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Cladistic Analysis of the Family Rhinocerotidae (Perissodactyla)

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Cladistic Analysis of the Family Rhinocerotidae (Perissodactyla) AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3143, 25 pp., 2 figures, 5 tables August 14, 1995 Cladistic Analysis of the Family Rhinocerotidae (Perissodactyla) ESPERANZA CERDENO' ABSTRACT A cladistic analysis ofthe family Rhinocerotidae otinae and Aceratheriinae are questioned. Within encompassing 45 taxa and 72 characters (referred the former, the elasmotheriines are separated into to craniomandibular [20], dental [26], and post- two groups: a new subtribe, Iranotheriina, is pro- cranial [26] features), and using Hyrachyus and posed, while Elasmotherium and Ninxiatherium two subfamilies of Hyracodontidae as outgroups, appear more closely related to Stephanorhinus and discovered 104 equally parsimonious cladograms. Coelodonta. The content of the subtribe Rhino- When the analysis was run with Hyrachyus as the cerotina is revised. The tribe Teleoceratini is re- only outgroup (removing the hyracodontid sub- moved from the Rhinocerotinae and included families), only six equally parsimonious trees were within the subfamily Aceratheriinae. A new ac- discovered. The discovered trees of the two anal- eratherine tribe, Alicornopini, is proposed for Al- yses are compared, which suggests a reinterpre- icornops, Peraceras, and Chilotheridium. The tation ofthe phylogenetic relationships within the analysis indirectly supports some synonymies pre- Rhinocerotidae. The subfamilies Diceratheriinae viously suggested, such as "Begertherium" = His- and Menoceratinae are not supported, as the gen- panotherium, and "Dicerorhinus" schleiermacheri era included within them appear as a paraphyletic = Lartetotherium schleiermacheri. group. Phylogenies of the subfamilies Rhinocer- INTRODUCTION The investigation offossil rhinocerotids has data on other groups of perissodactyls (e.g., long been considered a difficult task because horses), investigations of fossil rhinoceroses of their great intraspecific variation, as well are relatively scarce, at least at the family as the general homogeneity of the group at a level. Some of the most recent monographic higher taxonomic level. Unlike the body of works on rhinos are those of Guerin (1980), I Fulbright Postdoctoral Scholar, Department of Vertebrate Paleontology, American Museum of Natural History; Departamento de Paleobiologia, Museo Nacional de Ciencias Naturales, Jose Guti6rrez Abascal, 2; 28006 Madrid, Spain. Copyright © American Museum of Natural History 1995 ISSN 0003-0082 / Price $3.40 2 AMERICAN MUSEUM NOVITATES NO. 3143 TABLE 1 Genera and species of Hyracodontidae and Rhinocerotidae Studied, Geographical Distribution AF = Africa; AS = Asia; EU = Europe; NA = North America. HYRACODONTIDAE S. kirchbergensis EU Hyracodon Leidy, 1856 Coelodonta Bronn, 1831 H. nebraskensis NA C. antiquitatis EU/AS H. leydianus NA Hispanotherium Crusafont & Villalta, 1947 Ardynia Matthew & Granger, 1923 H. matritense EU/AS A. praecox AS "Begertherium" Beliajeva, 1971 A. kazachstanensis AS "-B." grimmi AS Prohyracodon Koch, 1897 "Beliajevina" Heissig, 1974 P. meridionale AS "B." tekkayai AS P. orientale EU Iranotherium Ringstrom, 1924 Eggysodon Roman, 1911 L mongoliense AS E. osborni EU I. cf. longirhinus AS RHINOCEROTIDAE Elasmotherium Fischer, 1808 Ronzotherium Aymard, 1886 E. shansiense AS R. fdlholi EU E. lagreli AS Protaceratherium Abel, 1910 Gaindatherium Colbert, 1934 P. albigense EU G. browni AS P. minutum EU G. vidali AS P. platyodon EU Ceratotherium Gray, 1867 P. fahlbuschi EU C. neumayri EU P. mirallesi EU C. simun AF Pleuroceros Roger, 1898 Subhyracodon Brandt, 1878 P. pleuroceros EU S. occidentalis NA Diaceratherium Dietrich, 1931 S. mitis NA D. lemanense EU S. tridactylum NA D. aurelianense EU Diceratherium Marsh, 1875 Brachypotherium Roger, 1904 D. armatum NA B. brachypus EU D. niobrarense NA D. annectens NA Prosantorhinus Heissig, 1974 P. germanicus EU Penetrigonias Tanner & Martin, 1976 P. douvillei EU P. hudsoni NA P. dakotensis NA Alicornops Ginsburg & Guerin, 1979 A. simorrense EU Trigonias Lucas, 1900 A. alfambrense EU T. osborni NA T. wellsi NA Hoploaceratherium Ginsburg & Heissig, 1989 H. tetradactylum EU Amphycaenopus Wood, 1927 A. platycephalus NA Aceratherium Kaup, 1932 A. incisivum EU Menoceras Troxell, 1921 M. arikarense NA "Mesaceratherium" Heissig, 1969 Floridaceras Wood, 1966 "M." gaimersheimensis EU F. whitei NA Lartetotherium Ginsburg, 1974 L. sansaniensis EU "F:AM 95544" Prothero (in press) NA Dicerorhinus Gloger, 1841 Peraceras Cope, 1880 "D." schleiermacheri EU P. superciliosum NA "D." pikermiensis EU/AS P. profectum NA P. hesei NA Stephanorhinus Kretzoi, 1942 Aphelops Cope, 1873 S. miguelkrusafonti EU A. megalodus NA S. megarhinus EU A. malacorhinus NA S. etruscus EU A. mutilus NA S. hemitoechus EU 1995 CERDENO: RHINOCEROTIDAE 3 TABLE 1-(Continued) Teleoceras Hatcher, 1894 Rhinoceros Linnaeus, 1758 T. medicornutum NA R. unicornis AS T. meridianum NA R. sondaicus AS T. major NA Diceros Gray, 1821 T. fossiger NA D. bicornis AF T. proterum NA T. hicksi NA Cerdefno (1989, 1992), Heissig (1989), For- However, duplication of these results is not telius et al. (1993), and Prothero (in press). possible because published data matrices are During the last decade, several cladistic phy- not available in the three cladistic analyses logenetic analyses ofrhinoceroses (sensu lato) noted above. The phylogenetic hypotheses have provided new hypotheses on their re- proposed by these authors are reflected in the lationships (Heissig, 1981, 1989; Groves, current classification of rhinoceroses (Proth- 1983; Prothero et al., 1986). The work of ero and Schoch, 1989). The research reported Prothero et al. (1986) is the most ambitious here is an attempt at a more rigorous phy- attempt; it includes the whole superfamily logenetic analysis of the family Rhinocero- Rhinocerotoidea and provides a good com- tidae. pilation of previous phylogenies of rhinos. MATERIAL AND METHODS Specimens ofmost genera were studied di- bridge); National Museum of Natural History- rectly by the author for more than 10 years. Smithsonian Institution (Washington). The material is housed in the following in- Extant species were reviewed from the col- stitutions: lection in the Department of Mammalogy, AMNH. Table 1 summarizes the species of Spain: Museo Nacional de Ciencias Naturales the genus reviewed by the author and indi- (MNCN), Museo del ITGE, Departamento de cates their geographical distribution. Paleontologia de la Universidad Complutense, Taxa in quotation marks correspond to and Museo Arqueologico Nacional (Madrid); those forms previously considered or sug- Instituto de Paleontologla M. Crusafont (Sa- gested to be synonymous, although a general badell), Museo Paleontologico (Valencia), So- consensus has not been reached: Hispano- ciedad de Ciencias Aranzadi (San Sebasti'an). therium = "Begertherium" and "Beliajevi- Portugal: Departamento de Estratigrafia e Paleo- na," Antunes and Ginsburg (1983)-Cerde-no biologia, Universidade Nova de Lisboa. (1989)-Fortelius and Heissig (1989). "Dicer- France: Institut de Paleontologie, MNHN (Paris; orhinus" schleiermacheri = ?Lartetotherium Departement des Sciences de la Terre de l'Uni- schleiermacheri-Cerdeino (1992). "Mesa- versite Claude Bernard (Lyon). ceratherium gaimersheimensis" = Acerathe- Netherlands: Instituut voor aardwetenschappen, rium paulhiacensus-Bonis (1973), "Mesa- Rijksuniversiteit (Utrecht). ceratherium gaimersheimensis" = Protacera- Germany: Universitlits-Institute fur Paliiontolo- therium gaimersheimensis, author's opinion. gie und historische Geologie (Munich), Senck- One specimen in particular ("F:AM 95544," enberg Museum (Frankfurt). a skull in the Frick collection, AMNH) has Italy: Museo di Geologia e Paleontologia (Flor- not been formally defined taxonomically, al- ence). though Prothero et al. (1986) and Prothero USA: American Museum of Natural History (in press) consider it to be a new genus. For (AMNH) (New York); Museum ofComparative "Begertherium" I have considered the type Zoology (MCZ) (Harvard University, Cam- species B. borrissiaki as well as B. grimmi, 4 AMERICAN MUSEUM NOVITATES NO. 3143 following the suggestion of Fortelius and ACKNOWLEDGMENTS Heissig (1989). "Dicerorhinus" pikermiensis (Geraads, 1988) seems to be a species very I am very grateful to Dr. R. H. Tedford for close to "Dicerorhinus" schleiermacheri, but the facilities he provided at the AMNH. I it is not included in the analysis because I also thank Drs. M. C. McKenna, R. H. Ted- have examined just a few ofits remains, and ford, K. Heissig, C. Guerin, S. Roig, and G. Geraads (1988) did not establish clear skel- C. Gould for the critical reading ofthe manu- etal characteristics for it. Groves (1983) dis- script, and their useful comments. The Ful- cussed the affinities of several Miocene spe- bright scholarship of the author is included cies assigned to the genus Dicerorhinus. Spe- in the Research Project PB91-0082, DGI- cies such as D. leakeyi (Hooijer, 1966), D. CYT, Spain. orientalis (Ringstrom, 1924), or D. piker- miensis (Geraads, 1988) are closely similar CHARACTER ANALYSIS to Lartetotherium sansaniensis and "Dicer- Dental morphology has been classically orhinus" schleiermacheri, and their detailed used as the basis for taxonomical studies, study would probably show their closer re- while postcranial characters have often been lationship with the genus Lartetotherium, as neglected. This analysis is mostly based on already suggested (Groves, 1983; Cerdeino, craniodental features, although
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