A comprehensive phylogeny of extinct and extant Rhizomyinae (Rodentia): evidence for multiple intercontinental dispersals Raquel López-Antoñanzas, Lawrence Flynn, Fabien Knoll To cite this version: Raquel López-Antoñanzas, Lawrence Flynn, Fabien Knoll. A comprehensive phylogeny of extinct and extant Rhizomyinae (Rodentia): evidence for multiple intercontinental dispersals. Cladistics, Wiley, 2013, 29 (3), pp.247 - 273. 10.1111/j.1096-0031.2012.00426.x. hal-01920777 HAL Id: hal-01920777 https://hal.archives-ouvertes.fr/hal-01920777 Submitted on 17 Dec 2020 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. 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Cladistics A comprehensive phylog eny of extinct and extant Rhizomyinae (Rodentia): evidence for multiple intercontinental dispersals Journal:For Cladistics Peer Review Manuscript ID: CLA-12-05-0553.R2 Manuscript Type: Article Date Submitted by the Author: n/a Complete List of Authors: López-Antoñanzas, Raquel; Museo Nacional de Ciencias Naturales-CSIC, Paleobiology Flynn, Lawrence; Harvard University, Peabody Museum Knoll, Fabien; Museo Nacional de Ciencias Naturales-CSIC, Paleobiology Paleontology, Phylogeny, Systematics, Taxonomy, Dispersal < Keywords: Biogeography < Applications Cladistics Page 1 of 68 Cladistics 1 2 3 Abstract 4 5 6 The subfamily Rhizomyinae is known from the Late Oligocene up to the present. 7 8 Today, this group comprises six species, which live in southern Asia and eastern Africa. 9 10 Despite the current moderate diversity of the rhizomyines, they had a greater 11 diversification and wider distribution in the past from Asia, their land of origin, to 12 13 Africa where they entered during the Early Miocene. So far 33 fossil species can be 14 15 referred to this group. A cladistic analysis involving fossil and living species has been 16 carried out. Prokanisamys spp. turned out to be the most basal taxa of the ingroup. This 17 18 analysis calls theFor monophyly Peer of several genera Review into question and allows proposing a 19 20 phylogenetic definition of the tribes Tachyoryctini and Rhizomyini. It also provides 21 information about the origin of the African rhizomyines and allows inferring multiple 22 23 dispersal phenomena from Asia to Africa in Early and Late Miocene times. 24 25 26 Keywords: Mammalia; Systematics; Cladistics; Cenozoic; Eurasia; Africa 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Cladistics Cladistics Page 2 of 68 1 2 3 Table of contents 4 5 Introduction 6 7 Material and methods 8 9 Systematics 10 11 Genus Rhizomys Gray, 1831 12 13 Rhizomys sinensis Gray, 1831 14 15 16 Rhizomys sumatrensis (Raffles, 1821) . 17 18 Rhizomys pruinosusFor Blyth, 1851Peer. Review 19 20 Rhizomys (Brachyrhizomys) shansius (Teilhard de Chardin, 1942) . 21 22 23 Rhizomys (Brachyrhizomys) shajius Flynn, 1993. 24 25 Genus Cannomys Thomas, 1915a. 26 27 Cannomys badius (Hodgson, 1841) . 28 29 30 Genus Tachyoryctes (Rüppell, 1836) . 31 32 Tachyoryctes splendens Rüppell, 1836 . 33 34 Tachyoryctes macrocephalus (Rüppell, 1842) . 35 36 37 Tachyoryctes pliocaenicus Sabatier, 1978 . 38 39 Tachyoryctes konjiti Sabatier, 1982 . 40 41 Tachyoryctes makooka Wesselman, Black et Asnake, 2009 . 42 43 44 Genus Anepsirhizomys Flynn, 1982a. 45 46 Anepsirhizomys opdykei Flynn, 1982a . 47 48 Anepsirhizomys pinjoricus (Hinton, 1933) . 49 50 51 Genus Miorhizomys Flynn, 2009. 52 53 Miorhizomys nagrii (Hinton, 1933) . 54 55 Miorhizomys micrus (Flynn, 1982a) . 56 57 58 Miorhizomys blacki (Flynn, 1982a) . 59 60 Cladistics Page 3 of 68 Cladistics 1 2 3 Miorhizomys pilgrimi (Hinton, 1933) . 4 5 Miorhizomys harii (Prasad, 1968) . 6 7 Miorhizomys tetracharax (Flynn, 1982a) . 8 9 10 Miorhizomys choristos (Flynn, 1982a) . 11 12 Genus Eicooryctes Flynn, 1982 . 13 14 Eicooryctes kaulialensis Flynn, 1982 . 15 16 17 Genus Protachyoryctes Hinton, 1933. 18 For Peer Review 19 Protachyoryctes tatroti Hinton, 1933 . 20 21 22 Genus Rhizomyides Bohlin, 1946. 23 24 Rhizomyides sivalensis (Lydekker, 1884) . 25 26 Rhizomyides carbonnelli (Brandy, 1979) . 27 28 Rhizomyides platytomeus Flynn, Heintz, Sen et Brunet, 1983 . 29 30 31 Rhizomyides mirzadi (Lang et Lavocat, 1968) . 32 33 Rhizomyides punjabiensis (Colbert, 1933) . 34 35 36 Rhizomyides lydekkeri (Hinton, 1933) and Rhizomyides saketiensis (Gupta, Verma et 37 Tewari, 1978) . 38 39 40 Genus Kanisamys Wood, 1937. 41 42 Kanisamys indicus Wood, 1937 . 43 44 45 Kanisamys sivalensis Wood, 1937 . 46 47 Kanisamys nagrii Prasad, 1968 . 48 49 Kanisamys potwarensis Flynn, 1982a . 50 51 52 Genus Nakalimys Flynn et Sabatier, 1984. 53 54 Nakalimys lavocati Flynn et Sabatier, 1984 . 55 56 Genus Pronakalimys Tong et Jaeger, 1993. 57 58 59 60 Cladistics Cladistics Page 4 of 68 1 2 3 Pronakalimys andrewsi Tong et Jaeger, 1993. 4 5 Genus Prokanisamys de Bruijn, Hussain et Leinders, 1981. 6 7 Prokanisamys arifi de Bruijn, Hussain et Leinders, 1981 . 8 9 10 Prokanisamys kowalskii ( Lindsay, 1996 ). 11 12 Prokanisamys benjavuni ( Mein et Ginsburg, 1985 ). 13 14 Prokanisamys major Wessels et de Bruijn, 2001 . 15 16 17 Prokanisamys sp. Wessels, Fejfar, Peláez-Campomanes et de Bruijn, 2003 . 18 For Peer Review 19 Phylogenetic analysis 20 21 22 Previous work 23 24 New analysis 25 26 Discussion 27 28 29 Prokanisamys 30 31 The African Pronakalimys and Nakalimys 32 33 Kanisamys 34 35 36 Protachyoryctes 37 38 Rhizomyides 39 40 “Rhizomyides ” mirzadi and “ R”. punjabiensis 41 42 43 Miorhizomys 44 45 The crown group 46 47 Tribe Tachyoryctini. 48 49 Tribe Rhizomyini. 50 51 52 Conclusion 53 54 Acknowledgements 55 56 57 References 58 59 60 Cladistics Page 5 of 68 Cladistics 1 2 3 4 López-Antoñanzas, R. et al. 1 5 6 7 8 A comprehensive phylogeny of extinct and extant 9 10 Rhizomyinae (Rodentia): evidence for multiple 11 12 intercontinental dispersals 13 14 Formatted: English (U.S.) 15 1 2 1 16 Raquel López-Antoñanzas , Lawrence J. Flynn and Fabien Knoll 17 18 1 For Peer Review 19 Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC, c/ 20 José Gutiérrez Abascal 2, Madrid 28006, Spain. 21 2Peabody Museum of Archaeology and Ethnology, Harvard University, Cambridge, 22 23 MA 02138. E-mails (RLA): [email protected] ; (LJF): [email protected], 24 (FK): [email protected] 25 26 27 28 29 30 Running title:Phylogeny of rhizomyineRhizomyinae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Cladistics Cladistics Page 6 of 68 1 2 3 4 López-Antoñanzas, R. et al. 2 5 6 7 8 Introduction 9 10 11 Living Rhizomyinae are fossorial muroid rodents from southern Asia and eastern 12 Africa (Fig. 1) that belong to the family Spalacidae (Jansa and Weksler, 2004; 13 14 Gogolevskaya et al., 2010 ). Despite their seceretive way of life, they are important to 15 the people who live in the same zones because, like other rodents (e.g., the cane rat 16 17 Thryonomys; López-Antoñanzas et al., 2004), they are agricultural pests, but also can be 18 a source of protein. For Peer Review 19 20 The living Asian forms (bamboo rats ) include Rhizomys (a trispecific genus) and 21 Cannomys (a monospecific genus). There is a single living African genus Tachyoryctes, 22 23 the mole rat, which includes 2 extant species, although that diversity may be greatly 24 underestimated (Musser and Carleton, 2005). No fossil of Cannomys has been found to 25 26 date. However, the Asian Rhizomys (Brachyrhizomys ) and the African Tachyoryctes are 27 known since Late Miocene time. Although the rhizomyines are today moderately 28 29 diversified, this subfamily, which is known from Late Oligocene times, has had a 30 flourishing history as reflected by the finding discovery of over 30 fossil species 31 32 distributed in 11 genera ( Pronakalimys , Nakalimys , Prokanisamys , Kanisamys , 33 Rhizomyides , Protachyoryctes , Tachyoryctes , Eicooryctes , Miorhizomys, 34 Anepsirhizomys Rhizomys Brachyrhizomys 35 and ( )) in Africa, Thailand, Afghanistan, 36 Pakistan, India, and China. 37 The systematic position of the rhizomyines is controversial. Some authors have 38 39 placed them in a distinct Family Rhizomyidae (Miller and Gidley, 1918; Ellermann, 40 1940, 1941; Simpson, 1945; Ellerman and Morrison-Scott, 1951, 1966; Colbert and 41 42 Hooijer, 1953; Flynn 1982a, 1982b, 1990, 1993; Flynn and Guo-qinQi , 1982; Flynn and 43 Sabatier, 1984; Flynn et al., 1983, 1995; Alemseged and Geraads, 2000; Wessels and de 44 45 Bruijn, 2001), including the tachyoryctines with them. Other authors considered the 46 rhizomyines as members (Lydekker, 1885; Palmer, 1897; Colbert, 1935) or a subfamily 47 48 (Thomas, 1896; Tullberg, 1899; Musser and Carleton, 2005; Flynn, 2009) of the 49 Spalacidae. As Winge (1887) originally did when he created the Tribe Rhizomyini, 50 51 some other specialists considered the rhizomyines as a tribe within the Muridae. 52 McKenna and Bell (1997) considered the rhizomyines a subfamily within the Muridae 53 54 and subdivided it into two tribes: the Rhizomyini and the Tachyoryctini. The results of 55 56 57 58 59 60 Cladistics Page 7 of 68 Cladistics 1 2 3 4 López-Antoñanzas, R. et al. 3 5 6 7 recent analyses of mitochondrial and nuclear genes support a monophyletic Spalacinae 8 9 plus Rhizomyinae, with Myospalacinae as sister group (Jansa and Weksler, 2004; 10 Gogolevskaya et al., 2010) and leave little doubt about the inclusion of the rhizomyines 11 12 in the Family Spalacidae.The results of recent analyses of nucleotide sequences support 13 a monophyletic Spalacinae plus Rhizomyinae, with Myospalacinae as sister group 14 15 (Norris et al., 2004; Gogolevskaya et al., 2010; see also Jansa and Weksler, 2004) and 16 leave little doubt about the inclusion of the rhizomyines in the Family Spalacidae.