Original articleNew 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
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 “Felis” vorohuensis 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). “Felis” geoffroyi 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
References (99)
Taphonomy of the Tres Arroyos 1 Rockshelter, Tierra del Fuego, Chile
Quaternary International
(2003)- et al.
Reading the entrails of chickens: molecular timescales of evolution and the illusion of precision
Trends in Genetics
(2004) Middle Pleistocene climate change recorded in fossil mammal teeth from Tarija, Bolivia, and upper limit of the Ensenadan Land-Mammal Age
Quaternary Research
(2000)- et al.
Molecular phylogeny of mitochondrial cytochrome b and 12s rRNA sequences in the Felidae: Ocelot and domestic cat lineages
Molecular Phylogenetics and Evolution
(1996) - et al.
Phylogeny and speciation of felids
Cladistics
(2000) - et al.
The Quaternary impact record from the Pampas, Argentina
Earth and Planetary Science Letters
(2004) - et al.
Changes of mammal assemblages in the Pampean region (Argentina) and their relation with the Plio-Pleistocene boundary
Palaeogeography, Palaeoclimatology, Palaeoecology
(1992) - et al.
Predominance of arid climates indicated by mammals in the pampas of Argentina during the Late Pleistocene and Holocene
Palaeogeography, Palaeoclimatology, Palaeoecology
(1999) - et al.
New radiocarbon chronology for the Guerrero Member of the Luján Formation (Buenos Aires, Argentina): paleoclimatic significance
Quaternary International
(2003) - et al.
The fossil record of the desert-adapted South American rodent Tympanoctomys (Rodentia, Octodontidae). Paleoenvironmental and biogeographic significance
Palaeogeography, Palaeoclimatology, Palaeoecology
(2002)
Paleoclimatic implications of the presence of Clyomys (Rodentia, Echimyidae) in the Pleistocene of Central Argentina
Palaeogeography, Palaeoclimatology, Palaeoecology
La fauna local Quequén Salado-Indio Rico (Pleistoceno tardío) de la provincia de Buenos Aires, Argentina. Aspectos paleoambientales y bioestratigráficos
Ameghiniana
Taphonomic and ecologic information from bone weathering
Paleobiology
Terrestrial vertebrate accumulations
Taphonomy and paleobiology
A new species of small cat (Felidae) from the late Pliocene–early Pleistocene (Uquian) of Argentina
Journal of Mammalogy
Gradual Miocene to Pleistocene uplift of the Central American Isthmus: evidence from tropical American tonnoidean gastropods
Journal of Paleontology
Magnetoestratigrafía de la secuencia cenozoica tardía de Camet (norte de Mar del Plata), Provincia de Buenos Aires. Actas de las 5e Jornadas Geológicas y Geofísicas Bonaerenses
Mar del Plata
Geología y registros magnéticos en la localidad de Camet: correlación con perfiles complementarios de Arroyo La Tapera y Santa Clara del Mar. Actas de las 1e Jornadas de Geología del Cuaternario
Paleomagnetismo y Magnetismo Ambiental, La Plata
Primeros datos geoquímicos y geocronologógicos a partir de algunas cineritas del área bonaerense
Building large trees by combining phylogenetic information: a complete phylogeny of the extant Carnivora (Mammalia)
Biological Reviews
An Analysis of the Natural Classifications of Mammalia for the Use of Students and Travelers
When clocks (and communities) collide: estimating divergence time from molecules and the fossil record
Journal of Paleontology
Catálogo de los Mamíferos de América del Sur. Parte I. Revista del Museo Argentino de Ciencias Naturales
Bernardino Rivadavia
Los félidos vivientes de la República Argentina. Revista del Museo Argentino de Ciencias Naturales
Bernardino Rivadavia
Densidad, biomasa y diversidad de mamíferos en la Puna y Cordillera Frontal
The late Miocene gomphothere Amahuacatherium peruvium (Proboscidea: Gomphotheriidae) from Amazonian Peru: implications for the Great American Faunal Interchange. Boletín del Instituto Geológico Minero y Metalúrgico
Serie D: Estudios Regionales
Pleistocene Mammals of the Cerrado and Caatinga of Brazil
Chronostratigraphy and “land-mammal ages” in the Cenozoic of southern South America: principle, practices, and the “Uquian” problem
Journal of Paleontology
Biostratigraphy and chronological scale of uppermost Cenozoic in the Pampean area, Argentina
Correlation of Pliocene to Holocene southern South America and European vertebrate-bearing units
Bolletino della Società Paleontologica Italiana
A middle-Pleistocene marine transgression in Central-eastern Argentina
Current Research in the Pleistocene
La última glaciación y deglaciación en el Estrecho de Magallanes: implicaciones para el poblamiento de Tierra del Fuego
Anales del Instituto de la Patagonia, Serie Ciencias Humanas
The geologic evidence of the Central American Isthmus
A molecular phylogeny of the Felidae: immunological distance
Evolution
The presence of Neochoerus Hay, 1926 (Rodentia, Hydrochoeridae) in Pleistocenic sediments of Southwestern Buenos Aires Province, Argentina
Quaternary of South America and Antarctic Peninsula
La fauna de vertebrados Pleistocenicos del Bajo San José (Provincia de Buenos Aires). Aspectos Paleoambientales
Ameghiniana
Phylogeographic patterns and evolution of the mitochondrial DNA control region in two Neotropical cats (Mammalia, Felidae)
Journal of Molecular Evolution
Adversaria Zoologica, fasciculus primus
Mémoires de la Société des Naturalistes de Moscou
The Pampas cat group (Genus Ly. Severtzov, 1858) (Carnivora: Felidae), a systematic and biogeographic review
American Museum Novitates
Andean mountain cat, Or. jacobita: morphological description and comparison with other felines from the Altiplano
Journal of Mammalogy
A multivariate comparison of some extant and fossil Felidae
Carnivore
SPA. (S)ankoff (P)arsimony (A)nalysis, version 1.1 (32 bit version). Program and documentation
Análisis arqueofaunístico del Sitio LM1, Llamuco, provincia del Neuquén, República Argentina
The Pleistocene fauna of Piauí (northeastern Brazil). Palaeocological and biochronological implications
Fumdhamentos
La faune Pléistocène de Tarija (Bolivie), note préliminaire
Bulletin du Muséum National d'Histoire Naturelle
Cited by (32)
Felines of the World: Discoveries in Taxonomic Classification and History
2019, Felines of the World: Discoveries in Taxonomic Classification and HistoryQuaternary biostratigraphy and biogeography of mountain region of Córdoba, Argentina
2017, GeobiosCitation Excerpt :Material and provenance: CORD-PZ 4480: complete left mandibular ramus (the crown of the p3 is missing), from a bioturbated dark sandy silt in Pampa de Olaen (FL 5; 31°8′4″ S, 64°34′38.9″ W, 1122 m a.s.l.). Biostratigraphical and geographical distribution: the fossil record of this genus spans from the late Pleistocene (Seymour, 1999; Prevosti, 2006) to Present days (Soibelzon and Prevosti, 2007). Species of this genus are distributed from Southeastern Bolivia, Southern and Western Paraguay, Southern Brasil, Uruguay and most of the Argentinian territory (Seymour, 1999).
Bone modification and destruction patterns of leporid carcasses by Geoffroy's cat (Leopardus geoffroyi): An experimental study
2012, Quaternary InternationalCitation Excerpt :Small felids from South America are diverse with eight known species. The present geographical distribution and fossil record suggest that at least five or six of these species evolved in South America, including Geoffroy’s cat, which last shared a common ancestor with the kodkod (Leopardus guigna) around 2.3 Ma (Nowell and Jackson, 1996; Johnson et al., 1999; Prevosti, 2006). Small cat remains have been found in both archaeological and paleontological sites from the Pampas to Patagonia (Mazzanti and Quintana, 2001; Prado et al., 2001; Martínez and Gutierrez, 2004; Zubimendi et al., 2010).
Comment on "The oldest South American Cricetidae (Rodentia) and Mustelidae (Carnivora): Late Miocene faunal turnover in central Argentina and the Great American Biotic Interchange" by D.H. Verzi and C.I. Montalvo [Palaeogeography, Palaeoclimatology, Palaeoecology 267 (2008) 284-291]
2009, Palaeogeography, Palaeoclimatology, PalaeoecologyLeopardus guigna (Carnivora: Felidae)
2023, Mammalian Species