Elsevier

Geobios

Volume 39, Issue 5, September–October 2006, Pages 679-694
Geobios

Original article
New material of Pleistocene cats (Carnivora, Felidae) from Southern South America, with comments on biogeography and the fossil recordNouveaux matériels de chats pléistocènes (Carnivora, Felidae) dans le sud de l'Amérique du Sud, commentaires sur la biogéographie et le registre fossile

https://doi.org/10.1016/j.geobios.2005.01.004Get rights and content

Abstract

In this paper the systematic position and age of several Pleistocene cat remains found in southern South American are studied, in an attempt to more fully document the scarce record of the group and clear up their obscure Quaternary history. The fossils are compared with a large sample of recent specimens by means of qualitative and quantitative characters, as well as multivariate methods (discriminant analysis). The age of previous records is restricted using recent chronostratigraphic and biostratigraphic studies. Ly. colocolo is recorded in the late Ensenadan (0.78–0.5 Ma BP) and Bonaerian/Lujanian (0.5 Ma–8.5 Ka BP) ages of the Pampean Region (Argentina) and in the late Pleistocene or Holocene of Tierra del Fuego (Chile). An incomplete hemimandible found in the Bonaerian of the Pampean Region is referred to cf. Herpailurus and could be the earliest record of this lineage. Two other remains could belong to On. geoffroyi, but their incompleteness and some differences prevent their assignation to this recent species. The age of “Felisvorohuensis is restricted to the late Ensenadan. The fossil record of the Ocelot Lineage is very fragmentary, but it is at least as old as late Ensenadan. Taphonomic biases are responsible for this poor fossil record and this fact could partially explain the hiatus with respect to the timing estimated by molecular divergence. The combination of data suggests that Ly. colocolo, On. guigna, On. geoffroyi and Oreailurus jacobita speciated in South America, supporting previous opinions. If the molecular divergence dates are right the recent diversity of this group could be explained by a minimum of five to six immigrations.

Résumé

La position systématique et l'âge de plusieurs restes de chats pléistocènes retrouvés dans le sud de l'Amérique du Sud sont étudiés, dans le but de compléter partiellement le pauvre registre fossile du groupe et d'éclaircir son histoire quaternaire, lacunaire. Les fossiles sont comparés avec un grand échantillon de spécimens récents, en utilisant la morphologie, la morphométrie et des méthodes d'analyse multivariée (Analyse Discriminante). Les âges proposés antérieurement sont rectifiés à l'aide de nouvelles études chronostratigraphiques et biostratigraphiques. Ly. colocolo est signalé dans l'Ensénadéen tardif (0,78–0,5 Ma AP) et le Bonaëréen/Lujanéen (0,5 Ma–8,5 Ka AP) de la Région Pampéenne (Argentine) et dans le Pléistocène tardif ou l'Holocène de la Terre de Feu (Chili). Une mandibule incomplète retrouvée dans le Bonaëréen de la Région Pampéenne est assignée à cf. Herpailurus et pourrait être le fossile le plus ancien de cette lignée. Deux autres restes pourraient correspondre à On. geoffroyi, mais leur incomplétude et quelques différences avec le matériel type empêchent de les assigner à cette espèce récente. L'âge de « Felis » vorohuensis est restreint à l'Ensénadéen tardif. Le registre fossile de la lignée de l'ocelot est très fragmentaire et commence certainement dans l'Ensénadéen tardif. Plusieurs biais taphonomiques sont responsables de la pauvreté du registre fossile et pourraient expliquer partiellement l'écart avec les âges proposés par divergence moléculaire. Les résultats suggèrent que Ly. colocolo, On. guigna, On. geoffroyi et Oreailurus jacobita ont leur origine en Amérique du Sud, ce qui étaye les opinions antérieures. Si les dates de la divergence moléculaire sont correctes, la diversité récente de ce groupe peut être expliquée par un minimum de 5–6 immigrations.

Introduction

The recent small felids of South America are diverse with eight species known (Wozencraft, 1993). Several species (e.g. Le. pardalis, Le. wiedii, On. tigrina; see Cabrera, 1958, Cabrera, 1961; Nowell and Jackson, 1996) inhabit forest and scrublands of the northern part of South America (i.e. northern Argentina and Uruguay, Bolivia, Brazil and northern countries), as well as Central America and southern North America (de Oliveira, 1997, de Oliveira, 1998a, de Oliveira, 1998b). Other cats (On. geoffroyi, Ly. colocolo) are found mainly in open environments of the southern part of South America (i.e. Uruguay, Chile, Argentina; Cabrera, 1961; Garcia Perea, 1994), whereas Or. jacobita and On. guigna are restricted to the high mountains of the Andes (Argentina, Chile, Bolivia and Peru) and to the Patagonian forest of Chile and Argentina, respectively (Nowell and Jackson, 1996; Yensen and Seymour, 2000; Quintana et al., 2000). The remaining species, Herpailurus yagouaroundi, is widely distributed from northern Patagonia to southern North America (De Oliveira, 1998b).

The Neotropical cat species are a monophyletic group (the Ocelot Lineage; e.g. Collier and O'Brien, 1985; Pecon Slattery et al., 1994), with the exception of H. yagouaroundi that is more closely related to Puma and other felids (Salles, 1992; Eizirik et al., 1998; Johnson et al., 1999; Seymour, 1999; Mattern and Mclennan, 2000). The time of divergence of this clade was calculated at 3–4 Ma BP (Wayne et al., 1989; Bininda Emmons et al., 1999) or 5–6 Ma BP (Pecon Slattery et al., 1994; Pecon Slattery and O'Brien, 1998; Johnson et al., 1999). The first date is coincident with the rise of the Panama bridge at 2.8–3.1 Ma AP (Coates and Obando, 1996), which connects Central and South America, but the second one predates this event.

Contrasting with their rich recent diversity and their early age of divergence and radiation as estimated by molecular methods, the fossil record of this group is scarce and relatively young. The oldest records correspond to “Felisvorohuensis and “Felis” sp. from the Ensenadan (late Pliocene to early middle Pleistocene, sensu Cione and Tonni, 1999), and “H. yagouaroundi” from the “Ensenadan” of Bolivia (Hoffstetter, 1963, Hoffstetter, 1986; Berta, 1983; Deschamps and Borromei, 1992; Berman, 1994; but see below). On. tigrina, H. yagouaroundi, Le. pardalis, Le. wiedii, have been found at paleontological sites in Brazil (late Pleistocene–Holocene; (Winge, 1895; Lund, 1950; Guérin et al., 1996; Lessa et al., 1998; Seymour, 1999). Seymour (1999) described several bones of Le. cf. Le. wiedii and of Oncifelis sp. or Leopardus sp. from the latest Pleistocene (≈ 13 Ka BP, see Marshall et al., 1984) site of Talara (Peru). Le. wiedii amnicola, Le. pardalis and Leopardus sp. reached southern North America during the last interglacial and glacial periods (Sangamon and Wisconsin, respectively; Werdelin, 1985; Hulbert and Pratt, 1998). “Felisgeoffroyi was mentioned for the late Pleistocene of the Buenos Aires province (Argentina; Berman, 1994; but see below) and “Felis lujanensis” for the Lujanian of Buenos Aires province. The latter may be a synonym of Ly. colocolo, but the holotype has been lost and the original description is confusing (Seymour, 1999). Fossils of Or. jacobita and On. guigna have not yet been found. Several specimens of “Felis” sp. have been mentioned in faunal lists of Pleistocene sites of Argentina (e.g. Deschamps and Borromei, 1992; Tonni and Scillato-Yané, 1997; Fig. 1), but without descriptions or figures. This situation is more common in the Holocene archeological and paleontological faunal lists (e.g. Goñi et al., 1996; Quintana, 2001; Tonni et al., 2002).

The objectives of this work are, the description of new and undescribed specimens of Pleistocene cats from the southern cone of South America, and a review the systematic status of other previously published specimens, using “classic” systematic methods, discriminant analysis and assessing the intraspecific variation of qualitative characters in the recent species. Second, the age of previous records is restricted with the aid of new paleomagnetic and biostratigraphic data (e.g. Bidegain et al., 1998, Bidegain et al., 2003; Cione and Tonni, 1999, Cione and Tonni, 2001). Finally, a discussion of the history of the Ocelot Lineage in the light of fossil and molecular evidences is included.

Section snippets

Materials and methods

The specimens described here were compared with recent and fossil individuals from several institutions (see Institutional Abbreviations). As the size of these specimens is small than that of Le. pardalis, and their morphology is clearly different (see Seymour, 1999: 227), only the smaller Neotropical cats (i.e. Or. jacobita, Le. wiedii, On. tigrina, On. geoffroyi, On. guigna, Ly. colocolo, H. yagouaroundi) and Felis catus were used for comparison. Of the recent specimens, only those with fully

Systematics

Order CARNIVORA Bowdich, 1821.

Family FELIDAE Fischer, 1817.

Subfamily FELINAE Fischer, 1817.

Genus Lynchailurus Severtzow, 1858.

Lynchailurus colocolo (Molina, 1782)

Material examined: MLP 90-XII-10-1 (Fig. 2(A, B) and Table 1): left fragmentary maxilla with the C and P3-P4; CI S/N° (cast MLP 01-V-20-1; Fig. 2(C, D), Table 1): left incomplete mandibular ramus with i2-p4 and a incomplete m1; IPUM 6872 (Fig. 2(E, F), Table 1): right hemimandible with p3-m1.

Geographic and stratigraphic distribution:

Fossil record, molecular divergence dates and the small felid immigrations to South America

The osteological, soft anatomy, chromosomal, and molecular evidence shows that the Ocelot Lineage is a monophyletic group (Collier and O'Brien, 1985; Herrington, 1986; Modi and O'Brien, 1988; Pecon Slattery et al., 1994; Johnson et al., 1996, Johnson et al., 1998, Johnson et al., 1999; Masuda et al., 1996; Pecon Slattery and O'Brien, 1998; Eizirik et al., 1998; Seymour, 1999; Mattern and McLennan, 2000), and several authors have suggested that this radiation occurred in Central and South

Conclusions

The C/I S/N°, the oldest record of Ly. colocolo, was found in late Ensenadan beds (0.78–ca. 0.5 Ma BP) in the Pampean region (Argentina). Other records of this species have occurred in the Bonaerian or Lujanian of the region. I confirm the presence of Ly. colocolo in Tierra del Fuego, an island not inhabited by any felids at present, but the age of this fossil could be Holocene or late Pleistocene due the bioturbation of the site.

If the determination of UNSGH 464 as Herpailurus is confirmed,

Acknowledgements

To Erika Hingst Zaher and Hussam Zaher for their help during my visit to Brazil. The AMNH Study Collection Grant allowed me to study the specimens of recent species deposited in this museum. To the collection curators and other people for their collaboration and the loan of recent specimens: C. De Muizon, M. Merino, O. Vaccaro, G. Musser, A. Dondas, S. Bargo, M. Reguero, P. Cardenas, D. Voglino, L. Soler, J. Pereyra, D. Flores, A. Rodríguez, D. Ibañez, R. Tedford and R. MacPhee. To Analía

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