RECORD OF TAYASSUIDS IN ?LATE PLIOCENE TO QUATERNARY … · 2020-03-19 · As one of the North American immigrants that entered South America during the Great American Biotic Interchange,

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Rev. bras. paleontol. 17(2):169-182, Maio/Agosto 2014© 2014 by the Sociedade Brasileira de Paleontologiadoi: 10.4072/rbp.2014.2.05

INTRODUCTION

The second stage of the Great American Biotic Interchange (GABI) took place when the Panamanian land bridge became complete some time during the late Pliocene (Webb, 1985). Among the groups that entered South America at this time are the tayassuids. They represent one of the fi rst mammalian immigrants from North America, after procyonids (Carnivora) and cricetid rodents (Prevosti et al., 2006; Woodburne et al., 2006; Woodburne, 2010), although the exact moment of tayassuids arrival is controversial. The oldest fossil tayassuid in South America dates to the beginning of the late Chapadmalalan (middle Pliocene, 3.3-4 Ma, Prevosti et al., 2006; 3.7 Ma Woodburne, 2010; Gasparini, 2013). But

Campbell et al. (2010) described two indeterminate forms of extinct peccaries, among other mammals of Neartic origin (gomphotheres, tapirs), collected from the Madre de Dios Formation (9.5 Ma), in southern Peru. These Peruvians records are considered controversial by some authors (Woodburne, 2010). Another issue related to the evolution of tayassuids that remains unclear is if both the extinct and living genera of tayassuids dispersed to South America from Central America as distinct lineages in multiple events, or diverged after a single dispersal event (Frailey & Campbell, 2012).

After their arrival in South America, the Tayassuidae had a great taxonomic diversity and a wide geographic distribution. The most abundant and diverse fossil record comes from Argentina (Gasparini, 2013). Members of the

RECORD OF TAYASSUIDS IN ?LATE PLIOCENE TO QUATERNARY DEPOSITS IN VENEZUELA

MARISOL MONTELLANO-BALLESTEROSDepartamento de Paleontología, Instituto de Geología, UNAM Circuito Exterior, Ciudad Universitaria,

Delegación Coyoacán, México D.F. 04510. [email protected]

ASCANIO D. RINCÓN & ANDRÉS SOLÓRZANOInstituto Venezolano de Investigaciones Científi cas, Laboratorio de Paleontología, Centro de Ecología, Caracas, Venezuela.

[email protected], [email protected]

ABSTRACT ‒ Tayassuidae is represented by three extant species whose distribution ranges from southwestern United States to north-central Argentina. As one of the North American immigrants that entered South America during the Great American Biotic Interchange, the oldest unquestionable record of the family dates to the middle Pliocene in Argentina. Intensive paleontological fi eldwork carried out in Venezuela resulted in many new localities containing vertebrate fossils remains. Among these sites, the fossil-rich tar pits and caves are particularly signifi cant. Dental remains of Tayassu pecari discovered in these sites are smaller than those described from other regions. The presence of Tayassu pecari in the ?late Pliocene Venezuelan locality represents the putative oldest record of this species in South America. A few specimens were identifi ed as Pecari tajacu. The Venezuelan tayassuid fossils document the presence of these two distinct North American immigrant lineages in northern South America, and provide information that improves our understanding of the distribution and diversity of the group in South America during the late Pliocene-early Holocene, following their entry into South American during the Great American Biotic Interchange.

Key words: caves, Pecari tajacu, tar pits, Tayassu pecari, Plio-Pleistocene, Venezuela.

RESUMO ‒ A família Tayassuidae está representada atualmente por três espécies, cuja distribuição varia do sudoeste dos Estados Unidos ao centro-norte da Argentina. Como um dos imigrantes norte-americanos que entraram na América do Sul durante o Grande Intercâmbio Biótico Americano, o mais antigo registro inquestionável da família data do Plioceno médio na Argentina. Intensivo trabalho de campo realizado na Venezuela resultou em muitas novas localidades contendo restos de fósseis de vertebrados. Entre esses sítios, os poços de piche e cavernas, ricos em fósseis, são particularmente signifi cativos. Restos dentários de Tayassu pecari descobertos nesses locais são menores do que aqueles descritos de outras regiões. A presença de Tayassu pecari na localidade venezuelana do Plioceno fi nal representa o registro mais antigo dessa espécie na América do Sul. Alguns espécimes foram identifi cados como Pecari tajacu. Os fósseis taiassuídeos venezuelanos documentam a presença destas duas linhagens distintas de imigrantes norte-americanos no norte da América do Sul, e fornecem informações que melhora a nossa compreensão sobre a distribuição e diversidade do grupo na América do Sul durante o Plioceno fi nal-Holoceno inicial, após a sua entrada nesse continente, durante o Grande Intercâmbio Biótico Americano.

Palavras-chave: cavernas, Pecari tajacu, poços de piche, Tayassu pecari, Plio-Pleistocene, Venezuela.

170 REVISTA BRASILEIRA DE PALEONTOLOGIA, 17(2), 2014

family have also been found in Brazil, Uruguay, Bolivia and Colombia (Rusconi, 1930, 1952; Stirton, 1947; Porta, 1969; Paula Couto 1975, 1981; Menegaz & Ortiz Jaureguizar, 1995; Ubilla et al., 2004; Villarroel et al., 1989; Gasparini et al., 2010).

According to Gasparini (2013), three genera of Tayassuidae are recognized in South America: Platygonus Le Conte, 1848 (middle Pliocene to early Pleistocene), with five extinct species; Catagonus Ameghino, 1904 (late Pliocene? to Recent), with fi ve species, one extant; and Tayassu Fischer, 1814 (middle Pleistocene to Recent), with two extant species. Wilson & Reeder (2005) have also recognized Pecari (Linnaeus, 1758) as a separate genus, a proposal that is followed here. Van Roosmalen et al. (2007) described a new species of living peccary in the Brazilian Amazon, Pecari maximus, although its validity has been recently questioned based on a molecular analysis of its DNA (Góngora et al., 2011).

Recently, Frailey & Campbell (2012) described two new genera and species of tayassuids, Sylvochoerus woodburnei and Waldochoerus bassleri, from the upper Miocene deposits on the western Amazon Basin, based on specimens that belong to the Harvey Bassler Collection of Peruvian Fossils, housed at the U.S. National Museum of Natural History and two specimens collected by the authors. According to Frailey & Campbell (2012) the dentition of Sylvochoerus resembles overall to that of Tayassu, and, that of Waldochoerus resembles Pecari, although they did not suggest any phylogenetic relationship with their fossil taxa and the living forms (Frailey & Campbell, 2012). If the late Miocene age of Waldochoerus and Sylvochoerus is confi rmed they would represent the oldest records of the family Tayassuidae in South America, and would suggest that these early tayassuids participated in the fi rst pulse of the Great American Biotic Interchange, at the same time the sloths reached North America.

In the last few years, intensive paleontological fi eldwork has been carried out in Venezuela, resulting in several new localities with vertebrate fossils. Among these there are important tar pits deposits, similar to Rancho La Brea in California, the best known being Mene de Inciarte in western Venezuela (Rincón, 2006 a,b; Rincón et al., 2006; Rincón et al., 2008). Tar pits deposits known as El Breal de Orocual (ORS16) discovered more recently in eastern of Venezuela, are producing an exceptional fauna that includes amphibians, reptiles, birds and mammals (Rincón et al., 2009; Holanda & Rincón, 2012; Costa-Fortier & Rincón, 2013). In addition, several caves are also known by its fossiliferous vertebrate remains: El Guácharo, Quebrada Honda, Del Miedo, El Zumbador, among others (Rincón, 2003). Rincón (2000) and McDonald et al. (2013) report fossil remains identifi ed as Tayassu sp. from Cueva de los Huesos, in the area of Cerro Pintado, Sierra de Perijá, Zulia State, Venezuela, (10º27’20”N/ 72º53’0”W), dated by radiocarbon to 14,150 ± 50 RC yr BP = (17,385 + 239 cal yr).

This paper describes the tayassuid material (except Platygonus) recovered from several ?late Pliocene-Quaternary localities in the northern part of Venezuela.

GEOLOGY, AGE AND LOCATION

The locality Mene de Inciarte (10°47’42.6’’N/ 72°14’20.8’’W) is in the vicinity of the Río Cachirí and the village of Cachirí at the foot of the Sierra de Perijá, State of Zulia, western Venezuela (Figure 1). This extensive asphalt seep or tar pits have a surface exposure 1,200 m long in a NE-SW direction and is up to 400 m wide at its widest part (Czaplewski et al., 2005). The local geology has been interpreted by Urbani & Galarraga (1991) as little-consolidated outcrops of sedimentary rocks corresponding to Pleistocene terraces. Jull et al. (2004) provided several radiocarbon dates from Mene de Inciarte using bone collagen and accelerator mass spectrometry. The youngest age, 25,500 ± 600 yr BP, was obtained from an osteoderm of Glyptodon sp. cf. G. clavipes.

Another tar pit locality, El Breal de Orocual, is located in the State of Monagas (9º50’48’’N/63º19’46.05’’W), near the town of Maturin, in eastern Venezuela (Figure 1). This tar seep partially overlies the Mesa Formation, which comprises fluvial-deltaic strata considered to have been deposited during the early to middle Pleistocene (González de Juana et al., 1980). Within the area of El Breal de Orocual two different fossil assemblages are known, one of these, ORS16 (close to an oil well numbered 16) was described by Rincón et al. (2009, 2011) the second ORS20 was only recently discovered and named after the closest oil well. The faunal assemblage from ORS16 suggests a probably late Pliocene-early Pleistocene age (Rincón et al., 2009; Rincón et al., 2011), while the ORS20 fauna suggests a late Pleistocene age.

In the State of Falcón, western Venezuela, there is a karstic system named Cerro Misión, located about 16 km south southwest of the town of Yaracal, where several caves have been discovered yielding fossil remains. The system includes the caves Cueva del Miedo (10º51’30”N/68º35’7”W) and Cueva de Rio Oro (10º50’27.1’’N/68º35’37.5’’W) (Figure 1). These caves are formed within the outcrops of the Capadare Formation, a thick sequence of massive middle Miocene reef limestone (Díaz de Gamero, 1985). Shark teeth and a bird remains had been recovered from the limestone of the Cueva del Miedo (Rincón & Stucchi, 2003), whereas late Pleistocene mammalian remains were recovered from the fl oor of the cave (Rincón, 2003; Rincón & White, 2007).

The Cueva El Guácharo (10°7’0”N/63°29’0”W) is located close to the town of Caripe, State of Monagas, eastern Venezuela (Figure 1). This cave is in the massive Aptian-Albian limestone of the El Cantil Formation and is probably the most important cave in Venezuela, with a horizontal extent of 9.425 m. Fossil remains were found in a paleosol in the Galería del Chorro (Morgan et al., 1988; Soibelzon & Rincón, 2007). The chronology of these fossils is not yet determined, but the presence of remains of Desmodus draculae and extant Diphylla ecaudata in cave strata have led to the assumption of late Pleistocene to early Holocene age (Morgan et al., 1988; Soibelzon & Rincón, 2007).

Another karstic system that contains some tayasuid remains is Cima de Los Escorpiones (10°08’02”N/

171MONTELLANO-BALLESTEROS ET AL. – TAYASSUIDS IN ?LATE PLIOCENE TO QUATERNARY IN VENEZUELA

64°28’29”W), located in Turimiquire Serrania, ten km from the town of Puerto La Cruz, State of Anzoátegui, western Venezuela. This cave is in the Santonian-Albian limestone of Querecual Formation, and is the deepest karstic system of Venezuela with a depth of 440 m (Gregoriani et al., 2008). The vertebrate fauna from this locality includes a jaw fragment of tayasuid, and some indeterminate perissodactyl remains.

MATERIAL AND METHODS

The fossil material is housed in the paleontological collection at the Instituto Venezolano de Investigaciones Científi cas (IVIC), Caracas, Venezuela. Within this collection the fossils from El Breal de Orocual are set apart and catalogued under the acronyms OR and OR20 depending of which site the material was collected. The specimens described were compared to several extant specimens from two collections: Colección Nacional de Mastozoología, at the Instituto de Biología, UNAM, Mexico City, and the Museum of Zoology, at the Biological Sciences Department at the University of Alberta, Edmonton, Canada. The graphs of measurements of the teeth were prepared using the data published by Woodburne (1969).Institutional abbreviations. IBUNAM, Instituto Biología, Universidad Nacional Autónoma de México, México City; IVIC, Colección de Paleontología, Instituto Venezolano de Investigaciones Científi cas, Caracas, Venezuela; ORS, Colección Paleontológica de El Breal de Orocual (ORS16

and ORS20), Caracas, Venezuela; UA, University of Alberta, Edmonton, Alberta, Canada.Morphological abbreviations. P/p, upper or lower premolars; M/m, upper or lower molars; d, deciduous.

SYSTEMATIC PALEONTOLOGY

Order ARTIODACTYLA Owen, 1848Suborder SUIFORMES Jaeckel, 1911

Infraorder SUOIDEA Gray, 1821Family TAYASSUIDAE Palmer, 1897

Tayassu Fischer, 1814

Tayassu pecari (Link, 1795)(Figures 2-3; Tables 1-2)

Referred material. From ORS 20, El Breal Orocual trench 20, State of Monagas: ORS20-039, lower jaw with left and right rami, with both sides of pm3-m2 and erupting m3; ORS20-040, left and right rami of a lower jaw with right pm2, pm3, dp4 and m1 and left pm2-pm3 and alveolus for right canine; ORS20-041, right lower jaw with alveoli for pm4, pm3, and pm2; m1-m2 and erupting m3; ORS20-042, left lower jaw with pm4-m3; ORS20-043, left lower jaw with m1-m2 and alveolus for pm4; ORS20-044, posterior part of left lower jaw with dp4 and m1 preserving ascending ramus and condyle; ORS20-046, fragment of right maxilla

Figure 1. Fossil localities in Venezuela. 1, Cerro Pintado and 2, Mene de Inciarte, State of Zulia; 3, Cueva del Miedo and 4, Cueva de Rio Oro, State of Falcón; 5, Cima de los Escorpiones, State of Anzoategui; 6, Cueva del Guácharo and 7, El Breal de Orocual (ORS16 and ORS20 tar pits), State of Monagas.


Figure 2. Tayassu pecari, lower jaws in different ontogenetic stages. ORS20-261, left lower jaw including mandibular symphysis of an old specimen with pm2-m3, A1, lingual, A2, labial, A3, occlusal views; ORS20-042, left lower jaw of an adult with pm4-m3, B1, lingual, B2, labial, B3, occlusal views; ORS20-40 right rami of a lower jaw of a juvenile with pm2, pm3, Dp4 and m1, C1, lingual, C2, labial, C3, occlusal views. Scale bars = 50 mm.

A1

B1

C1

B2B3

C2 C3

A2 A3

A B CFigure 3. Tayassu pecari, IVIC-P-2753, almost complete left lower jaw with c-m3, A, lingual, B, labial, C, occlusal views. Scale bars: A, B = 50 mm; C = 20 mm.

with PM4-M2; ORS20-047, left lower jaw with pm4-m1; ORS20-048, right M3; ORS20-049, fragment of left lower jaw with erupting m2; ORS20-050, fragment of right alveolus for pm2, pm3-m1; ORS20-051, fragment of right maxilla with M1 or M2; ORS20-052, right M2; ORS20-053, right M2; ORS20-054, left PM4; ORS20-055, pm4; ORS20-057, left M2; ORS20-058, right M1 or M2; ORS20-261, left lower jaw with pm2-m3 with mandibular symphysis; ORS20-311, right PM3; from Cueva del Miedo, State of Falcón: IVIC-P-2753, almost complete left lower jaw with c-m3; IVIC-P-2760, fragment of right lower jaw with dpm4-m1; IVIC-P-2761, right m3; IVIC-P-2762, fragment of right maxilla with M1-M2; IVIC-P-2763, left m3; IVIC-P-2764, fragment of a lower jaw with right m3; from Cima de los Escorpiones, State of Anzoategui: IVIC-P-2766, fragment of left lower jaw with m1-m2; from Cueva El Guácharo, State of Monagas, IVIC-P-2769, M1 or M2; from ORS16, El Breal Orocual trench 16, State of Monagas: ORS16-294, m3 fi gured in Rincón et al. (2009) as Tayassu sp. Age. Probably late Pliocene-early Holocene.

Description. The lower jaws and lower teeth were the most common elements, whereas a few upper teeth or maxillae were found (Figure 2). In the locality ORS20, nine fragments of lower jaws bearing teeth were recovered. The ontogenetic age of each individual was determined after the stage of teeth eruption (Margarido et al., 2007). According to this criteria the Venezuelan sample includes two juveniles (ORS20-040, ORS20-044); three subadults (ORS20-039, ORS20-041, ORS20-043), and three adults (ORS20-042, ORS20-050, ORS20-261), the last one corresponds to old individual. ORS20-047 and ORS20-049 are considered to be between subadult or adult stages, because the m2 is already erupted. Two specimens are from Cueva del Miedo, IVIC-P-2753, an adult (Figure 3), and IVIC-P-2760, a juvenile.

These remains are identifi ed as Tayassu pecari because the cusps are bulbous, stacked and with a wide base (Woodburne, 1968). There are also similarities in the development and position of the accessory cusps. In the lower dentition, anterior and posterior cingula in m1 and m2 are present; the lower premolars are molarized, especially the pm4, which has four cusps; the difference in height between the trigonid and talonid


Table 1. Measurements (mm) of lower dentition of Tayasu pecari. *Range of measurements taken from Woodburne (1968).

Specimenpm2 pm3 pm4 Dpm4 m1 m2 m3

L W L W L W L L W L W L WIVIC-P-2753 9 5.3 10 6.7 12 8.7 14.7 11 15.5 11.6 19 11.6IVIC-P-2760 12 11 8.6IVIC-P-2761 22.9 15IVIC-P-2761 22.8 15IVIC-P-2763 21.6 18.2IVIC-P-2763 22 13.2IVIC-P-2764 16.7 10.4IVIC-P-2764 17.6 10.7IVIC-P-2766 15.5 11.6 16.5 12.7ORS16-294 23.2 13.4ORS20-039l 10 7 12 9 14 11.2 15.7 14ORS20-039r 10 6.5 12.6 9.5 14 11.2 15.7 12.8ORS20-040 7.7 4 8.5 5.6 16 15 10.3ORS20-041 12.9 9.6 14.5 11.8ORS20-042 11.7 8.6 13.1 11.2 15.7 12.7 20 12.1ORS20-043 12.1 10 13.9 11.2ORS20-044 14.2 12.2 9.2ORS20-046 7.4 3.7 10 5.7 brokenORS20-047 12.5 9.3 15 12ORS20-049 15.8 11.7ORS20-050 8.2 6.7 11 8.6 11.6 9.9ORS20-055 11 10ORS20-261 8.5 6 10.5 8.6 12 10.5 13 11.1 15.4 13 18 18PZUNNE 10 5.95 11.3 7.55 13.55 11.2IBUNAM 15182 9.7 6.5 10 7.3 12.3 9.2 14.7 11 14.6 12.4 17.9 11.8IBUNAM 15183 9.3 6.8 10.8 7.2 13 9.7 14.9 14.9 11 15.3 12.8 19.5 12.6IBUNAM 36704 9.4 5.7 11.1 6.5 13.1 9.6 13.8 10.8 15 12.6 17.9 12.2

Tayassu pecari* 8.75-10.60

4.95-6.60

9.55-12.10

6.45-8.70

11.7-14.0

9.20-12.65

14.0-16.25

10.9-14.4

15.15-18.35

12.35-16.6

18.45-24.30

12.65-16.20

Table 2. Measurements (mm) of upper dentition of Tayasu pecari. *Range of measurements taken from Woodburne (1968).

SpecimenPM2 PM3 PM4 M1 M2 M3

L W L W L W L W L W L WIVIC-P-2762 14.6 12.9 15.6 13.6MBLUZ-P-5.123 16.3 14ORS20-046 12 11 13.4 10.5 15.3 12.6ORS20-051 15.3 12.4ORS20-052 15.7 14ORS20-053 15.3 13ORS20-054 12 10.4ORS20-057 16.6 13.7ORS20-058 17.4 14.7ORS20-311 12.7 12IBUNAM1046right 8.8 8 9.8 10.5 10.7 9.7 13.6 11.5 15.3 12.6 14.8 12IBUNAM1046left 8.7 8 10.4 9.8 10.3 10 13.8 11.7 14.8 14.2 15.3 13.4IBUNAM36704 8.6 8.5 10 11 11.5 11.5 13.4 12 15.4 14 17.2 14.4IBUNAM15183 8.4 8.4 9.6 9.6 11 11 13.6 11.8 15.6 14.5 16.8 13.4IBUNAM15182 8 8 10 10 11.3 11.2 14.7 11.6 14.6 13 15.7 12.4

Tayassu pecari* 8.5-11.5 8.45-10.55

9.80-12.30

9.90-12.25

10.85-13.45

11.15-13.80

12.65-16

12.15-15.60

11.95-17.70

14.85-19.3

14.85-19.30

13.65-19.65


is not as pronounced as in Pecari. The cusps of the pm4 are as high as those of m1. The last lower molar is longer than wide. For measurements of the lower dentition, see Table 1.

In the upper dentition, the P4 is quadrangular, with four cusps; the upper molars have well developed anterior and posterior cingula; the accessory cusps cross the transverse valley. ORS20-054 is identifi ed as a molarized P4. In contrast to the M1, its posterior side is straight and the anterior is more convex, the anterior and posterior cusps are present, and the accessory cusps are placed anteriorly. The M3 ORS20-048, is longer than wide, the anterior part is wider than the posterior, and it bears a strong anterolingual cingulum. For measurements of the upper dentition, see Table 2.Discussion. The morphology of the lower and upper dentitions is characteristic of the genus Tayassu. However, the size of the Venezuelan dental series and isolated teeth does not fi t exactly within the size range of Tayassu pecari (Woodburne, 1968; the present data based on specimens deposited in UA and IBUNAM). In general, most of the Venezuelan specimens are smaller than Tayassu and larger than Pecari (Figures 4,5).

The premolars of the juvenile specimen (ORS20-040) fall within the range size of Pecari but the molars do not. The teeth of the subadults (ORS20-041 and ORS20-043) fi t in the range of the largest Pecari; in the adult (ORS20-039) one tooth falls in the range of Pecari, but the other teeth in that of Tayassu. ORS20-261, left lower jaw with complete dental series, shows some teeth in the size range of Pecari and others in the Tayassu.

It is interesting to notice that considering the length, the upper dentition falls within the range of of Tayassu, but considering the width they fall in the range of P. tajacu (Figure 5).

Woodburne (1968) and later Frailey & Campbell (2012) noted that there are differences between Pecari and Tayassu in the shape of the lower jaw. The post digastric sulcus lies below the m3 in Tayassu, and below m2-m3 in Pecari. In the Venezuelan material identifi ed as Tayassu, in the old individuals (ORS20-261, IVIC-P-2753) it lies below m3, but in ORS20-042 (an adult) it lies between m2-m3. In the three specimens, the sulcus is not pronounced, so maybe this character is variable during the ontogenetic development. In IVIC-P-2765 identifi ed as Pecari tajacu, this structure is not pronounced and it lies below the m3 (Figures 2-3,6).

According to Woodburne (1968) the dental dimensions of Tayassu pecari and Pecari tajacu usually overlap, but he considered the length and width of m1, length of m2 and width of m3 to be useful to distinguish the two genera. In the Venezuelan sample there are some specimens (ORS20-044, ORS20-043, ORS20-041, ORS20-050, IVIC-P-2761, IVIC-P-2764 and IVIC-P-2753, see Table 1, Figures 4,5) with these teeth smaller than those of T. pecari. Comments. Currently, Tayassu pecari has its southernmost geographic distribution in Argentina, and it´s northernmost in southern México. In Venezuela, this species now inhabits the evergreen forests in the state of Zulia, and the deciduous and semideciduous forest areas of the states of Cojedes, Carabobo, Sucre, Monagas and Bolívar, with a more restricted distribution than that of Pecari tajacu (Bisbal & Jiménez,

1995). During ?Plio-Pleistocene times T. pecari was present in the states of Falcón and Anzoátegui, where now is absent.

Pecari Reichenbach, 1835

Pecari tajacu (Linnaeus, 1758)(Figure 6; Tables 3-4)

Referred material. From ORS 20, El Breal Orocual trench 20, State of Monagas: ORS20-045, fragment of lower jaw with alveoli for pm2-pm3, and erupting pm4, part of diastema canine-pm; from Cueva Rio de Oro, State of Falcón: IVIC-P- 2765, fragment of right lower jaw with pm4-m3 and an associated incomplete skull, lacking muzzle, with the left zygomatic arch broken;and preserving the right upper teeth series PM2-M3, left PM4-M3; Mene de Inciarte, State of Zulia: IVIC-P-2768 fragment of right maxilla with M1-M3.Age. Late Pleistocene.Description. The fragment of the lower jaw (ORS20-045) is identifi ed as Pecari tajacu because the erupting pm4 is molariform: the anterior cuspids are higher and more strongly developed than the posterior ones. IVIC-P-2765 collected in the Cueva Río de Oro (Figure 6), is a small specimen (see measurements in Table 3) including the right dentary and a partially preserved skull. The dentary includes pm4-m3 with worn m1 and m2 indicating it is from a quite old adult. The partial skull lacks the muzzle, has a broken left zygomatic arch; and includes right PM2-M3, and left PM4-M3. In the right row, a tooth is erupting medial to the PM3. This is an anomalous condition because all the upper teeth are permanent and this specimen corresponds to an old individual. The morphology of the dental series corresponds to that of Pecari, but the most anterior premolar shows a more Tayassu-like shape, it is rounded and is not triangular as those of Pecari (Figure 6). In addition, the cusps of m3 are not bulbous as in T. pecari. The fragment of maxilla with M1-M3 collected in Mene de Inciarte (IVIC-P-2768) is identifi ed as P. tajacu because the fi rst molar is subquadrangular in outline, the four major cusps are developed and paracone and metacone are slightly anterior to their lingual counterparts. A transverse valley separates the anterior from the posterior areas and it is somewhat interrupted, in labial profi le, by the metaconule. Labial and lingual cingula are present. The second upper molar is larger than the fi rst, has a strong labial and an incipient lingual cingulum, but otherwise as the M1. In the M3, the cingula are well developed and the shape is triangular, narrowing posteriorly, differing from T. pecari where this tooth is more rectangular. They basically follow the morphology described by Woodburne (1969) and fi t within the size range of P. tajacu (Figure 5). For measurements of upper dentition see Table 4.Comments. It is curious that among all the material collected from ORS20 there is only one lower fourth premolar referred to Pecari tajacu, but it bears all the characteristics of this species: the distinct difference in height between trigonid and talonid and its size fi ts perfectly in the size range of P. tajacu. Currently, this species is more widely distributed


Figure 4. Graphs of length and width of lower molars of the Venezuelan tayassuids within the range of size of Woodburne (1968).


Figure 5. Graphs of length and width of upper molars of the Venezuelan tayassuids within the range of size of Woodburne (1968).


Figure 6. Pecari tajacu. IVIC-P-2768 fragment of right maxilla with M1-M3, A1, lingual, A2, labial, A3, occlusal views; IVIC-P- 2765, fragment of right lower jaw with pm4-m3, B1, lingual, B2, labial, B3, occlusal views; incluir IVIC-P-2765, incomplete skull, C1, dorsal, C2, lateral, C3, occlusal view of right upper teeth series PM2-M3, left PM4-M3. Scale bars: A1-A3 = 10 mm; B1-B2, C1-C3 = 50 mm; B3 = 30 mm.

A1

B1

C1 C2 C3

B2 B3

A2 A3

than Tayassu pecari, extending northward to the southwestern part of United States. In Venezuela, it is found through all the country, except in the Andean region. It inhabits deciduous, semideciduous and evergreen forests of the Cordillera de la Costa, Andean foot plains, Amazonian, savannas and the forests along the rivers in Los Llanos (Bisbal & Jiménez, 1995). Currently, there is no record of P. tajacu in the State of Zulia.

DISCUSSION

Gasparini (2013) summarized the history of the South American Tayassuidae but did not include the Venezuelan record of Platygonus (Rincón et al., 2009). According to Gasparini (2013) the oldest record of Platygonus dates from middle Pliocene and during early-middle Pleistocene its distribution decreased, and its youngest and last record dates from the Toscas del Rio de La Plata, Buenos Aires Province, Argentina (Soibelzon et al., 2008). The oldest record of Catagonus dates from late Pliocene? and it is still present in the Gran Chaco of Argentina, Paraguay and Bolivia. As a fossil, Pecari tajacu had been found in Pleistocene faunas in Guatemala (Woodburne, 1969) and Colombia (Porta,

1969; Villarroel et al., 1989) (Figure 7). Pecari tajacu has a late Pleistocene record in Florida (Hulbert et al., 2009) and also is known from archaeological sites of Holocene age (Czaplewski, 2012).

The oldest record of Tayassu pecari dates from middle Pleistocene in South America; it has been recorded mainly in Argentina, with a few records in Brazil, Uruguay, and Colombia (Porta, 1969; Paula Couto, 1975, 1981; Ubilla, 2004; Ubilla et al., 2004; Gasparini & Ubilla, 2011; Gasparini et al., 2013) (Figure 7). The presence of Tayassu from El Breal Orocual, which is probably late Pliocene – early Pleistocene, represents the oldest record of this taxon in South America.

Both genera (Tayassu and Pecari) have been found as subfossils in Holocene deposits in Peru, Argentina, Venezuela, Panamá and Brazil (Cartelle & Hartwig, 1996; Eisenberg et al., 1979; Gasparini et al., 2013; Leigh & Wright, 1990; Terborgh, 1990; Politis & Messineo, 2008).

The origin of the genera Tayassu and Pecari has been diffi cult to establish. So far there are two hypotheses about the origin and divergence of these tayassuids: (i) the tayassuids migrated from North America to Central America and South America where they diversifi ed via adaptive radiation during the Pleistocene less than 2 million years ago (Wetzel, 1977;


Table 3. Measurements (mm) of lower dentition of Pecari tajacu. *Range of measurements taken from Woodburne (1968).

Specimenp2 p3 p4 m1 m2 m3

L W L W L W L W L W L W

ORS20-045 11.5 7

IVIC-P-2765 10 8 11 8.7 13 9.7 14.9 10

IBUNAM 41107 6.7 4.2 7.9 6 9.5 8 11.1 8.8 12 9.6 14.6 10

IBUNAM 335 8.1 5.5 8.5 6 11 8 11.6 9 13.8 12 17.5 12

IBUNAM 24580 7.7 4.2 9.9 6 11 9 12.3 9.3 13 10.6 15.5 10.4

UA 3223right 6 4 7.8 5.1 10.4 8 12.4 9.8 14.6 11.4 15.2 11.4

UA 771 7.4 4 7.5 5.5 9.3 7 12 9.8 14.9 10.3

UA 7985left 6 3 8 4.9 13.5 8 11.4 9 12.8 11.4

UA 7985right 5.5 3 11.4 9 12.9 11.4

UA3223left 8 5.1 10.4 8 11.4 10 14.5 12 15.6 12

UA5247left 7.3 + 8.5 5 12 8 11 10.5 13 11 12.9+ 10

UA5247right + 5.5 9 6 10.3 9 11.5 10.5 13 11.5 17.2 12

UA7986left 8 8 10 10 12.5 10.5 14 9

UA7986right 8.5 7 10 9.5 12.5 10.5 13.2 9

Pecari tajacu* 7.1-8.95 3.8-5.45 7.60-10.35

5.15-7.20 9.0-12.4 6.35-

9.9510.30-13.3

8.0-10.80

12-14.95

9.75-13.0

12.30-20.5

9.45-12.40

Table 4. Measurements (mm) of upper dentition of Pecari tajacu. *Range of measurements taken from Woodburne (1968).

SpecimenP2 P3 P4 M1 M2 M3

L W L W L W L W L W L W

IVIC-P-2768 12 10.1 13.4 10.9 11.9 10

IVIC-P-2765r 7 6.9 8.1 7.2 8.9 8.9 9.6 10 13 11 15 12

IVIC-P-2765l 8.8 9 10.3 9.5 12.7 10.6 15.1 10.8

IBUNAM 3358.8 8 8.3 9.3 10 11 11.4 11.4 12.2 11.8 14.6 13.5

IBUNAM 245807.9 7.2 9.2 9 10 10.5 11.6 9.6 12.5 10.6 13.6 12.2

IBUANM 411077.3 7 7.7 9.4 8.7 9.5 10.6 10 11.7 10.4 12.4 11

UA 3223left 7 7.3 8.1 9.2 9.3 10 12 11 13.6 11.8 11.5 11

UA 3223right 7.2 7.2 7.3 7 9.3 10 11.4 11 13.5 12 11.4 10

UA 5247right 7.2 8.5 8.5 8.4 9.7 10.5 11 11.5 12.5 12.5 14.05 11.5

UA 771left 6.1 5.7 8 8.5 9.5 10.3 12 11.4 13.2 12.5

UA 771right dp2? 7 6 9 9.5 10 10 11.6 11

UA5247left 9 8.5 9.3 9.8 10.5 10.9 11.3 12.5 12

UA7982left 7 5.5 9 7 9.1 8.2 11 10 12.1 11

UA7982right 7 5 9.5 7 10 7.5 11 9.8 13 11

UA7986left 7.1 6.8 8 8.1 8.8 9.1 9 10 12 10.5 11 10

UA7986right 7.5 6.5 8.1 8.1 8.1 9.1 9.5 10.05 12.1 11 12 9.5

Pecari tajacu* 6.95-9.7 5.75-9.05 8.0-10.2 7.35-

10.55 8.9-11.5 8.70-11.7

9.80-12.85 9.10-13 11.10-

14.4010.05-14.35

11.50-16.50

10.30-13.20


Recent molecular phylogenetic analyses suggested that Tayassu and Catagonus are more closely related to each other than to Pecari (Theimer & Keim, 1998; Góngora & Morán, 2005). Based on global nucleotide substitution rate Theimer & Keim (1998) estimated that the divergence between Catagonus and Tayassu occurred between 1.7-2.3 Ma and 3.2-7.4 Ma between Pecari and Catagonus/Tayassu. Góngora & Morán (2005) suggested two possible ranges for the earlier event, 7.7-6.2 Ma and 13.2-4.3 Ma (these two ranges resulting from different methodologies). Based on their chronology, the fi rst split between Pecari and Catagonus/Tayassu sometime during the Miocene-Pliocene, would have taken place after the arrival of their last common ancestor to South America. If the age and the relationships of the fossils Peruvian tayassuids are confi rmed, the timing of the splitting of the two lineages implied by molecular methods should be earlier.

Unfortunately, the new Venezuelan records do not provide any information in pro or against the proposal of whether the extinct and living genera of tayassuids dispersed to South America from Central America as distinct lineages in the late Miocene in multiple dispersal events, or diverged in South America, after a single dispersal event, this issue remains unclear. However, the presence of these taxa in Venezuela testifi es to the presence of well-established populations and a wide distribution probably from the late Pliocene.

CONCLUSIONS

The fossil localities in Venezuela document the northern-most South American biotas present in South America during the Great American Biotic Interchange, which is otherwise absent in low tropical latitudes.

The description of the peccaries from probably late Pliocene-Quaternary of Venezuela is signifi cant and adds another taxon to the list of faunal assemblage from this part of South America that was previously practically unknown. If the age of late Pliocene of ORS 16 is confi rmed, the presence of Tayassu pecari in Venezuela would represent the oldest record of this taxon in South America. Compared to the living specimens of Tayassu pecari, the Venezuelan specimens are smaller.

ACKNOWLEDGMENTS

The authors thank the Instituto Venezolano de Investi-gaciones Científicas (Project Paleomapas de Venezuela #1096), and Universidad Nacional Autónoma de México, through the Programa de Intercambio Académico of the Coordinación de Investigación Científi ca, for supporting the fi eld work in Venezuela and the visits of MMB to Caracas (2010, 2012 and 2013) and the visits of AR and AS to Mexico City (2010, 2011). Thanks to H.G. McDonald for correcting the English and suggestions to the final manuscript; to G. Parra, A. Carballo, Petróleos de Venezuela S.A, (PDVSA) and Instituto de Patrimonio Cultural, Venezuela, for the facilities during the development of the project. Finally, thanks to D. Prothero and an anonymous reviewer who provided comments and new information that improved the manuscript.

Figure 7. Fossil localities with Tayassu and Pecari remains in South and Central America.

Woodburne, 1968, 1969); (ii) these peccaries diverged in North America during the late Miocene less than 10 million years ago before they independently colonized South America (Wright, 1989).

Unfortunately, the fossil record of tayassuids in Central America is very poor, with the Pleistocene record of Pecari tajacu in Guatemala (Woodburne, 1969), the early-middle Miocene record of Cynorca in Panama (MacFadden et al., 2010), and the Hemphillian indeterminate tayassuid in Costa Rica (Laurito & Valerio, 2012). The fossil record in Central American therefore, does not currently provide any information that can help to elucidate the origin and evolution of the tayassuid lineages in South America. If we accept the Miocene age of the two new tayassuid forms described from Peru: Waldochoerus and Sylvachoerus (Frailey & Campbell, 2012) and that they are close related to the two modern lineages Pecari and Tayassu, this would imply that these two genera were already separated since late Miocene in South America.

Another issue that remains controversial is the relationship between the species and genera of tayassuids. Based on cranial and dental characters, Wetzel et al. (1975) and Wetzel (1977) proposed that Tayassu and Pecari are more closely related to each other than to Catagonus. In contrast, based on his analysis using osteological and dental characters Wright (1989, 1993, 1998) considered Pecari and Catagonus to be more closely related, whereas Tayassu was considered to be a member of a separate clade along with other extinct species.


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Received in January, 2014; accepted in July, 2014.

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RECORD OF TAYASSUIDS IN ?LATE PLIOCENE TO QUATERNARY … · 2020-03-19 · As one of the North American immigrants that entered South America during the Great American Biotic Interchange,