)

Genus Mastacomys Thomas 1882 (1 species)

Genus Mastomys Thomas 1915 (8 species)

Genus Maxomys Sody 1936 (17 species)

Genus Melasmothrix Miller and Hollister 1921 (1 species)

Genus Melomys Thomas 1922 (23 species)

Genus Mesembriomys Palmer 1906 (2 species)

Genus Micaelamys Ellerman 1941 (2 species)

Genus Microhydromys Tate and Archbold 1941 (2 species)

Genus Micromys Dehne 1841 (1 species)

Genus Millardia Thomas 1911 (4 species)

Genus Muriculus Thomas 1902 (1 species)

Genus Mus Linnaeus 1758 (38 species with 5 subspecies)

Genus Mylomys Thomas 1906 (2 species)

Genus Myomyscus Shortridge 1942 (4 species)

Genus Nesokia Gray 1842 (2 species)

Genus Nesoromys Thomas 1922 (1 species)

Genus Nilopegamys Osgood 1928 (1 species)

Genus Niviventer J. T. Marshall, Jr. 1976 (17 species)

Genus Notomys Lesson 1842 (9 species)

Genus Oenomys Thomas 1904 (2 species)

Genus Palawanomys Musser and Newcomb 1983 (1 species)

Genus Papagomys Sody 1941 (2 species)

Genus Parahydromys Poche 1906 (1 species)

Genus Paraleptomys Tate and Archbold 1941 (2 species)

Genus Paramelomys Rümmler 1936 (9 species)

Genus Paruromys Ellerman 1954 (1 species)

Genus Paulamys Musser 1986 (1 species)

Genus Pelomys Peters 1852 (5 species)

Genus Phloeomys Waterhouse 1839 (2 species)

Genus Pithecheir Lesson 1840 (2 species)

Genus Pithecheirops Emmons 1993 (1 species)

Genus Pogonomelomys Rümmler 1936 (2 species)

Genus Pogonomys Milne-Edwards 1877 (5 species)

Genus Praomys Thomas 1915 (16 species)

Genus Protochromys Menzies 1996 (1 species)

Genus Pseudohydromys Rümmler 1934 (4 species)

Genus Pseudomys Gray 1832 (24 species)

Genus Rattus Fischer 1803 (66 species)

Genus Rhabdomys Thomas 1916 (2 species)

Genus Rhagamys Major 1905 (1 species)

Genus Rhynchomys Thomas 1895 (2 species)

Genus Solomys Thomas 1922 (5 species)

Genus Sommeromys Musser and Durden 2002 (1 species)

Genus Spelaeomys Hooijer 1957 (1 species)

Genus Srilankamys Musser 1981 (1 species)

Genus Stenocephalemys Frick 1914 (4 species)

Genus Stochomys Thomas 1926 (1 species)

Genus Sundamys Musser and Newcomb 1983 (3 species)

Genus Taeromys Sody 1941 (7 species)

Genus Tarsomys Mearns 1905 (2 species)

Genus Tateomys Musser 1969 (2 species)

Genus Thallomys Thomas 1920 (4 species)

Genus Thamnomys Thomas 1907 (3 species)

Genus Tokudaia Kuroda 1943 (2 species)

Genus Tryphomys Miller 1910 (1 species)

Genus Uromys Peters 1867 (10 species)

Genus Vandeleuria Gray 1842 (3 species)

Genus Vernaya Anthony 1941 (1 species)

Genus Xenuromys Tate and Archbold 1941 (1 species)

Genus Xeromys Thomas 1889 (1 species)

Genus Zelotomys Osgood 1910 (2 species)

Genus Zyzomys Thomas 1909 (5 species)

Subfamily Otomyinae Thomas 1896

Genus Myotomys Thomas 1918 (2 species)

Genus Otomys F. Cuvier 1824 (19 species)

Genus Parotomys Thomas 1918 (2 species)

Discussion: Phylogenetic analyses of mitochondrial and nuclear gene sequences support the monophyly of Gerbillinae and Murinae ( Adkins et al., 2001; Montgelard et al., 2002; Robinson et al., 1997); Deomyinae , Gerbillinae , and Murinae ( Catzeflis et al., 1995; Conroy and Cook, 1999; Dubois et al., 1999; Hänni et al., 1995; Martin et al., 2000); and those three subfamilies plus Otomyinae ( Jansa and Weksler, 2004; Michaux and Catzeflis, 2000; Michaux et al., 2001 b). Until analyses of molecular data recovered a monophyletic clade consisting of Acomys , Lophuromys , Uranomys , and Deomys (Deomyinae) , the first three genera had been listed in classifications and checklists under Murinae . They share many of the diagnostic cranial traits associated with murines and the defining "upper first molars with three cusps in the anterior two transverse chevrons" and positions of the two lingual cusps relative to the adjacent central cusps ( Carleton and Musser, 1984; Flynn et al., 1985:605). Deomys was either viewed as the sole member of Deomyinae or an unusual dendromurine (see Deomys account). Gerbils have been treated as a separate family within Muroidea or subfamily within Cricetidae (separated from Muridae ) or more encompassing Muridae (see Gerbillinae introduction). Although strongly differentiated in external and cranial characters among muroids, most gerbil species lack longitudinal enamel crests on their molars, as found in cricetids, and resemble murines in this derived trait, which has been the primary basis for postulating a sister-group relationship between Gerbillinae and Murinae ( Flynn et al., 1985; Tong and Jaeger, 1993).

Despite past treatments of otomyines as an independent family ( Roberts, 1951), subfamily of Nesomyidae ( Chaline et al., 1977; Lavocat, 1978) or Cricetidae ( Misonne, 1974) , or in an inclusive Muridae generally comparable to Muroidea ( Carleton and Musser, 1984; Ellerman, 1941; Thomas, 1896), morphological information from fossil and living samples, along with sequences from mitochondrial and nuclear genes, unequivocally link Vlei and Whistling Rats with African murines (see Otomyinae introduction). Including Otomyinae within a restricted Muridae (separate from Cricetidae ) was already formulated by Tullberg (1899), Miller and Gidley (1918), and Simpson (1945). Cranial, dental, and mandibular conformation of otomyines lacks derived traits that link them with any other African clade, living or fossil, and resembles the cranial configuration of arvicanthine genera (especially Arvicanthis , Pelomys , and Mylomys ). Furthermore, there is an impression that enlarged M3s (with four to 10 laminae) and m1s (four to seven laminae), diagnostic of otomyines, are not found elsewhere among murids and are only seen in nonmurid groups ( Misonne, 1969). The species of late Miocene Microtia from the Gargano fissure fillings in southern Italy are typical European Neogene murines, possibly derived from Stephanomys or Apodemus , with enlarged M3s and m1s accommodating four to seven transverse, tubercular and chevron-like laminations ( Freudenthal, 1976; Freudenthal and Suárez, 1999; Jaeger and Hartenberger, 1989; Millien and Jaeger, 2001; Millien-Parra, 2000 b; Zafonte and Masini, 1992). The laminae of otomyines are flat and lack indication of the murine triserial pattern and cuspidation (except in early fossils) but the proliferation of laminae in Microtia and otomyines suggests a deep shared genetic heritage.

We isolate the African Leimacomys in its own subfamily to highlight the enigmatic nature of its mosaic character pattern and uncertain phylogenetic affinities. Usually associated with dendromurines (see Leimacomyinae introduction), its cranial configuration is unlike any genus in that group and instead resembles murines, gerbils, and deomyines such as Lophuromys . Molar occlusal patterns were thought to match those in the dendromurine Steatomys , but in many ways are more similar to gerbillines ( Denys et al., 1995) or extinct myocricetodontines (C. Denys, in litt., 2001). While nesting Leimacomyinae within Muridae seems to reflect morphology, its status as an independent clade, or alternatively as a member of either Deomyinae or Gerbillinae , will require testing with a broader survey of African genera in Muridae and Nesomyidae in which the focus is on morphological traits other than just dentition, and the incorporation of DNA gene sequences.

Evolutionary origin of the Muridae as constituted here may be traced back through extinct myocricetodontines (early Miocene and late Miocene-early Pliocene in Arabian Peninsula, early to late Miocene in Pakistan, middle Miocene of China, middle to late Miocene in North and East Africa, early to middle Miocene of Turkey, and late Miocene-early Pliocene in Spain; Qiu, 2001; Wessels, 1996) to early Miocene cricetids such as Spanocricetodon in Pakistan and and the morphologically similar Notocricetodon of Africa ( de Bruijn et al., 1996). Tong and Jaeger (1993) speculated that myocricetodontines are paraphyletic, producing both gerbils and murines, which may be a sister-group to African dendromurines and cricetomyines; they estimated divergence of gerbillines-murines from dendromurines-cricetomyines at 18 million years before present. Such a relationship is reflected in the phylogenetic analyses of nuclear IRBP gene sequences by Jansa and Weksler (2004) in which those deomyines, gerbillines, and otomyines sampled (here = Muridae ) form a major sister-group to exemplars of nesomyines, dendromurines, cricetomyines, mystromyines, and petromyscines (here = Nesomyidae ) .