CidarisRevista Ilicitana de Paleontología y Mineralogía

G R U P O C U L T U R A L P A L E O N T O L Ó G I C O D E E L C H E

Núm. 302010

VIII Encuentro de Jóvenes Investigadores en PaleontologíaVOLUMEN DE ACTAS

Cidaris (2010).30 - VIII EJIP, pág. 169-172 169

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. joan.madurell@icp.cat; david.alba@icp.cat; josep.aurell@icp.cat.

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. salvador.moya@icp.cat.

ABSTRACTFossil 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.

RESUMENFélidos 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

Pumas (also known as cougars or mountain lions) are classifi ed into the species Puma concolor (Linnaeus 1771), which is currently distributed though the American conti-nent. Nonetheless, there are also puma-like fossil remains recorded throughout Eurasia (Hemmer et al., 2004). In the past, they were attributed to Panthera schaubi Viret, 1954, and even a new genus, Viretailurus Hemmer 1964, was erected on their basis. Following Hemmer (2001), however, we attribute them to Puma pardoides (Owen 1846). Unfortunately, the remains of these middle-sized

carnivorans are very scarce, so that their anatomy is poor-ly known. As a consequence, their phylogenetic status and taxonomic attribution remain uncertain.

In this work, we report the mandibular remains of P. pardoides from several Early Pleistocene sites from Spain. Besides providing a detailed comparison with extant pumas, we also compare this fossil material with extant snow leopards, Uncia uncia (Schreber 1775). This is required, given the fact that similarities with the latter species have been noted in some puma-like fossil material from the French locality of Saint Vallier (Olive, 2006).

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Finally, the implications of Eurasian puma-like cats for the understanding of the origins of extant pumas are also discussed from a paleobiogeographic viewpoint.

2. MATERIALS AND METHODSThe remains of P. pardoides reported here were recov-

ered from the Spanish localities of La Puebla de Valverde (Teruel), Cueva Victoria (Murcia) and Vallparadís in Ter-rassa (Barcelona). This material is housed at the Institut Català de Paleontologia (former Institut de Paleontologia M. Crusafont in Sabadell; Barcelona, Spain, acronym IPS), except for the Vallparadís remains (acronym EVT), which are temporarily housed at the Centre de Restau-ració de Béns Mobles de Catalunya in Valldoreix (CRB-MC, Barcelona, Spain).

3. SYSTEMATIC PALEONTOLOGYOrder Carnivora Bowdich 1821

Family Felidae Fischer Von Waldheim 1817

Subfamily Felinae Fischer Von Waldheim 1817

Genus Puma Jardine 1834

Puma pardoides (Owen 1846)

3.1. SYNONYMYFelis pardoides Owen 1846: original description of the species.Panthera schaubi Viret 1954: original description of the species.Viretailurus schaubi (Viret 1954) in Hemmer (1964): original description of the genus.Panthera schaubi Viret 1954 in Kurtén (1976).Felis (Puma) sp. in Sotnikova (1976).Panthera cf. schaubi Viret 1954 in Kurtén and Crusafont (1977).Panthera schaubi Viret 1954 in Turner and Antón (1997).Puma pardoides (Owen 1846) in Hemmer (2001).Felidae indet. (Puma size) in Petrucci (2008).Puma pardoides (Owen 1846) in Madurell-Malapeira et al. (2010).

3.2. REFERRED SPECIMENS

La Puebla de Valverde: Left mandibular corpus with c1, dp3 alveolus, and broken dp4 and m1 (IPS36127). Cueva Victoria: Right mandibular corpus with c1, p3, p4 and m1 (IPS46144). Left p3 (IPS46145). Vallparadís: Partial left mandibular corpus with p4 and broken m1 (EVT4178).

3.3. AGEThe oldest remains reported here are those from the

MN17 locality of La Puebla de Valverde (Kurtén and Cru-

safont-Pairó, 1977). Cueva Victoria has an Early Pleisto-cene age of ca. 1.1 Ma (Blain et al., 2008 and references therein). And fi nally, Vallparadís has a late Early Pleisto-cene age of >0.8 Ma (Alba et al., 2008).

3.4. MORPHOLOGICAL COMPARISONSThe Iberian specimens of P. pardoides from La Puebla

de Valverde, Cueva Victoria and Vallparadís share many dental similarities to the extant P. concolor and the P. par-doides remains from the Late Pliocene of St. Vallier; these include: (1) the location of the three mental foramina; (2) the position of the masseteric fossa, which reaches the level of the m1 protoconid; (3) p3 with a well-developed mesial accessory cusp, more protruding than the distal cusp; (4) p4 with symmetrical protoconid (in lateral view), and well-developed and similarly-sized accessory cusps; and (5) m1 with a protoconid slightly larger and higher than the paraconid, and quite vertically oriented.

The Iberian remains described in this paper also show several similarities with the extant U. uncia: (1) stoutly-built lower mandibular corpus; (2) number and position of the mental foramina; (3) masseteric fossa reaching the level of the m1 protoconid, in lateral view; (4) p3 with a well-developed, circular parastylid that is more protrud-ing than the posterior accessory cuspid; and (5) p4 with a symmetrical protoconid in lateral view and with two well-developed and similarly-sized accessory cuspids. Never-theless, the Iberian remains further differ from U. uncia by m1 protoconid slightly larger and more protruding than the paraconid, with a small cuspid at the base of the protoconid and, in some cases, a lingual bulge between the two main cuspids.

To sum up, the Iberian fossil remains described in this paper display many similarities to the extant P. concolor and the fossil P. pardoides from St. Vallier, while they can be distinguished from U. uncia on dental grounds despite some similarities. This justifi es the attribution of the puma-like remains reported in this paper to P. par-doides.

4. DISCUSSION

4.1. DISTRIBUTION AND CHRONOLOGY

The recorded chronological range of P. pardoides in the Iberian Peninsula, on the basis of the fossil remains reported in this work, spans from the MN17 of La Pueb-la de Valverde up to >0.8 Ma in the case of Vallparadís. Outside of the Iberian Peninsula, P. pardoides is record-ed from the MN16b of Perrier-Étouaires (Hugeney et al., 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-feld (above the base of the Jaramillo chron, ca. 1 Ma) in Germany (Hemmer, 2001); probably Stránská skála (late Early Pleistocene) in the Czech Republic (Hemmer,

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The Eurasian puma-like cat Puma pardoides

2001); possibly from the MN17 locality of Varshets in Bulgaria (Spassov, 2000); the MN16 site of Kvabebi in Georgia (Hemmer et al., 2004); and the Early Villafran-chian localities of Shamar and Beregovaya in Mongo-lia (Sotnikova, 1976), which respectively correspond to the MN16a and MN16b (Vislobokova et al., 1993). Ad-ditional puma-like fossil remains have been described from the localities of Vallonnet (Jaramillo chron, ca. 1 Ma) (Moullé, 1992) in France and Tegelen (MN17) in The Netherlands (Hemmer, 2001. Recently, Petrucci (2008) noted that an ulna and a fi fth metacarpal from the Early Pleistocene locality of Pirro Nord in Italy resemble puma-like cats in both size and morphology.

4.2. PHYLOGENY AND PALEOBIOGEOGRAPHYThe evolutionary origins of the puma lineage are far

from being defi nitively settled. Johnson et al. (2006) ar-gued that the cheetah-like cat Miracinonyx Adams, 1979 and the puma lineage might have shared a last common ancestor in the Middle to Late Pliocene of America, which would imply an American origin of the genus Puma. However, this contention is not supported by the fossil evidence, since P. concolor is only recorded from ca. 0.4 Ma in the American continent (Van Valkenburgh et al., 1990). Alternatively, Eurasian puma-like cat remains (P. pardoides) deserve further consideration regarding the or-igin of the American P. concolor from both a phylogenetic and paleobiogeographic perspectives.

P. pardoides is recorded from Western Europe to cen-tral Asia from the MN16 until the MN17, and hereafter this taxon is not recorded (with the possible exception of

the scarce remains from Pirro Nord) until above the base of the Jaramillo chron in Untermassfeld and several other sites. In the Iberian Peninsula, this species also ranges from the Late Pliocene to the latest Early Pleistocene, thus coinciding with the range of the species elsewhere in Eurasia. The disappearance of P. pardoides from the Eurasian fossil record occurs at the Early/Middle Pleis-tocene transition. This disappearance matches the arrival of leopards into this continent, which became a common element from the late Middle to the Late Pleistocene (Pal-ombo et al., 2008).

The close phylogenetic relationship between the Eur-asian P. pardoides and the extant P. concolor from the American continent, inferred from morphological simi-larities, suggests that the former species must have dis-persed across the Bering Strait during the Middle Pleisto-cene, before its fi rst record in America (Van Valkenburgh et al., 1990). Given the long gap of about 400 kyr between the last European record of puma-like cats and their fi rst American record, it is possible to hypothesize that, be-fore dispersing into America, pumas inhabited northern Asia during the early Middle Pleistocene. A revision of the Asian fossil material would be required in order to de-cipher whether puma-like remains have been incorrectly determined due to their similarities with leopards and snow leopards.

5. SUMMARY AND CONCLUSIONSThe Iberian record of the Eurasian felid Puma pardoides

includes the localities of La Puebla de Valverde (MN17), Cueva Victoria (base of the Jaramillo subchron) and Vall-

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.

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paradís (below the Brunhes-Matuyama boundary). The Iberian specimens share many dental features with those from Saint-Vallier (France), and further closely resemble the extant Puma concolor. Although the material described in this paper shares some features with the snow leopard (Uncia uncia), it can be clearly distinguished from the latter on dental grounds. Recently, there have been some claims that these similarities between Old World puma-like cats and snow leopards might imply a close phylo-genetic relationship. However, the lack of fossil remains from U. uncia precludes an accurate assessment of their affi nities. On the contrary, the puma-like fossil remains reported in this work signifi cantly contribute to our under-standing of the evolutionary history of the puma lineage. First, they show that Eurasian pumas are recorded from the Late Pliocene until the latest Early Pleistocene. More-over, these remains further confi rm that Eurasian puma-like cats (Puma pardoides) are indeed closely related to American pumas (Puma concolor). When taken together, and from a paleobiogeographic viewpoint, this evidence is consistent with an Eurasian origin of the puma lineage.

6. ACKNOWLEDGMENTSFieldwork at Vallparadís was fi nanced by Gestió

d’Infraestructures, S.A., under the supervision of the Gen-eralitat de Catalunya. We thank S. Val, M. Valls, and pre-parators from Origen Conservació i Restauració, S.C.P., for the excellent preparation of the specimens. The col-laboration of the Ajuntament de Terrassa is also acknowl-edged. This work has been possible thanks to the Spanish Ministerio de Ciencia e Innovación (project CGL2008-00325/BTE and Programa Ramón y Cajal RYC-2009-04533 for DMA), the Departament d’Innovació, Univer-sitats i Empresa of the Generalitat de Catalunya (Grup de Paleoprimatologia i Paleontologia Humana, 2009 SGR 754, GRC) and the National Science Foundation (project RHOI-Hominid-NSF-BCS-0321893).

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