Paleontología Mexicana

First record of the collared peccary Dicotyles tajacu (Artiodactyla, Tayassuidae), in the Gliptodonte locality, Villaflores municipality, Chiapas

Primer registro del pecarí de collar Dicotyles tajacu (Artiodactyla, Tayassuidae), en la localidad de Gliptodonte, municipio de Villaflores, Chiapas.

Carbot-Chanona, Gerardo1,*; Gómez-Pérez, Luis Enrique1

1 Museo de Paleontología “Eliseo Palacios Aguilera”, Departamento de Paleontología. Dirección de Gestión, Investigación y Educación Ambiental, Secretaría de Medio Ambiente e Historia Natural. Calzada de Los Hombres Ilustres s/n, Tuxtla Gutiérrez, Chiapas, 29000, México.

*gfcarbot@gmail.com

Abstract

Tayassuidae (peccaries) is a family of artiodactyls exclusively of America, that was widely distributed in North America during the Pleistocene. Nevertheless, records of this family are scarce in Mexico. The only valid species reported from the Late Pleistocene in Mexico were Platygonus compressus and Dicotyles tajacu, based on a few specimens. In this study, we report a new peccary specimen from the Gliptodonte locality (Late Pleistocene, Rancholabrean NALMA), Villaflores municipality, in the southern State of Chiapas. The new specimen consists of a distal part of the left humerus, which shows morphological differences from other North American Pleistocene and recent peccaries (Mylohyus, Platygonus, and Tayassu), allowing positive identification of the collared peccary D. tajacu. Thus, a new record of the species for the State of Chiapas and the first record of the Gliptodonte locality in the municipality of Villaflores is added, expanding the distribution range of the species during the Late Pleistocene in Mexico.

Keywords: collared peccary, Pleistocene mammals, Rancholabrean NALMA, southern Mexico.

Resumen

Tayassuidae (pecaríes) es una familia de artiodáctilos exclusiva de América, que se distribuyó ampliamente en América del Norte durante el Pleistoceno. Sin embargo, los registros de esta familia son escasos en México. Las únicas especies válidas reportadas para el Pleistoceno Tardío de México son Platygonus compressus y Dicotyles tajacu, con base en muy pocos especímenes. En este estudio reportamos un nuevo ejemplar de pecarí de la localidad Gliptodonte (Pleistoceno Tardío, NALMA Rancholabreano), municipio de Villaflores, en el sureño Estado de Chiapas. El nuevo espécimen consiste en la parte distal del húmero izquierdo, que muestra diferencias morfológicas con otros pecaríes del Pleistoceno y del Reciente de América del Norte (Mylohyus, Platygonus y Tayassu), lo que permitió la identificación del pecarí de collar D. tajacu. Se suma así un nuevo registro de la especie para el Estado de Chiapas y el primer registro de la localidad Gliptodonte en el municipio de Villaflores, ampliando el rango de distribución de la especie durante el Pleistoceno Tardío en México.

Palabras clave: Mamíferos pleistocénicos, NALMA Rancholabreano, pecarí de collar, sureste de México.

1. Introduction

Tayassuidae Palmer, 1897, is a New World family of artiodactyls that diverged in the Eocene from a common Asian ancestor with the family Suidae, but has radiated successfully in North America, with a chronological range from the late Eocene to the present (Harris and Li-Ping, 2007; Prothero, 2009, 2021). This family was one of the first to cross South America during the Great American Biotic Interchange in the middle Pliocene (GABI 1; Woodburne, 2010).

Tayassuidae were very diverse at the end of the Neogene and the beginning of the Quaternary, with the Pleistocene being the geological epoch with most genera and species. During this time interval, six genera were recognized in South America and five in North America. The genera Brasiliochoerus Rusconi 1930; Catagonus Ameghino, 1904; Parachoerus Rusconi, 1930; Dicotyles Cuvier, 1816; Platygonus Le Conte, 1848 and Tayassu Fischer, 1814 are present in South America (Gasparini et al., 2013; Parisi Dutra et al., 2017). The genera Mylohyus Cope, 1889; Platygonus, and Dicotyles, have been recognized in the Pleistocene of North America, but Tayassu does not have a fossil record until now (Kurtén and Anderson, 1980; Mayer and Wetzel, 1987; Hulbert et al., 2009; Czaplewski, 2012; Prothero, 2021). Recently, a new taxon, Muknalia minima, was established based on an almost complete mandibular ramus, recovered from the Muknal cave (Late Pleistocene-early Holocene), which is part of the Ox Bel Ha system, in Quintana Roo, Yucatán (Stinnesbeck et al., 2017). However, M. minima were relegated as a junior synonym of Pecari tajacu (=Dicotyles tajacu [Linnaeus, 1758]) by Schubert et al. (2020). Another Pleistocene Mexican record of peccaries is the flat-headed peccary Platygonus compressus Le Conte, 1848, documented for El Cedazo, Aguascalientes; Tequixquiac, State of Mexico; Valsequillo, Puebla and the Chapala Basin, Jalisco (Lucas, 2008; Ferrusquía-Villafranca et al., 2010). Other described species from the Pleistocene of Mexico are P. ticuli Mones, 1974 from Chapala and Zocoalco, Jalisco (Mones, 1974) and P. alemanii Dugès, 1891 from Moroleón, Guanajuato (Dugès, 1891). However, P. ticuli has been synonymized with P. vetus Leidy, 1882 by Lucas (2008), while P. alemanii does not differ from P. compressus, so it can be synonymized with this species. The collared peccary Dicotyles tajacu has been documented for the Loltún and Actún Spukil caves in Yucatán (Arroyo-Cabrales and Álvarez, 2003), and for the Santa Marta shelter in Ocozocoautla, Chiapas, based on cranial and appendicular material from the layer XVI, corresponding to the Pleistocene (García-Bárcena, 1976; Acosta et al., 2018).

The fossil record of peccaries in Mexico is poorly known. Therefore, the aim of this work is to report for the first time the presence of the collared peccary Dicotyles tajacu in the Gliptodonte locality, Villaflores municipality, based on the distal part of a humerus collected from the Late Pleistocene (NALMA Rancholabrean) fluviolacustrine sediments that crop out in that locality. This discovery contributes to the knowledge of the Late Pleistocene fauna in southern Mexico.

2. Study area and geological setting

The Gliptodonte locality (locally named Rancho Argentina) is located in the vicinity of the town Villaflores, at 16°12'55''N and 93°16'20''W (Figure 1). This locality is registered with the code 2PSP00000134 in the National Database of Paleontological Monuments in the Dirección de Registro Público de Monumentos y Zonas Arqueológicos e Históricos of Instituto Nacional de Antropología e Historia (INAH).

The sedimentary sequence is formed by alternating fluvial sediments of 8.0 m thick (Figure 2). The base is 1.6 m thick and is formed by olive silt, thin sand, and brown clay organic. Over the base section, there is a 0.30 m layer of brown silty clay. In this section, fossil remains of the glyptodont Glyptotherium cylindricum (Brown, 1912), the ground sloth Eremotherium laurillardi (Lund, 1842), the horses Equus mexicanus (Hibbard, 1955), E. conversidens Owen, 1869, and the with-tailed deer Odocoileus virginianus Zimmermann, 1780 have been recovered (Montellano-Ballesteros and Carbot-Chanona, 2010; Gómez-Pérez and Carbot-Chanona, 2012; Jiménez-Hidalgo et al., 2019; Carbot-Chanona et al., 2022). The next section is a 1.0 m layer composed of dark brown clay and brown clay-silt. Above this section, there is a 0.40 m layer of fine sand to silt section without boulders. The next layer is formed by 0.70 m of brown fine clay. Above this section is a 0.6 m of clay with organic matter; in this section there are G. cylindricum, E. conversidens, and O. virginianus fossil remains (Gómez-Pérez and Carbot-Chanona, 2012; Jiménez-Hidalgo et al., 2019). The next layer is formed by 0.60 m of sandy silt. Above there is a 0.40 m of fine-grained sand. Over this section, there is a 0.50 m layer of silt with medium sand lenses. The next layer is formed by 0.70 m medium sand with rounded pebbles and the presence of manganese lines. Finally, the upper part of the sequence consists of 1.2 m coarse sand with large lithoclasts.

The faunal association present in the locality has been referred to Rancholabrean NALMA (Gómez-Pérez and Carbot-Chanona, 2012).

3. Material and methods

The specimen described here consists of the distal half of a completely permineralized left humerus that presents a median state of conservation and was recovered from horizontal fluvial sediments that crop out in the Gliptodonte locality in the municipality of Villaflores, Chiapas.

The anatomical terminology used in this study is based on Schmid (1972) and Bortolami and Callegari (1995). The measurements were obtained with a digital caliper with an error range of 0.1 mm and were taken as shown in Figure 3. This study follows the proposal of Acosta et al. (2020), who argue that the valid binomina for the collared peccary is Dicotyles tajacu, and not Pecary tajacu, Tayassu tajacu, or Dicotyles crassus.

Since artiodactyl taxa of similar sizes existed in the Late Pleistocene of North America, such as deer (Cervidae Gray, 1821), antilocaprids (Antilocapridae Gray, 1866), and peccaries (Tayassuidae), a first comparison was conducted with recent specimens of these groups to assign the material studied here to a particular family. For this purpose, osteological material of the withe-tailed deer Odocoileus virginianus (specimens IHNCR-001 and IHNCR-007), the pronghorn Antilocapra americana Ord, 1815 (DP 1555), the with-lipped peccary Tayassu pecari (Link, 1975) (IHNCR-042) and the collared peccary Dicotyles tajacu (specimens DP 638, DP 5099, DP 7779 and IHNCR-048) was used. Subsequently, the fossil material was compared in detail morphologically and meristically with the tayasuid species present in the North American Rancholabrean NALMA, Mylohyus fossilis (Leidy, 1860) and P. compressus, and with the living T. pecari and D. tajacu (Figures 5 and 6, Table 1).

3.1. Institutional abbreviations

DP, Osteological Collection, Laboratorio de Arqueozoología “M. en C. Ticul Álvarez Solórzano” (historically Departamento de Prehistoria), Instituto Nacional de Antropología e Historia, Ciudad de México, México. IHNFG, Instituto de Historia Natural, Fósil Geográfico (acronym historically used for the paleontological collection now housed at Secretaría de Medio Ambiente e Historia Natural), Tuxtla Gutiérrez, Chiapas, México. IHNCR, Instituto de Historia Natural, Reference Collection, Museo de Paleontología “Eliseo Palacios Aguilera”, Secretaría de Medio Ambiente e Historia Natural, Tuxtla Gutiérrez, Chiapas, México.

3.2. Morphological abbreviations

afc, articulation of the flexor complex; lc, lateral condyle; mc, medial condyle; le, lateral epicondyle; me, medial epicondyle; of, olecranon fossa; rf, radial fossa; stf, supratrochlear foramen, tr, trochlea.

3.3. General abbreviations

NALMA, North American Land Mammal Age.

4. Results

4.1. Systematic Paleontology

Class Mammalia Linnaeus, 1758

Order Artiodactyla Owen, 1848

Suborder Suiformes Jaekel, 1911

Family Tayassuidae Palmer, 1897

Genus Dicotyles Cuvier, 1816

Dicotyles tajacu (Linnaeus, 1758)

Figures 4 (3D model), 5, and 6, Table 1

4.2. Referred material

IHNFG-2718, distal half of left humerus (Figure 4), collected by Valente Villanueva in 2003 and later donated to the Museo de Paleontología “Eliseo Palacios Aguilera”. The specimen is housed in the paleontological collection of the Secretaría de Medio Ambiente e Historia Natural, State of Chiapas, southern Mexico.

4.3. Location and horizon

Gliptodonte locality, Villaflores, Chiapas, southeastern Mexico. Late Pleistocene, Rancholabrean NALMA.

4.4. Description

The IHNFG-2718 specimen corresponds to the distal half of the left humerus (Figure 4). The diaphysis-epiphysis suture was completely ossified, indicating that it was an adult specimen. The lateral and medial borders of the articular end as well as the borders of the articular condyles were rounded, indicating that the material underwent a process of dragging after fossilization. In the anterior view, it can be seen that the medial condyle is slightly wider than the lateral condyle. The medial epicondyle projected slightly medially, whereas the lateral epicondyle extended laterally. The articulating surface of the flexor complex lies distally on the medial epicondyle and is projected distally. In the medial view, the flexor complex joint extends from the posterior part of the medial epicondyle to the medial part of the epicondyle. The olecranon fossa is wide and shallow, and the supratrochlear foramen is located inside. In the posterior view, the lateral supracondylar crest was observed to be incipiently protruding.

5. Discussion

5.1. Comparison of IHNFG-2718 with medium-sized artiodactyls

Medium-sized artiodactyls of the families Antilocapridae (Stockoceros conklingi Stock, 1930), Cervidae (Odocoileus virginianus and Navahoceros fricki Schultz and Howard, 1935), and Tayassuidae (Platygonus compressus, Mylohyus fossilis, and Dicotyles tajacu) inhabited North America during the Rancholabrean (Kurtén and Anderson 1980). Therefore, a first comparison was made between IHNFG-2718 and living representatives of these families, to include the specimen in one of them.

IHNFG-2718 is an adult specimen because the epiphysis-diaphysis suture was completely ossified. Its dimensions are smaller than those of the humeri of Antilocapra americana (DP 1555) and Odocoileus virginianus (IHNCR-001 and IHNCR-007). Both A. americana and O. virginianus were approximately 15% larger than IHNFG-2718. In contrast, the dimensions of IHNFG-2718 were comparable to those of an adult specimen of T. pecari and D. tajacu (Table 1).

Detailed morphological comparisons between IHNFG-2718, O. virginianus, A. americana, and T. pecari revealed distinctive features in the humerus of these species. In anterior view, the medial condyle and the lateral condyle in IHNFG-2718 were of similar proportions, as observed in the humerus of T. pecari and D. tajacu, whereas in the humerus of O. virginianus and A. americana, the medial condyle was wider laterally-medially than the lateral condyle (Figure 5). In the posterior view, IHNFG-2718 presents the supratrochlear foramen in the olecranon fossa, which is present in T. pecari and D. tajacu, but not in O. virginianus or A. americana. Additionally, the inner lower border of the olecranon fossa in T. pecari and D. tajacu was almost horizontal, a feature observed in IHNFG-2718. In contrast, in O. virginianus and A. americana, the border was lateromedially inclined (Figure 5). Another striking feature is that the articulating surface of the flexor complex is more distally prominent in T. pecari, D. tajacu, and IHNFG-2718 than in O. virginianus and A. americana. Additionally, the olecranon fossa is wider and deeper in IHNFG-2718 than in O. virginianus and A. americana, but is similar to the olecranon fossa of T. pecari and D. tajacu.

All the morphological and meristic features of IHNFG-2718 indicate that it is a tayasuid, similar to T. pecari and D. tajacu.

5.2. Comparison of IHNFG-2718 to another North American tayasuids

The genera of tayasuids present in the Rancholabrean of North America, Platygonus, and Mylohyus, and the living Tayassu and Dicotyles, differ from each other mainly by craniomandibular features and size (see Prothero, 2021). Of the four taxa, Mylohyus is the largest, a trait that is reflected in the postcranial bones, which are slightly larger than those of Platygonus, Tayassu, and Dicotyles (Figure 5, Table 1). In addition to size, there were morphological differences between the humeri of the four genera. The humerus in Mylohyus, Tayassu, and Dicotyles is similar in general morphology and proportion and is slenderer than the humerus of Platygonus (Lundelius, 1960). In Mylohyus, the crest on the lateral border of the olecranon fossa is farther from the fossa and lies on the lateral surface of the humerus (Lundelius, 1960). This feature is similar in Tayassu and Dicotyles, while in Platygonus the crest is located on the back of the humerus (Lundelius, 1960). Additionally, the medial border of the olecranon fossa is narrower and more angular in Mylohyus than in Platygonus, a feature that is similar in Tayassu and Dicotyles (Lundelius, 1960). In the specimen IHNFG-2718, the morphological characteristics present in Tayassu and Dicotyles are observed, in addition to the fact that its measurements fall within the size range of these two genera (Table 1), so it is ruled out that it belongs to Mylohyus or Platygonus.

Although there are differences in size between the extant species of peccaries, Tayassu pecari (total length= 905-1,390 mm; height at shoulders= 400-530 mm; body mass= 25-40 kg; Mayer and Wetzel, 1987) and Dicotyles tajacu (total length= 870-940 mm; height at shoulders= 400-480 mm; body mass= 18-27 kg; Hall, 1981), these differences in size are not noticeable in the dimensions of the humeri compared in this study (Figure 6, Table 1). However, a detailed comparison between the humeri of these species allowed us to identify some morphological differences.

In the posterior view, the humerus of T. pecari presents a slight elevation on the internal lateral border of the olecranon fossa, which extends distally to the middle part of the lateral epicondyle, forming a small “ridge” (Figure 6). In contrast, in the D. tajacu humerus, the medial lateral border of the olecranon fossa, this “ridge” is not present. In the specimen IHNFG-2718, the “ridge” of T. pecari humerus was not observed either, but the internal lateral edge of the olecranon fossa was similar to that of D. tajacu (Figure 6). Based on this morphological characteristic, the specimen IHNFG-2718 is assigned to Dicotyles tajacu, thereby adding a new record of the species for the state of Chiapas and the first record for the Gliptodonte locality, in the municipality of Villaflores, thus expanding the distribution range of the species during the Late Pleistocene in Mexico.

5.3. Paleoenvironmental significance

Today, D. tajacu inhabits deserts, arid woodlands, oak woodlands, and tropical areas from the southern U.S.A. through to northern Argentina (Grubb and Groves, 1993). This species has unusual climatic tolerance, because it occurs in forests where average temperatures are around 27 °C, but some populations have also been found in desert areas where temperatures reach 45 °C (Bodmer and Sowls, 1993). The distribution of D. tajacu is clearly tropical and semi-tropical, and agrees with the presence of neotropical mammals in the Gliptodonte locality, such as E. laurillardi and G. cylindricum (Gómez-Pérez and Cabot-Chanona, 2012; Carbot-Chanona et al., 2022), and with the palynological fossil record, that includes Compositae, Mimosoideae, Gramineae, Solanaceae, and Pinaceae (Carbot-Chanona et al., 2008), indicating that this was the main type of environment in Chiapas during the final Pleistocene.

6. Conclusions

Although the material described here consists only of a distal fragment of the left humerus, an exhaustive morphological comparison with the humeri of various species of living and fossil North American medium-sized artiodactyls allowed us to refer the specimen IHNFG-2718 as belonging to the collared peccary Dicotyles tajacu, a species that lives in several regions of the State of Chiapas. With this finding, we document a new record of this species for the Late Pleistocene in Mexico, as well as the first record of the species for the Gliptodonte locality.

The presence of D. tajacu in Chiapas allows us to know the fauna present in southeastern Mexico during the Late Pleistocene. In future studies, these findings could provide data to understand which taxa managed to prevail over the extinction caused in the Pleistocene-Holocene interval as a consequence of climate change.

Conflicts of interest

The authors declare that they have none.

Acknowledgments

The material described in this work was recovered thanks to the project “Rescate de la paleobiodiversidad de Chiapas” sponsored by the government of the State of Chiapas. The digital 3D model is part of the “Estrategias para la divulgación de la ciencia a través de la virtualización 3D del Patrimonio Paleontológico” project, sponsored by the Instituto de Ciencia, Tecnología e Innovación del Estado de Chiapas. We thank Joaquín Arroyo-Cabrales and Ana Fabiola Guzman† for allowing us access to the osteological collection at Laboratorio de Arqueozoología “M. en C. Ticul Álvarez Solórzano”, Instituto Nacional de Antropología e Historia. The Platygonus photograph was provided by the University of Florida. We also thank Eduardo Jiménez-Hidalgo for the comments on the early draft of this manuscript, and the two anonymous reviewers that provided useful comments that improved the manuscript, also Josep Moreno-Bedmar for their editorial support.

References

Acosta, G., Beramendi, L. E., González, G., Rivera, I., Eudave, I., Hernández, E., Sánchez, S., Morales, P., Cienfuegos, E., & Otero, F. (2018). Climate change and peopling of the Neotropics during the Pleistocene-Holocene transition. Boletín de la Sociedad Geológica Mexicana, 70(1), 1–19. https://doi.org/10.18268/bsgm2018v70n1a1

Acosta, L., Garbino, G., Gasparini, G., &Parisi Dutra, R. (2020). Unraveling the nomenclatural puzzle of the collared and white-lipped peccaries (Mammalia, Cetartiodactyla, Tayassuidae). Zootaxa, 4851(1), 60–80. https://doi.org/10.11646/zootaxa.4851.1.2

Ameghino F. (1904). Nuevas especies de mamíferos cretáceos y terciarios de la República Argentina. Anales de la Sociedad Científica Argentina, 58, 35–291.

Arroyo-Cabrales J. & Álvarez, T. (2003). A preliminary report of the late Quaternary mammal fauna from Loltún cave, Yucatán, Mexico. In Schubert, B. W., Mead, J. I., Graham, R. W. (Eds.). Ice Age cave faunas of North America. (pp. 262–272). Denver: Museum of Nature and Science.

Bodmer, R. E., & Sowls, L. K. (1993). The Neotropical tayassuids Tayassu and Catagonus: The collared peccary (Tayassu tajacu). In Oliver, W. L. R. (ed.). Pigs, peccaries, and hippos, (7–13). International Union for Conservation of Nature and Natural Resources.

Bortolami, R., & Callegari, E. (1995). Anatomia comparata dei mammiferi domestici. Italia: Edizione Agricole della Calderini s.r.l.

Brown, B. (1912). Brachyostracodon, a new genus of glyptodonts from Mexico. Bulletin of the American Museum of Natural History, 31(17), 167–177.

Carbot-Chanona, G., Gómez-Pérez, L. E., & Coutiño-José, M.A. (2022). A new specimen of Eremotherium laurillardi (Xenarthra, Megatheriidae) from the Late Pleistocene of Chiapas, and comments about the distribution of the species in Mexico. Boletín de la Sociedad Geológica Mexicana, 74(2), 1–12. https://doi.org/http://dx.doi.org/10.18268/ BSGM2022v74n2a070322

Carbot-Chanona, G., Vázquez-Bautista, D., Montellano-Ballesteros, M., Domínguez-Vázquez, G., & Islebe, G. A. (2008). Reconstrucción paleoambiental del Pleistoceno tardío de Chiapas (abstract). In IX Seminario de Interno de Investigación, Instituto de Historia Natural, p. 5.

Cope, E. D. (1889). Vertebrata of the Swift Current River. Nos. II. American Naturalist, 23, 151–155.

Cuvier, G. (1816). Le règne animal distribué d’après son organisation, pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. Vol. 1. Les mammifères et les oiseaux. A. Belin, Paris, xxxvii, 540 pp.

Czaplewski, N. J. (2012). Pleistocene peccaries (Mammalia: Tayassuidae) from western Oklahoma. The Southwestern Naturalist, 57(1), 112–117. https://doi.org/10.1894/0038-4909-57.1.112

Dugès, A. (1891). Platygonus alemanii nobis, fósil Cuaternario. Naturaleza, 2, 15–18.

Ferrusquía-Villafranca, I., Arroyo-Cabrales, J., Martínez-Hernández, E., Gama-Castro, J., Ruiz-González, J., Polaco, O.J., & Johnson, E. (2010). Pleistocene mammals of Mexico: A critical review of regional chronofaunas, climate change response and biogeographic provinciality. Quaternary International, 217(1), 53–104.

Finch, W. I., Whitmore jr., F. C., & Sims, J. D., 1970, Stratigraphy, morphology, and paleoecology of a fossil peccary herd from western Kentucky. U.S. Geological Survey Professional Paper, 790, 1–25 .

Fischer, G. (1814). Zoognosia tabulis synopticis illustrata. Vol. 3. Quadrupeda reliqua, Ceti Monotrymata: Nicolai Sergeidis Vsevolozsky, Moscow .

García-Bárcena, J. (1976). Excavaciones en el abrigo de Santa Marta, Chis (1974 ). Informe Preliminar, Mecanoescrito en Archivo Técnico del INAH.

Gasparini, G. M., Ubilla, M., & Tonni, E. P. (2013). The Chacoan peccary, Catagonus wagneri (Mammalia, Tayassuidae), in the late Pleistocene (northern Uruguay, South America): paleoecological and paleobiogeographic considerations. Historical Biology, 25(5–6), 679–690. https://doi.org/10.1080/08912963.2012.742519

Gómez-Pérez, L. E., & Carbot-Chanona, G. (2012). Contribución al estudio de los megamamíferos del Pleistoceno Tardío del municipio de Villaflores, Chiapas, México. Lacandonia, 6(1), 31–41.

Gray, J. E. (1821). On the natural arrangement of vertebrose animals. The London Medical Repository Monthly Journal and Review, 15, 296–310.

Gray, J. E. (1866). Notes on the Proghorn Buck (Antilocapra), and its position in the system. Annals and Magazine of Natural History, 18(10), 323–326.

Grubb, P., & Groves, C. (1993). The Neotropical tayassuids Tayassu and Catagonus: Taxonomy and description In Oliver, W.L.R. (eds.). Pigs, Peccaries, and Hippos, (pp. 5-7). International Union for Conservation of Nature and Natural Resources.

Hall, E. R. (1981). The Mammals of North America, Vol. II. New York: John Wiley & Sons.

Harris, J. M., & Liu, L. (2007). Superfamily Suoidea. In Prothero, D.R. and Foss, S.E. (eds.). The evolution of Artiodactyls, 130–150. Johns Hopkins University Press.

Hibbard, C. W. (1955). Pleistocene vertebrates from the upper Becerra Formation, Valley of Tequixquiac, Mexico, with notes on other Pleistocene forms. University of Michigan, Contributions from the Museum of Paleontology, 12 (5), 47–96.

Hulbert, R. C., Morgan, G. S., & Kerner, A. (2009). Collared peccary (Mammalia, Artiodactyla, Tayassuidae, Pecari) from the Late Pleistocene of Florida. In Papers on Geology, Vertebrate Paleontology, and Biostratigraphy in Honor of Michael O. Woodburne. Museum of Northern Arizona Bulletin 65, Flagstaff, Arizona, 543–556.

Jaekel, O. (1911). Die Wirbeltiere. Eine Übersicht über die Fossilen und Lebenden Formen: Berlin: Verlag Gebrüder Borntraeger.

Jiménez-Hidalgo, E., Carbot-Chanona, G., Guerrero-Arenas, R., Bravos-Cuevas, V. M., Holdridge, G. F., & Israde-Alcántara, I. (2019). Species diversity and paleoecology of Late Pleistocene horses from southern Mexico. Frontiers in Ecology and Evolution, 7, 394, 1–18. Doi: 10.3389/fevo.2019.00394

Kurtén, B., & Anderson, E. (1980). Pleistocene mammals of North America. New York: Columbia University Press.

Le Conte, J. L. (1848). On Platygonus compressus: a new fossil pachyderm. Memoirs of the American Academy of Arts and Sciences, 3, 257–274.

Leidy, J. (1860). Description of vertebrate fossil. In Holmes, F.S. (ed.). Post-Pleiocene fossils of South Carolina (99–122). Charleston: Russell & Jones.

Leidy, J. (1882). On an extinct peccary: Proceedings of the Academy of Natural Science of Philadelphia, 301–302.

Link, H. F. (1795). Ueber die Lebenskraefte in naturhistorischer Rücksicht und die Classification der Saeugthiere. Beytraege zur Naturgeschichte. Vol. 2: K.C. Stiller, Rostock und Leipzig, 126 pp.

Linnaeus, C. (1758). Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1. 10th Edition. L. Salvius, Holmia (Stockholm), 824 pp.

Lucas, S. G. (2008). Late Cenozoic fossil mammals from the Chapala rift basin, Jalisco, Mexico. New Mexico Museum of Natural History and Science Bulletin, 44, 39–49.

Lund, P. W. (1842). Blik paa Brasiliens dyreverden för sidste jordomvaeltning. tredje afhandling: Forsaettelse af Pattedyrene. Det Kongelige Danske Videnskabernes Selskbas Naturvidenskabelige og Mathematiske Afhandlinger, 9, 137–208.

Lundelius Jr., E. L. (1960). Mylohyus nasutus long-nosed peccary of the Texas Pleistocene. Bulletin of the Texas Memorial, 1, 1–40.

Mayer, J. J., & Wetzel, R. M. (1987). Tayassu pecari. Mammalian Species, 293, 1–7.

Mones, A. (1974). Nueva especie de pecarí fósil (Platygonus) del Estado de Jalisco. Anales Instituto Nacional de Antropología e Historia, 51, 1970-1971, 119–128.

Montellano-Ballesteros, M., & Carbot-Chanona, G. (2010). Presencia de Odocoileus (Artiodactyla, Cervidae) en el Pleistoceno de Chiapas, México. In Cervantes, F.A., Hortelano Moncada, Y., & Vargas Cuenca, J. (eds.). 60 Años de la colección Nacional de Mamíferos del instituto de Biología, UNAM (pp. 291–298). México: Universidad Nacional Autónoma de México.

Ord, G. (1815). North American zoology In Gudrie’s Geography (2nd American edition): Johnson and Warner: Philadelphia, Pennsylvania, 292–308.

Owen, R. (1848). Description of teeth and portions of jaws of two extinct Anthracotherioid quadrupeds (Hypopotamus vectianus and Hyop. bovinus) discovered by the Marchioness of Hastings in the Eocene deposits on the NW coast of the Isle of Wight: with an attempt to develp Cuvier’s idea of the classification of Pachyderms by the number of their toes. Quaterly Journal of the Geological Society of London, 4, 103–141.

Owen, R. (1869). On fossil remains of equines from Central and South America referable to Equus conversidens, Ow., Equus tau, Ow., and Equus arcidens, Ow. Philosophical Transactions of the Royal Society of London, 159, 559–573.

Palmer, T. S. (1897). Notes on the nomenclature of four genera of tropical american mammals. Proceedings of the Biological Society of Washington, 11, 173–174.

Parisi Dutra, R., Casali, D. M., Missagia, R. V., Gasparini, G. M., Perini, F. A., & Cozzuol, M. A. (2017). Phylogenetic systematics of peccaries (Tayassuidae: Artiodactyla) and a classification of South American tayassuids. Journal of Mammalian Evolution, 24(3), 345–358. https://doi.org/10.1007/s10914-016-9347-8

Prothero, D. R. (2009). The early evolution of the North American peccaries (Artiodactyla: Tayassuidae). Museum of Northern Arizona Bulletin 65(1969), 509–541. http://www.donaldprothero.com/files/92367802.pdf

Prothero, D. R. (2021). The systematics of North American peccaries (Mammalia: Artiodactyla: Tayassuidae). New Mexico Museum of Natural History & Science Bulletin, 85, 1–76 .

Prothero, D. R., & Grenader, J., 2012, a New primitive species of the flat-headed peccary. Journal of Paleontology, 86(6), 1021–1031.

Rusconi, C. (1930). Las especies fósiles argentinas de pecaríes y sus relaciones con las del Brasil y Norteamérica. Annales del Museo Nacional de Historia Natural Bernardino Rivadavia, 36, 121–241.

Schmid, E. (1972). Atlas of Animal Bones: For Prehistorians, Archaeologists and Quaternary Geologists. Amsterdam and New York: Elsevier Publishing Company.

Schubert, B. W., Samuels, J. X., Chatters, J. C., & Arroyo-Cabrales, J. (2020). Muknalia minima from the Yucatán of Mexico is synonymous with the Collared Peccary, Pecari Tajacu (Artiodactyla: Tayassuidae). Open Quaternary, 6(1), 1–9. https://doi.org/10.5334/oq.84

Schultz, C. B., & Howard, E. B. (1935). The fauna of Burnet Cave, Guadalupe Mountains, New Mexico. Proceedings of the Academy of Natural Sciences of Philadelphia, 87, 273–298.

Stinnesbeck, S. R., Frey, E., Stinnesbeck, W., Avíles Olguín, J., Zell, P., Terrazas Mata, A., Benavente Sanvicente, M., González González, A., Rojas Sandoval, C., & Acevez Nuñez, E. (2017) A new fossil peccary from the Pleistocene-Holocene boundary of the eastern Yucatán Peninsula, Mexico. Journal of South American Earth Sciences, 77, 341–349. https://doi.org/10.1016/j.jsames.2016.11.003

Stock, C. (1930). Quaternary antelope remains from a second cave deposits in the Organ Mountains, New Mexico. Los Angeles County Museum Publications, 2, 1–18.

Woodburne, M. O. (2010). The Great American Biotic Interchange: Dispersals, tectonics, climate, sea level and holding pens. Journal of Mammalian Evolution, 17(4), 245–264. https://doi.org/10.1007/s10914-010-9144-8

Zimmermann, A. W. (1780). Geographische Geschichte des Menschen und der vierfüssigen Thiere. In der Weygandschen Buchhandlung, Leipzig, 2, 1–432.

Paleozoología

Figure 1. Location of the Gliptodonte locality, in Villaflores, Chiapas, southern Mexico.

Figure 2. Generalized stratigraphic section of the Gliptodonte locality, Villaflores, Chiapas, southern Mexico

Paleozoología

Figure 3. Schematic drawing of the humerus showing the measurements taken. 1) Lateral-medial length through the epicondyles, and 2) anteroposterior width.

Figure 4. 3D model of the IHNFG-2718 left humerus. To see the model with better resolution, access in https://sketchfab.com/3d-models/humero-fosil-de-pecari-de-collar-40207bbe0d3d46b4a45550ea2aca46bf

Figure 5. Comparison between North American medium-sized artiodactyls. A–D, left humerus (IHNFG-2718), in anterior (A), medial (B), posterior (C), and lateral (D) view. E–H, left humerus of Tayassu pecari (IHNCR-042), in anterior (E), medial (F), posterior (G), and lateral (H) view. I–L, right humerus (flipped images) of Platygonus compressus (UF-61610), in anterior (I), medial (J), posterior (K), and lateral (L) view. M–O, right humerus (flipped images) of Mylohyus fossilis (RMM 4567), in anteromedial (M), posterior (N), and posterolateral (O) view. P–S, left humerus of Odocoileus virginianus (INHCR-001), in anterior (P), medial (Q), posterior (R), and lateral (S) view. T–W, left humerus of Antilocapra americana (DP 1555), in anterior (T), medial (U), posterior (V), and lateral (W) view. Scale bar equal 50 mm.

Figure 6. Comparison between (A) IHNFG-2718, (B) Dicotyles tajacu (IHNCR-048), and (C) Tayassu pecari (IHNCR-042) in posterior view. Scale bar equal 30 mm. The yellow arrow shows the slight elevation on the internal lateral border of the olecranon fossa present in T. pecari, but not in D. tajacu and IHNFG-2718.

Table 1. Comparative measurements (in mm) of tayassuids during the Late Pleistocene in North America. Data were taken from Finch et al. (1970)a and Lundelius (1969)b.

Specimens Lateral-medial length through the epicondyles Anteroposterior width IHNFG-2718 29 26 Tayassu pecari IHNCR-045 27 24 Dicotyles tajacu DP 608 27.5 23 Dicotyles tajacu DP 5099 25.4 22.2 Dicotyles tajacu DP 7779 29 24.8 Dicotyles tajacu IHNCR-048 27 23 aPlatygonus compressus USNM 26098 42 -- aPlatygonus compressus USNM 26100 45 -- aPlatygonus compressus USNM 26102 40.5 -- Platygonus compressus UF 61610 38.6 36.6 Mylohyus nasutus (=M. fossilis) RMM 4567 46.7 33 bMylohyus nasutus (=M. fossilis) TMM 933-3232 42.5 45.6