June 2009 Notes 217 PANTHERA LEO ATROX (MAMMALIA: CARNIVORA: FELIDAE) IN CHIAPAS, MEXICO MARISOL MONTELLANO-BALLESTEROS* AND GERARDO CARBOT-CHANONA Departamento de Paleontologı́a, Instituto de Geologı́a, Universidad Nacional Autónoma de México, Circuito Exterior, Ciudad Universitaria, Deleg. Coyoacán, 04510 México, D.F. (MM-B) Museo de Paleontologı́a Eliseo Palacios Aguilera, Instituto de Historia Natural, Calzada de Los Hombres Ilustres s/n, Parque Madero 29000, Tuxtla Gutiérrez, Chiapas, México (GC-C) *Correspondent: marmont@servidor.unam.mx ABSTRACT—A well-preserved right lower jaw with complete P3-M1 and an isolated canine of a large felid was found in the southern state of Chiapas, Mexico. It is identified as the American lion Panthera leo atrox. Its presence in Chiapas documents its southernmost distribution in North America. RESUMEN—Una mandı́bula derecha bien conservada de un gran félido con P3-M1 y un canino aislado fueron encontrados en el estado sureño de Chiapas, México. Fue identificado como león americano Panthera leo atrox. Su presencia en Chiapas documenta la distribución más al sur del continente americano. In 1950, Eliseo Palacios-Aguilera first reported the mammalian fossil remains of edentates and proboscideans recovered from the Quaternary deposits of Villaflores and Villa Corzo, these municipalities are located in central Chiapas. Recovery of fossils was sporadic until 2004, when the staff of the Museo de Paleontologı́a del Instituto de Historia Natural started a systematic study in those areas. As a result of paleontological field surveys, remains of Mammuthus, Cuvieronius, Eremotherium, Glyptotherium, Equus, Odocoileus, and Bison have been recovered (CarbotChanona et al., 2004; Carbot-Chanona and Vázquez-Bautista, 2006; Montellano-Ballesteros and Carbot-Chanona, in press). One of the most important fossil specimens collected is a fragmentary lower jaw of a large felid that is the object of this note. Taxonomic history of the genus and species is complex, Leidy (1853) originally described Felis atrox. In 1873, he described Felis imperialis as a smaller separate species. However, Kurtén and Anderson (1980) synonymized F. imperialis with P. leo atrox. Merriam (1909) and Frick (1930) described the subspecies F. atrox bebbi and F. atrox alaskensis, respectively. With the detailed study of Merriam and Stock (1932) on the fossil-felid material from Rancho La Brea, California, it has been possible to examine morphological variation within the population. They synonymized F. THE SOUTHWESTERN NATURALIST 54(2):217–222 imperialis with F. atrox and considered as nomina dubia the proposed subspecies bebbi and alaskensis. Simpson (1941) moved F. atrox to the genus Panthera and suggested that P. atrox was not a lion, but might be a giant jaguar. He considered it to be distinct from P. onca and named it P. jaguarius atrox. Currently, the genus Felis includes small felids, while Panthera includes large cats such as lions, tigers, and leopards (Turner and Antón, 1997). The abbreviations used in the paper are: IHNFG, Instituto de Historia Natural, Fósil Geográfico, Tuxtla Gutiérrez, Chiapas; C, canine; M, molar; P, premolar (measurements are in millimeters). Class Mammalia Linnaeus, 1758 Order Carnivora Bowdich, 1821 Family Felidae Gray, 1821 Subfamily Pantherinae Pockock, 1917 Genus Panthera Oken, 1816 Panthera leo atrox Leidy, 1853 Type: Felis atrox Leidy, 1853:319. Left lower jaw with broken C and P3-M1 from Pleistocene at Natchez, Mississippi. Felis imperialis Leidy, 1873:259. Type: upper jaw with P3, from Pleistocene gravels at Livermore Valley, California. Felis imperialis Matthew, 1902:321. Felis atrox bebbi Merriam, 1909:301. Felis atrox Freudenberg, 1910:33. Felis imperialis Freudenberg, 1910:31. 218 The Southwestern Naturalist vol. 54, no. 2 FIG. 1—Views of the partial right lower jaw of Panthera leo atrox with P3-M1 (IHNFG-2678): a) lingual; b) labial; c) occlusal; an isolated canine d) outer, e) inner. Referred Specimen—The specimen is a partial right lower jaw with complete P3-M1 and an isolated canine that probably belongs to the same individual (IHNFG-2678; Fig. 1). Locality and Age—The fossil specimen was from La Tejerı́a, which is a quarry for construction materials. This site is near the town of San Pedro Buenavista, in the Municipio of Villa Corzo, June 2009 Notes 219 TABLE 1—Comparative measurements (mm) of lower dentitions of Panthera leo atrox from Mexico (Chiapas and Tequixquiac) and Rancho La Brea, California. Character Chiapasa IHNFG-2678 Length of canine Width of canine Length of P3 Width of P3 Length of P4 Width of P4 Length of M1 Width of M1 21.9 15.6 17.5 9.7 26.2 12.5 26.7 13.2 a b c Tequixquiacb 102 — — — — 23.5–24 — 24–25 — Rancho La Brea, Californiac 21.8–30.4 15.1–22.0 17.0–21.6 8.9–13.2 25.8–32.3 12.0–16.9 26.9–33.9 13.0–17.5 Herein. Freudenberg (1910). Merriam and Stock (1932). Chiapas. The fossil was collected by Misael Nanduca, owner of the La Tejerı́a, from a 20-cm thick layer of a dark, fine-grained, sandy clay. The Pleistocene age is suggested based on presence of an unknown species of Glyptotherium sp., Equus conversidens, and probably Cuvieronius tropicus. Description and Comparisons—The canine has the occusal end broken. P3 is double rooted with the posterior root more robust. Tooth crown bears a distinct small anterior and a slightly larger posterior basal tubercle. Apex of the main cuspid is slightly posterior to the middle of the cuspid. Posterior edge of the main cuspid is sharp. Talonid of P3 is small with a shallow basin. Merriam and Stock (1932) mentioned that the anterior basal tubercle is present in most P3s of FIG. 2—Measurements (mm) taken from specimen IHNFG-2678: L, length; W, width; P, premolar; M, molar; C, canine. Panthera leo atrox from Rancho La Brea. P4 shows anterior and basal tubercles of nearly equal size, separated from the main cusp by a notch. Posterior tubercle is divided in the type specimen (Merriam and Stock, 1932:figure 133). Principal cusp of P4 is higher than long and the anterior slope is longer than the posterior slope; in the lingual side of the basal posterior tubercle, there is a shallow basin. This tooth is bordered posterolingually by a distinctive cingulum. M1 also is double rooted, with talonid exhibiting a cingulum as in the material from Rancho La Brea. Protoconid is slightly longer anteroposteriorly and higher than the paraconid. These cuspids meet medially to form a well-defined carnassial notch. In comparison to the material from Rancho La Brea, California, no significant morphological difference was noted. Measurements are in Table 1 and Fig. 2; size of the specimen from Chiapas is in the size range of the sample from Rancho La Brea. Geographic Distribution of Panthera leo atrox in Mexico—The American lion, P. l. atrox is distinguished easily from other Pleistocene felines by its great size and relatively long and slender bones (Kurtén and Anderson, 1980). It was widely distributed in Canada and United States, reaching Mexico during Rancholabrean times; however, it is not known where it occurred in Mexico (Bell et al., 2004). Kurtén and Anderson (1980) mentioned its presence in Peru, but Seymour (1983, in Yamaguchi et al., 2004) reinterpreted the material as a jaguar (P. onca) of large size. Freudenberg (1910) in his work on the carnivores of the Plio-Pleistocene Mexican faunas, 220 The Southwestern Naturalist referred a fragmented skull with I3, C, P3-P4 as Felis imperialis; a mandible with P4-M1 to Felis cf., imperialis and an upper-jaw fragment without teeth to Felix (5Panthera) atrox. Simpson (1941) suggested that some of the material described by Freudenberg (1910) as Felis cf., imperialis was a jaguar (P. onca), and that the rest belonged to a small F. atrox or to a large extinct Mexican race of P. onca. Simpson (1941) could not find reliable differences between P. onca and P. atrox in the preserved elements. Unfortunately, all fossil material described by Freudenberg is lost and, therefore, cannot be restudied. Miller (1943) and later, Arroyo-Cabrales and Polaco (2003) mentioned Felis (Panthera) atrox as part of the megafauna collected in the San Josecito Cave, Nuevo León, but neither of them mentioned material that was collected. Aviña (1969) reported a P2 from Villa Corzo, Chiapas, which was identified by Alvarez as F. (P.) atrox. The illustrated tooth in Aviña does not correspond to a P2 but to a P3, and the catalog number is different from that labeled in the figure. Compared to the P3 figured in Merriam and Stock (1932:plate 33:figures 2 and 2a) and size of material from California, this specimen from Chiapas is similar to F. (P.) atrox. In the El Cedazo local fauna, Aguascalientes, Mooser and Dalquest, (1975) referred a radius to this cat as F. (P.) atrox because of its large size. Lorenzo and Mirambell (1981) mentioned this felid at Rancho La Amapola, San Luis Potosı́. Rufolo (1998) questionably referred an incomplete distal end of a humerus of a juvenile to ?P. leo atrox, which was in the collection from Lago de Chapala housed at the Los Angeles County Museum. From information presented above, it is evident that the current record of this felid is scarce. The record at La Tejerı́a confirms the southernmost record of this big cat in North America. It is absent in local faunas described for Central America (Cisneros, 2005). Systematic History of Panthera leo atrox—Comparative morphological analysis of Pleistocene and Holocene lions of different geographic populations suggests existence of two evolutionary lines: the spelaea group (Europe) and the leo group of Africa and southern Asia. Some authors favor the taxonomic combination of these groups within Panthera leo (Kurtén, 1965; Turner and Antón, 1997) and others prefer a taxonomic separation at the species level in P. spelaea and P. leo (Baryshnikov and Boeskorov, vol. 54, no. 2 2001; Burger et al., 2004). In their attempt to reconstruct the phylogenetic position of P. leo spelaea (Pleistocene European cave lion), Burger et al. (2004) used mitochondrial cytochrome-b genes and suggested that this extinct lion is the sister clade of extant lions, and that this feline lineage was isolated from the African and Asian lions since their dispersal over Europe ca. 600,000 years ago. They concluded that the question of whether extinct cave lions of Europe and extant lions of Africa and Asia should be recognized as different species may be a matter of convention. The same controversy exists with the Pleistocene American lion; some authors considered P. atrox as a subspecies of the African lion P. leo (e.g., Kurtén and Anderson, 1980; Anderson, 1989; Turner and Antón, 1997); others considered that it is distinct from P. leo spelaea, which lived in Eurasia during the Pleistocene (e.g., Simpson, 1941; Whitmore and Foster, 1967; Martin and Gilbert, 1978; Wheeler and Jefferson, 2003). There is no molecular study dealing with the Pleistocene American lion that would provide clarification of lineages. The fossil record of felids in Mexico is scarce, so geographic and temporal distribution and paleodiversity are not possible to determine at present. This attests to the geographical importance of the find. The fossil material described here is identified as P. l. atrox, which indicates that the most southern distribution documented for this lion in the Pleistocene was Chiapas. Its assignment as a subspecies of P. leo follows the idea that sometime during the Pleistocene, the species P. leo dispersed from Africa, invading Europe and Asia where P. l. spaleae, P. l. vereshchagini, and P. l. fossilis are known. It later crossed Beringia and reached North America, living there in open habitats. We thank M. Nanducá, who kindly donated fossil material described in this paper, and the staff of the Museo de Paleontologı́a, Instituto de Historia Natural (E. Ovalles-Damián, D. Vázquez-Bautista, M. A. Coutiño-José and A. S. Núñez-Vera) who participated in the field work. 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