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Madurell-Malapeira E Cidaris Revista Ilicitana de Paleontología y Mineralogía Núm. 30 2010 VIII Encuentro de Jóvenes Investigadores en Paleontología VOLUMEN DE ACTAS GRUPO CULTURAL PALEONTOLÓGICO DE ELCHE THE EURASIAN PUMA-LIKE CAT PUMA PARDOIDES (OWEN 1846) (CARNIVORA, FELIDAE): TAXONOMY, BIOGEOGRAPHY AND DISPERSAL EVENTS EL FÉLIDO EURASIÁTICO PARECIDO AL PUMA PUMA PARDOIDES (OWEN 1846) (CARNIVORA, FELIDAE): TAXONOMÍA, BIOGEOGRAFÍA Y EVENTOS DE DISPERSIÓN Joan Madurell-Malapeira1, David M. Alba1, Salvador Moyà-Solà2 and Josep Aurell-Garrido1 1Institut Català de Paleontologia, Universitat Autònoma de Barcelona, Edifi ci ICP, Campus de la UAB s/n, 08193 Cerdanyola del Vallès, Barcelona, Spain. [email protected]; [email protected]; [email protected]. 2ICREA at Unitat d’Antropologia Biològica (Dept. BABVE) and Institut Català de Paleontologia, Universitat Autònoma de Barcelona, Edifi ci ICP, Campus de la UAB s/n, 08193 Cerdanyola del Vallès, Barcelona, Spain. [email protected]. ABSTRACT Fossil puma-like felids (Puma pardoides) are recorded from several late Pliocene to early Pleistocene Eurasian localities, but the interpretation of the phylogenetic relationships between them and the extant puma (Puma concolor) remains con- troversial. In the past, extinct puma-like cats were included into several genera and species, and a close relationship with both pumas (Puma concolor) and snow leopards (Uncia uncia) has been suggested. In this work, we describe the fossil remains of puma-like cats from the Iberian Peninsula, which cover the whole known chronological distribution of this species in Eurasia. On the basis of morphological comparisons, it is concluded that P. pardoides is closely related to living pumas, which supports a likely Eurasian origin of the puma lineage. Keywords: La Puebla de Valverde, Cueva Victoria, Vallparadís, Villafranchian, puma-like cats. RESUMEN Félidos fósiles parecidos a los pumas (Puma pardoides) han sido registrados en varias localidades euroasiáticas desde el Plioceno superior al Pleistoceno inferior, pero la interpretación de las relaciones fi logenéticas entre ellos y el puma actual (Puma concolor) es aún controvertida. En el pasado, los félidos fósiles parecidos a los pumas han sido clasifi cados en diversos géneros y especies, y se ha sugerido una relación cercana con los pumas (Puma concolor) y los leopardos de las nieves (Uncia uncia). En este trabajo, describimos los restos fósiles de félidos parecidos a los pumas provenientes de la Península Ibérica, los cuales abarcan todo el rango cronológico conocido de esta especie en Eurasia. En base a criterios morfológicos, se concluye que P. pardoides está estrechamente emparentado con los pumas actuales, lo que apoya un probable origen euroasiático del linaje del puma. Palabras clave: La Puebla de Valverde, Cueva Victoria, Vallparadís, Villafranquiense, félidos parecidos a los pumas. 1. INTRODUCTION carnivorans are very scarce, so that their anatomy is poor- ly known. As a consequence, their phylogenetic status and Pumas (also known as cougars or mountain lions) are taxonomic attribution remain uncertain. classifi ed into the species Puma concolor (Linnaeus 1771), which is currently distributed though the American conti- In this work, we report the mandibular remains of nent. Nonetheless, there are also puma-like fossil remains P. pardoides from several Early Pleistocene sites from recorded throughout Eurasia (Hemmer et al., 2004). In Spain. Besides providing a detailed comparison with the past, they were attributed to Panthera schaubi Viret, extant pumas, we also compare this fossil material with 1954, and even a new genus, Viretailurus Hemmer 1964, extant snow leopards, Uncia uncia (Schreber 1775). This was erected on their basis. Following Hemmer (2001), is required, given the fact that similarities with the latter however, we attribute them to Puma pardoides (Owen species have been noted in some puma-like fossil material 1846). Unfortunately, the remains of these middle-sized from the French locality of Saint Vallier (Olive, 2006). Cidaris (2010).30 - VIII EJIP, pág. 169-172 169 Joan Madurell-Malapeira et al. Finally, the implications of Eurasian puma-like cats for safont-Pairó, 1977). Cueva Victoria has an Early Pleisto- the understanding of the origins of extant pumas are also cene age of ca. 1.1 Ma (Blain et al., 2008 and references discussed from a paleobiogeographic viewpoint. therein). And fi nally, Vallparadís has a late Early Pleisto- cene age of >0.8 Ma (Alba et al., 2008). 2. MATERIALS AND METHODS 3.4. MORPHOLOGICAL COMPARISONS The remains of P. pardoides reported here were recov- ered from the Spanish localities of La Puebla de Valverde The Iberian specimens of P. pardoides from La Puebla (Teruel), Cueva Victoria (Murcia) and Vallparadís in Ter- de Valverde, Cueva Victoria and Vallparadís share many rassa (Barcelona). This material is housed at the Institut dental similarities to the extant P. concolor and the P. par- Català de Paleontologia (former Institut de Paleontologia doides remains from the Late Pliocene of St. Vallier; these M. Crusafont in Sabadell; Barcelona, Spain, acronym include: (1) the location of the three mental foramina; (2) IPS), except for the Vallparadís remains (acronym EVT), the position of the masseteric fossa, which reaches the which are temporarily housed at the Centre de Restau- level of the m1 protoconid; (3) p3 with a well-developed ració de Béns Mobles de Catalunya in Valldoreix (CRB- mesial accessory cusp, more protruding than the distal MC, Barcelona, Spain). cusp; (4) p4 with symmetrical protoconid (in lateral view), and well-developed and similarly-sized accessory cusps; 3. SYSTEMATIC PALEONTOLOGY and (5) m1 with a protoconid slightly larger and higher than the paraconid, and quite vertically oriented. Order Carnivora Bowdich 1821 Family Felidae Fischer Von Waldheim 1817 The Iberian remains described in this paper also show several similarities with the extant U. uncia: (1) stoutly- Subfamily Felinae Fischer Von Waldheim 1817 built lower mandibular corpus; (2) number and position Genus Puma Jardine 1834 of the mental foramina; (3) masseteric fossa reaching the level of the m1 protoconid, in lateral view; (4) p3 with a Puma pardoides (Owen 1846) well-developed, circular parastylid that is more protrud- ing than the posterior accessory cuspid; and (5) p4 with a 3.1. SYNONYMY symmetrical protoconid in lateral view and with two well- developed and similarly-sized accessory cuspids. Never- Felis pardoides Owen 1846: original description of the theless, the Iberian remains further differ from U. uncia species. by m1 protoconid slightly larger and more protruding Panthera schaubi Viret 1954: original description of the than the paraconid, with a small cuspid at the base of the species. protoconid and, in some cases, a lingual bulge between Viretailurus schaubi (Viret 1954) in Hemmer (1964): the two main cuspids. original description of the genus. Panthera schaubi Viret 1954 in Kurtén (1976). To sum up, the Iberian fossil remains described in this paper display many similarities to the extant P. concolor Felis (Puma) sp. in Sotnikova (1976). and the fossil P. pardoides from St. Vallier, while they Panthera cf. schaubi Viret 1954 in Kurtén and Crusafont can be distinguished from U. uncia on dental grounds (1977). despite some similarities. This justifi es the attribution of Panthera schaubi Viret 1954 in Turner and Antón the puma-like remains reported in this paper to P. par- (1997). doides. Puma pardoides (Owen 1846) in Hemmer (2001). Felidae indet. (Puma size) in Petrucci (2008). 4. DISCUSSION Puma pardoides (Owen 1846) in Madurell-Malapeira et al. (2010). 4.1. DISTRIBUTION AND CHRONOLOGY 3.2. REFERRED SPECIMENS The recorded chronological range of P. pardoides in the Iberian Peninsula, on the basis of the fossil remains La Puebla de Valverde: Left mandibular corpus with c1, reported in this work, spans from the MN17 of La Pueb- dp3 alveolus, and broken dp4 and m1 (IPS36127). Cueva la de Valverde up to >0.8 Ma in the case of Vallparadís. Victoria: Right mandibular corpus with c1, p3, p4 and m1 Outside of the Iberian Peninsula, P. pardoides is record- (IPS46144). Left p3 (IPS46145). Vallparadís: Partial left ed from the MN16b of Perrier-Étouaires (Hugeney et al., mandibular corpus with p4 and broken m1 (EVT4178). 1989) and the MN17 of Saint-Vallier (Argant, 2004) both in France; the MN16b or MN17 locality of Newborn, Red Crags in Great Britain (Owen, 1846); Untermass- 3.3. AGE feld (above the base of the Jaramillo chron, ca. 1 Ma) The oldest remains reported here are those from the in Germany (Hemmer, 2001); probably Stránská skála MN17 locality of La Puebla de Valverde (Kurtén and Cru- (late Early Pleistocene) in the Czech Republic (Hemmer, 170 Cidaris The Eurasian puma-like cat Puma pardoides Figure 1. Hemimandible IPS46144 from Cueva Victoria: A, buccal view; B, lingual view; C, occlusal view. D, Left hemiman- dible EVT4178 from Vallparadís, in lingual view. E, Left juvenile hemimandible IPS36127 from La Puebla de Valverde, in buccal view. 2001); possibly from the MN17 locality of Varshets in the scarce remains from Pirro Nord) until above the base Bulgaria (Spassov, 2000); the MN16 site of Kvabebi in of the Jaramillo chron in Untermassfeld and several other Georgia (Hemmer et al., 2004); and the Early Villafran- sites. In the Iberian Peninsula, this species also ranges chian localities of Shamar and Beregovaya in Mongo- from the Late Pliocene to the latest Early Pleistocene, lia (Sotnikova, 1976), which respectively correspond to thus coinciding with the range of the species elsewhere the MN16a and MN16b (Vislobokova et al., 1993). Ad- in Eurasia. The disappearance of P. pardoides from the ditional puma-like fossil remains have been described Eurasian fossil record occurs at the Early/Middle Pleis- from the localities of Vallonnet (Jaramillo chron, ca. 1 tocene transition. This disappearance matches the arrival Ma) (Moullé, 1992) in France and Tegelen (MN17) in of leopards into this continent, which became a common The Netherlands (Hemmer, 2001. Recently, Petrucci element from the late Middle to the Late Pleistocene (Pal- (2008) noted that an ulna and a fi fth metacarpal from the ombo et al., 2008).
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