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New material of Tapirus (Perissodactyla: Tapiridae) from the Pleistocene of

southern Brazil

Article  in  Revista Mexicana de Ciencias Geológicas · November 2012

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308 Revista Mexicana
Holanda et al. de Ciencias Geológicas, v. 29, núm. 2, 2012, p. 308-318

New material of Tapirus (Perissodactyla: Tapiridae) from

the Pleistocene of southern Brazil

Elizete C. Holanda1,*, Ana Maria Ribeiro2, and Jorge Ferigolo2

1
Programa de Pós-Graduação em Geociências, Universidade Federal do Rio Grande do Sul,
Campus do Vale, Caixa Postal 15001, CEP 91501-970, Porto Alegre, RS, Brazil.
2
Museu de Ciências Naturais, Fundação Zoobotânica do Rio Grande do Sul,
Av. Dr. Salvador França, 1427, Jardim Botânico, CEP 90690-000, Porto Alegre, RS, Brazil.
* [email protected]

ABSTRACT

New material of Tapirus from the late Pleistocene of Rio Grande do Sul State is described, and
other specimens previously collected are reviewed. The material comes from Alegrete (Sanga da Cruz
Creek), Dom Pedrito (São Luiz cattle ranch), Iraí, Quaraí (Cerro da Tapera), and Santa Vitória do
Palmar (Hermenegildo Beach). The specimens from Iraí are referred to the extant species, T. terrestris,
on the basis of their size and morphology. Specimens from Sanga da Cruz Creek, Hermenegildo Beach,
São Luiz cattle ranch and Cerro da Tapera are very fragmentary or present no diagnostic characters,
and are tentatively assigned to Tapirus cf. T. terrestris. We confirm the presence of the T. terrestris in the
upper Pleistocene of Rio Grande do Sul state. Currently, there are neither archaeological nor historical
records of this species for the southern region of this state.

Key words: late Pleistocene, Perissodactyla, Tapirus, Rio Grande do Sul, Brazil.

RESUMEN

Nuevos materiales del género Tapirus de Pleistoceno tardío del Estado de Rio Grande do Sul son
presentado, mientras otros especímenes previamente recolectados son revisados. El material descripto
proviene de las municipalidades de Alegrete (Sanga da Cruz), Dom Pedrito (Estancia São Luiz), Iraí,
Quaraí (Cerro da Tapera), y Santa Vitória do Palmar (Playa del Hermenegildo). Los especímenes de las
localidades de Iraí fueron comparados con otras especies del género, y debido a su morfología y tamaño,
ellos fueron referidos a la especie reciente, T. terrestris. Los especímenes de Sanga da Cruz, Playa del
Hermenegildo, Estancia São Luiz y Cerro da Tapera están muy fragmentados o no presentan caracteres
diagnósticos, y fueron tentativamente asignados a Tapirus cf. T. terrestris. Nosotros confirmamos la
presencia de T. terrestris en el Pleistoceno tardío del Estado de Rio Grande do Sul. Actualmente, no hay
registros arqueológicos ni históricos de esta especie para la región sur de este estado.

Palabras clave: Pleistoceno tardío, Perissodactyla, Tapirus, Rio Grande do Sul, Brasil.

Holanda, E.C., Ribeiro, A.M., Ferigolo, J., 2012, New material of Tapirus (Perissodactyla: Tapiridae) from the Pleistocene of southern Brazil: Revista
Mexicana de Ciencias Geológicas, v. 29, núm. 2, p. 308-318.
 Tapirus from the Pleistocene of southern Brazil 309

INTRODUCTION Soliani (1973) identified remains of T. terrestris at

Arroio Chuí, and Oliveira (1992) cites T. terrestris from the
Family Tapiridae contains a single extant genus, Coastal Plain, but specimens mentioned by these authors
Tapirus Brisson, 1762, with four extant species; one species could not be located in any collection, and at moment, it is
inhabits southeastern Asia and three are found in Neotropical not possible to confirm the presence of tapirs at this site.
America (Brooks et al., 1997). Tapirs reached South America Paula Couto (1943) recorded the first fossil tapir from Iraí
during the Great American Biotic Interchange, where they Municipality, a lower molar, probably m3. Souza Cunha
are today the largest terrestrial mammals (Stehli and Webb, (1959) reported postcranial fragments of T. terrestris, also
1985). The species of Tapirus live in the neotropical zone from Iraí; specimens that are re-described in this study.
and are associated with humid tropical and subtropical for- Holanda et al. (2005) recorded the first tapir from Dom
est. They are browsers and frugivores, playing a critical role Pedrito Municipality, described herein.
in structuring of tropical forests as seed dispersers (Fragoso
and Huffman, 2000), and are indicators of ancient forested
environments (DeSantis and MacFadden, 2007). GEOLOGICAL SETTING
Some species of Tapirus have been described from
Pleistocene deposits of South America based on mandibular The tapir material described herein comes from five
and dentary fragments. These are T. tarijensis Ameghino, localities in Rio Grande do Sul state. Sanga da Cruz Creek
1902 (Tarija, Bolivia, dubious stratigraphic provenance), is a tributary of the Ibicuí River, situated 23 km north of
T. rioplatensis Cattoi, 1957 (Buenos Aires Province, Alegrete City (Da Rosa, 2003; Ribeiro and Scherer, 2009;
Argentina - early Pleistocene) and T. oliverasi Ubilla, 1983 Figure 1). The specimen described here comes from Sanga
(Department of Montevideo, Libertad Formation, Uruguay da Cruz, however the precise point of collection is lacking.
– early Pleistocene). Most mammals from Sanga da Cruz are found in con-
Three late Pleistocene species have been described glomeratic facies that yielded a thermoluminescence date
based on cranial material; T. cristatellus Winge, 1906 between 14,925 ± 800 and 14,830 ± 750 years BP (Milder,
from Minas Gerais and Bahia states, Brazil (Cartelle, 2000; Da Rosa, 2003). At another point near the outfall of
1999; Holanda et al., 2007), T. mesopotamicus Ferrero and the Ibucuí River is a mudstone facies where Miller (1987)
Noriega, 2007 from the Mesopotamia region of Entre Rios reported Glossotherium robustum, and this yielded a 14C
Province, Argentina, and T. rondoniensis Holanda, Ferigolo date of 12,770 ± 220 years BP.
and Ribeiro, 2011 from southwestern Amazonia. The Cerro da Tapera outcrop is situated on the right
Late Pleistocene records of T. terrestris (Linnaeus, margin of the Quaraí River, Quaraí Municipality, about 200
1758) are known from many places in Argentina, Brazil, m downstream from the outfall of Cati Creek (30º51’98”S,
Uruguay and Venezuela (Paula Couto, 1980; Rancy, 1981; 56º29’30”W; Figure 1). The deposit here is typically fluvial,
Simpson and Paula Couto, 1981; Tonni, 1992; Ubilla, with several levels of fine to medium sandstone and brown
1996; Hyrooka, 2003; Porpino and Santos, 2003; Salles et mudstone (Ribeiro et al., 2008). The fossils collected at
al., 2006; Ferrero et al., 2007; Holanda and Rincón, 2009; this locality are isolated fragments of Propraopus gran-
Oliveira, 2010). In addition to these records, fragmentary dis, Morenelaphus sp., Glyptodontidae, Mylodontinae,
material from the late Pleistocene of Uruguay, Argentina, Toxodontidae, Cervidae, Carnivora and indeterminate frag-
Brazil, Peru and Venezuela have been referred to Tapirus, ments (Ribeiro and Scherer, 2009). According to Ribeiro
but without specific determination (Rusconi, 1928; Cattoi, et al. (2008) these deposits yielded thermoluminescence
1951; Rolim, 1974; Ubilla, 1983; Marshall et al., 1984; dates from two levels of 11,000 ± 2,000 and 13,000 ± 2,150
Sedor et al., 2004; Holanda and Cozzuol, 2006; Holanda years BP.
and Rincón, 2009). The São Luiz cattle ranch is a private property within
the limits of the Dom Pedrito Municipality, in southern
Rio Grande do Sul state. A fossil tapir specimen was
Previous studies of Tapirus in southern Brazil found here associated with Stegomastodon waringi. Many
Glyptodontidae osteoderms were recovered at the same
Specimens of Tapirus from Rio Grande do Sul state locality. At Iraí, tapir fossils were found during an excava-
are scarce, incomplete, and comprised almost exclusively of tion for extraction of mineral water in 1936 (Tupi Caldas,
dental fragments and postcranial remains. Bombin (1976) 1939); material here was also associated with S. waringi
referred late Pleistocene material from the Touro Passo according to Gadens Marcon (2008). No biostratigraphic or
Formation (Uruguaiana Municipality) to the species T. ter- geochronologic studies have been done for these two sites,
restris, but specimens mentioned by the author could not be but both are stated to be late Pleistocene in age (Ribeiro
located in any collection. The presence of fossil tapirs from and Scherer, 2009).
Touro Passo Creek was confirmed by Kerber and Oliveira “Hermenegildo Beach” on the Coastal Plain, is part of
(2008) at the Ponte Velha I outcrop (Figure 1), based on the Pelotas Basin that contains deposits of an extensive sys-
lower molar remains. tem of alluvial fans, and four lagoon-barrier systems, known
310 Holanda et al.

Figure 1. Map of the localities where reported

fossil tapir were in Rio Grande do Sul state.
Black triangle is previously known locality and
grey triangles are localities reported here. 1.
Sanga da Cruz Creek; 2. São Luiz cattle ranch;
3. Iraí; 4. Cerro da Tapera; 5. Hermenegildo
Beach; 6. Ponte Velha I; a. “line of Paca” of
Ihering (1892).

as I to IV (Villwock and Tomazelli, 1995). Lagoon-Barrier specimens (see Appendix). The dental terminology follows
System III represents the third transgression/regression Butler (1952) and the taxonomy follows Colbert (2005).
sequence of the Pleistocene. Tapir fossils were exhumed The measurements follow Simpson (1945), Hue (1907)
at the beach, on Hermenegildo’s parcel (53°15’S and and Hulbert (2005). The descriptive statistics were done
33°42’W; Lopes et al., 2001). The lagoonal depression that using the most recent available version of Palaeontological
is now drained by the Chuí Creek, where mammalian fos- Statistics – PAST software (Hammer et al., 2001).
sils are found, is correlated with this system (Villwock and The abbreviations used in this study are as follows:
Tomazelli, 1995). Recently, Lopes et al. (2010) performed DGM, Departamento Nacional de Produção Mineral, Rio
Electron Spin Resonance dating on fossils from Chuí Creek de Janeiro, Brazil; MACN, Museo Argentino de Ciencias
and the coastline. The Chuí Creek specimens yielded ages Naturales Bernardino Rivadavia, Buenos Aires, Argentina;
between 42 and 38 ka, much younger than previously esti- MCN, Museu de Ciências Naturais, Fundação Zoobotânica
mated (Lopes et al., 2001). Specimens from the coastline do Rio Grande do Sul, Porto Alegre, Brazil; MCP, Museu
produced a wide age range, around 6 x 104 years, indicating, de Ciências of Pontifícia Universidade Católica do Rio
according to the authors, that the fossils came from deposits Grande do Sul, Porto Alegre, Brazil; MLP, Museo de La
of different ages that were successively reworked and re- Plata, Argentina; MN, Museu Nacional do Rio de Janeiro,
deposited in younger sediments by sea-level oscillations. Brazil; MZUSP, Museu de Zoologia of Universidade de
The associated mammalian fauna includes the following: São Paulo, Brazil.
Tardigrada, Cingulata, Litopterna, Notoungulata, Carnivora,
Rodentia, Proboscidea, Artiodactyla and Perissodactyla
(Oliveira, 1992; Buchmann, 2002; Ribeiro and Scherer, SYSTEMATIC PALEONTOLOGY
2009).
Order Perissodactyla Owen, 1848
Family Tapiridae Burnett, 1830
MATERIALS AND METHODS Genus Tapirus Brisson, 1762
Tapirus terrestris (Linnaeus, 1758)
New material described herein is housed in the paleo- (Figure 2, Tables 1-3)
vertebrate collections of Museu de Ciências Naturais (MCN)
and Museu de Ciências e Tecnologia (MCP). The specimens Referred material and provenance. MN2099-V, incom-
from Iraí reported by Souza Cunha (1959) are re-described plete left humerus; MN2098-V, right tibia; DGM79M, left
here. The material was compared to both recent and fossil calcaneum, Iraí Municipality.
 Tapirus from the Pleistocene of southern Brazil 311

Figure 2. Tapirus terrestris, a - b: MN2099-V, left humerus in cranial and caudal views (left to right); c-f: DGM79M, left calcaneum in dorsal, plantar, medial
and lateral views (left to right); g-j: MN2098-V, right tibia in cranial, caudal, proximal and distal views (left to right and top to bottom). Abbreviations:
c: capitulum; cf: coronoid fossa; ht: humeral trochlea; ie: intercondylar eminence; lc: lateral condyle; mc: medial condyle; mm: medial malleolus; of:
olecranon fossa; pl: popliteal linea; pn: popliteal notch; se: sulcus extensorius; tc: tibial crest; tco: cochlea; tt: tuberosity tibial; a: medial astragalar facet;
b: proximal astragalar facet; c: distal astragalar facet; t: tuberosity; 1: coracoid process; 2: sustentacular process. Scale bar = 30 mm.

Description. The distal end of a left humerus MN2099-V is oval and flat, and is thicker and narrower than the lateral
has the condyles slightly worn (Figure 2a - b, Table 1); the one, which is nearly circular and craniocaudally convex. The
distal articular face is semi-cylindrical and oblique, higher condyles are separated posteriorly by a poorly visible pop-
medially than laterally. The humeral trochlea is slightly liteal notch, and axially by the intercondylar eminence. The
rectangular, has a pronounced constriction on its centre, and intercondylar eminence is formed by a high and laminated
is wider and higher than the capitulum, which is narrow and medial tubercle, and by a low and robust lateral tubercle,
low. Both the coronoid and olecranon fossae are covered which are separated by an irregular central depression. The
by sediment, but it is possible observe that the olecranon tibial tuberosity is large and rounded. Extending distally
fossa is narrower and higher than the coronoid fossa. The from the tibial tuberosity is the tibial crest, which has a
ectepicondyle is low and prominent, and is lateral directed. noticeable medial inclination. The lateral surface of the
The lateral supracondyloid crest extends to the caudodistal tibia is smooth and slightly sprained in spiral. The medial
margin of the ectepicondyle. The entepicondyle is robust surface is broad and flat on the proximal half, and narrow
and high. and convex on the distal half. The caudal surface is narrow
The tibia MN2098-V is more complete but the ex- and is divided in two parts by an oblique popliteal line. The
tremities are slightly abraded (Figure 2g-j; Table 2). Its proximal region of the popliteal line has a rugous concave
proximal half is slightly triangular and nearly cylindrical. area. The distal region of the popliteal line has a smooth
The proximal end is relatively broad. The medial condyle and convex area. The distal end is slightly rectangular
312 Holanda et al.
Table 1. Measurements from the humeri of Tapirus. Abbreviations: CDD: is deep and rugose. In the plantar surface there is a slightly
craniocaudal diameter of distal portion; MDD: mediolateral diameter of visible groove. On lateral surface, distal to the coracoid
distal portion; TMD: mediolateral diameter of humeral trochlea; where N
> 1, first line is mean ± standard deviation, second line is observed range, process, there is a rough and slightly rounded projection
and third line is number of observations (N). Data of T. cristatellus from for the insertion of the external ligament.
Winge (1906) and the cave provenance. Remarks. The distal end of the left humerus (MN2099-V)
CDD MDD TMD was erroneously considered by Souza Cunha (1959) to be
a femur. The postcranial materials from Iraí are similar
T. terrestris 60.38 ± 3.45 68.64 ± 2.92 41.05 ± 2.33 in size and morphology to those of T. terrestris. The only
55.3 - 67 64.4 - 75.24 37.5 - 46
(16) (16) (16) fossil tapir with preserved postcranium known from the
Pleistocene of South America is T. cristatellus. Almost all
MN2099-V 63.1 70 42
measurements of T. cristatellus are much larger than for T.
T. cristatellus -- 64.5 --
terrestris (Tables 1-3).
Escrivania 1
Comparing the humerus (MN2099-V), tibia (MN
T. cristatellus -- 62 --
2098 PV) and calcaneum (DGM79M) measurements, this
Escrivania 5
material is similar in size with T. terrestris in all measure-
ments (Tables 1-3). The specimens are very gracile, as in
T. terrestris. The tibia and calcaneum measurements are
and much smaller than the proximal end. In the astragalar smaller than for T. cristatellus (Tables 2-3). Although T.
notch, facets for the proximal trochlea of the astragalus are pinchaque and T. bairdii have considerable overlap in their
deep and asymmetric. These facets are separated sagittally dental measurements with T. terrestris (Simpson, 1945),
by a low and convex ridge. This ridge extends between the postcranium of T. pinchaque (Roulin, 1829) has been
two processes located at the cranial and caudal margins of described as smaller than in T.terrestris. T. bairdii (Gill,
the cochlea. The medial malleolus is very prominent and 1865) and T. indicus Desmarest, 1819 are larger than T.
forms the craniomedial wall of the astragalar notch, which terrestris (Simpson, 1945 – including measures from T.
is slightly inclined in the direction of the facet. bairdii; Eisenmann and Guérin, 1992).
The calcaneum DGM79M is long and thin, and its
ends are lightly abraded (Figure 2c-f, Table 3). The distal Tapirus cf. T. terrestris
half is transversely wide, and the axial portion is thickened (Figures 3-5, Tables 4-5)
on the facet for the cuboid. The tuberosity is quite promi-
nent, mediolaterally compressed and dorsoplantarly thick. Referred material and provenance. MCP-4290-PV,
On the more axial face, the tuberosity is rugose and con- fragment of left maxilla, São Luiz cattle ranch; MCN-
vex. On the distal half, the dorsal surface has three facets PV-3515, MCN-PV-6968, MCN-PV-8085-8087, dental
for the astragalus. The sustentacular process is a medial crowns, MCN-PV-8843, symphyseal portion of mandible,
projection and has an oval concave facet for the astragalus. MCN-PV-10069, proximal portion of left metatarsal III,
Dorsoproximally, the coracoid process holds a concave Hermenegildo Beach; MCN-PV-9700, right scapula, Cerro
proximal facet, and its proximomedial border is convex. The da Tapera; MCN-PV-7071, left metacarpal II, Sanga da
distalmost and smallest articular surface for the astragalus Cruz Creek.
is narrow and nearly flat. The distal end of the calcaneum is Description. The incomplete left maxilla (MCP-4290-PV)
narrower lateromedially than the rest of the body and bears bearing M2 and M3, is from an adult. Both molars show
the cuboid facet. The groove between the articular facets considerable occlusal wear (Figure 3a). Only parts of the

Table 2. Measurements from the tibia of Tapirus. Abbreviations: L: length; MDP: mediolateral diameter of proximal portion; CDP: craniocaudal diameter
of proximal portion; MDB: mediolateral diameter of the body; MDD: mediolateral diameter of distal portion; CDD: craniocaudal diameter of distal por-
tion; MDDA: mediolateral diameter of distal articular surface; LTC: length of tibial crest. Where N > 1, first line is mean ± standard deviation, second
line is observed range, and third line is number of observations (N). Data of T. cristatellus from Winge (1906) and the cave provenance.

L MDP CDP MDB MDD CDD MDDA LTC

T. terrestris 258.44 ± 9.0 75.55 ± 3.4 65.90 ± 5.1 26.05 ± 1.89 47.97 ± 2.2 40.56 ± 2.8 39.26 ± 3.28 96.01 ± 7.04
244 -275 70.25 -84.3 57.1 – 75 23.2 -28.7 44.9 – 53 35.8 – 45 34.3 - 46.8 85 - 105.5
(16) (16) (16) (16) (16) (16) (16) (16)
MN2098-V 268 66.62 54 28.9 52 44.8 37.3 92
T. cristatellus -- -- -- -- 57 -- -- --
Escrivania 1
T. cristatellus -- -- -- -- 55 -- -- --
Escrivania 5
 Tapirus from the Pleistocene of southern Brazil 313

Table 3. Measurements of the calcaneus in Tapirus. Abbreviations: L: and parastyle are absent. MCN-PV-8085 is a fragment of
length; MDB: mediolateral diameter of the body; MDD: mediolateral a left protolophid, probably from a dp3 (Figure 4g-h). The
diameter of distal portion; where N > 1, first line is mean ± standard devia-
tion, second line is observed range, and third line is number of observations mesial cingulum is very well marked. The protoconid and
(N). Data of T. cristatellus from Winge (1906) and the cave provenance. metaconid are higher than the cross crest that unites them,
forming a slightly V-shaped protolophid. MCN-PV-8086
L MDT CDP MW
is also a crown fragment, but a hypolophid, probably of
T. terrestris 100.79 ± 4.8 43.06 ± 1.17 24.41 ± 1.43 35.44 ± 2.79 a right m2, showing little wear and abrasion (Figure 4i-j).
94.1 - 110.6 39 - 48.82 22.5 – 27 33.04 – 38 Its distal cingulum and subcingulum are conspicuous. The
(11) (11) (11) (11)
entoconid is higher than the hypoconid, and the cross crest
DGM79M 101 45.6 22.2 33 is lower, forming a V-shaped hipolophid.
T. cristatellus 113 -- -- -- The scapula (MCN-PV-9700) is well preserved and
Escrivania 1 has the robust form with deep musculature fossae indica-
T. cristatellus 111 -- -- -- tive of a full adult (Figure 5a-b). The specimen is elongate
Escrivania 5
and narrow. The lateral surface is divided into two slightly
asymmetric fossae by a prominent scapular spine. The free
crest of the spine is thickened, directed caudally, and bears
no acromion. There is a prominent and robust tuber in the
alveolar and jugal processes of the maxilla are preserved. middle of the free crest of the spine. The supraspinous fossa
The fragment is broken anterior to M2, and extends poste- is wider than the infraspinous fossa, and slightly triangular.
rior to M3. The posterior portion of the jugal process is a On the surface of the supraspinous fossa there is a muscular
lateroposteriorly directed blade, forming a concavity in the line, very thickened, that divides it into two slightly deep
posteroventral border of the maxilla.
The M2 and M3 (MCP-4290-PV) have a trapezoidal
form, with the protoloph wider than the metaloph. The
protocone of M2 is fractured and exhibits much wear on the
lophs (Figure 3). The protoloph and metaloph are separated
lingually by a transverse groove, but buccally the paracone
and metacone are united due to wear, forming a pronounced
longitudinal crest (ectoloph). The mesial cingulum is worn
completely and the distal cingulum is still sharp. The M2
parastyle is well developed. The M3 protocone is also
fractured and shows less wear than the M2. The M3 meta-
loph and protoloph have wear on the mesial face. The M3
parastyle is smaller than in the M2. The M3 has a greater
subcingulum, and is separated from the paracone by a
conspicuous buccal groove. The mesial cingulum shows
wear and the distal cingulum is much more defined. Both
the M2 and M3 have a lingual cingulum on the metalophs.
The mandible fragment (MCN-PV-8843) preserves
alveoli for the canines and right p2 (Figure 4a). The diastema
is 35 mm long, and the portion bearing alveoli for incisors
is lacking. The symphysis is solidly fused, concave and
very robust (46.5 mm in width), extending posteriorly to a
point anterior to p2. The labial wall of the mental foramen
is located posteroventral to the p2 alveolus.
Specimen MCN-PV-3515 is a left protoloph, prob-
ably a DP4, showing little wear or abrasion (Figure 4b-c).
The protocone is higher than the paracone; the parastyle
is small and separated from the paracone by a transverse
groove. The mesial cingulum is very visible. Specimen
MCN-PV-6968 is a right protoloph, probably a DP3/P4
(Figure 4d-e). It displays only the protocone and part of
the cross crest with little wear. The mesial cingulum is
Figure 3. Tapirus cf. T. terrestris, with the second (M2) and third (M3)
hardly visible. Specimen MCN-PV-8087 is a crown frag- upper molar, a - b: MCP-4290PV, left maxilla in occlusal and lateral views.
ment of an upper molar, showing wear in the protoloph and Abbreviations: hy: hypocone; me: metacone; pa: paracone; past: parastyle.
metaloph, and much abrasion (Figure 4f). The cingulum Scale bar = 30 mm.
314 Holanda et al.

Figure 4. Tapirus cf. T. terrestris, a: MCN- PV-8843, symphyseal portion of mandible in occlusal view; b-c: MCN-PV-3515, left protoloph in mesial and
occlusal views; d-e: - MCN-PV-6968, right protoloph in mesial and occlusal views; f: MCN-PV-8087, fragment of an upper molar in occlusal view; g-h:
MCN-PV-8085, left protolophid in mesial and occlusal views; i-j: MCN-PV-8086, right hypolophid in distal and occlusal views. Abbreviations: entd:
entoconid; hyd: hypoconid; med: metaconid; pa: paracone; past: parastyle; pr: protocone; prd: protoconid; 1: p2 alveolus. Scale bar in a = 30 mm and
b - f = 10 mm.

fossae. The medial surface is irregular, and presents the concave dorsally and convex palmarly, and the lateral facet
slightly rugose facies serrata. The subscapular fossa is for the magnum, narrower, flat lateromedially, and slightly
much deeper and larger. convex dorsopalmarly (Figure 5f). Medially and palmarly
The cranial border is thickener and more rounded than to the trapezoid facet is a small facet, slightly convex to the
the caudal border, and slightly oblique in relation to the trapezium. Lateral to facet for the magnum is a low but thick,
spine. Distal to the cranial border, it forms a deep and high slightly flat facet for metacarpal III. The body has two faces:
scapular notch. The dorsal border is much thickened in the one dorsal, slightly flat and turned dorsomedially; and one
most axial portion, which is most prominent and inclined palmar, more convex and very rough. The distal condyle
medially. The ventral border is the most robust part of the is robust, dorsally convex, and palmarly divided by a high
scapula. The glenoid cavity is shallow; its lateral border crest in two convexities. Lateral to each condyle is a small
forms a larger arc than the medial border, and it extends semicircular notch.
cranially around the contour of the lateral supraglenoid The metatarsal III (MCN-PV-10069) has one larger
tuberosity. The supraglenoid tuberosity projects ventrally, facet for the ectocuneiform in the proximal end. The ecto-
with a medial inclination. Dorsal to supraglenoid tuberosity cuneiform facet is slightly triangular and concave, with the
is the robust and rugose coracoid process. lateral margin higher than medial margin (Figure 5i). Lateral
The scapula measurements (in millimeters) are: maxi- to the ectocuneiform facet are two facets for metatarsal IV;
mum length: 299; craniocaudal diameter: 142; maximum the dorsal facet is slightly triangular and the plantar one is
width of supraspinous fossa: 65.6; craniocaudal diameter of oval, these separated by a deep groove. The plantar facet
neck: 47.2; craniocaudal diameter of articular surface: 78; is on a prominent process, although partly broken, and is
craniocaudal diameter of glenoid cavity: 43; lateromedial directed lateroplantarly. Medial to the ectocuneiform facet
diameter of glenoid cavity: 46.5. there are two smaller facets for metatarsal II, separated by
The metacarpal II (MCN-PV-7071) has a proximal end a shallow groove. The dorsal facet is larger than plantar
slightly narrower than the distal (Figure 5d). There are two facet, and slightly oval.
proximal facets separated by a high ridge; the medial facet Remarks. The specimen from Dom Pedrito (São Luiz cattle
for the trapezoid, thicker than broad, slightly rectangular, ranch) was compared to material from other species of
 Tapirus from the Pleistocene of southern Brazil 315

Figure 5. Tapirus cf. T. terrestris, a - b: MCN- PV-9700, right scapula in medial and lateral views (left to right); d and f: MCN-PV-7071, left metacarpal II
in dorsal and proximal views; h - i: MCN-PV-10069, left metatarsal III in dorsal and proximal views. T. terrestris, c: MCN 2848, right scapula in lateral
view; e and g: MCN 2532, left metacarpal II in dorsal and proximal views. Abbreviations: ef: ectocuneiform facet; fs: facies serrata; mf: magnum facet;
ml: muscular line; sf: suprascapular fossa; sn: scapular notch; ss: spine of the scapula; st: supraglenoid tuberosity; t: tuber; tf: trapezoid facet. Scale bar
in a - e = 30 mm and f - i= 10 mm.

Tapirus wherever possible. The dental characters in MCP- metacarpal II (MCN-PV-7071) and the mediolateral diam-
4290-PV are very similar to those found in T. terrestris. eter of proximal portion of metatarsal III (MCN-PV-10069;
Considering its size, the M2 is smaller than that in T. Table 5). All other measures of metacarpal II and metatarsal
indicus or in T. cristatellus, but is similar in size to that in III are as in T. terrestris, MCN-PV-7071 and -10069 are
T. terrestris and T. mesopotamicus (Table 4). Regarding both smaller than in T. cristatellus (Table 5).
the M3, all compared species overlap, although T. indicus These specimens are here tentatively assigned to
and T. cristatellus specimens are larger than those of Tapirus cf. T. terrestris despite having a somewhat larger
T. terrestris. size, considering that there is otherwise no difference
The Hermenegildo Beach materials are very fragmen- with T. terrestris, T. mesopotamicus shows an overlap in
tary, abraded, and show signs of reworking, but they do not some dental measurements with T. terrestris, although the
differ from T. terrestris. The postcranial specimens (MCN- length total of the skull is 390 mm (Ferrero and Noriega,
PV-7071, MCN-PV-9700, and MCN-PV-10069) are robust, 2007), whereas T. terrestris has a mean length of 351.7 mm
but are within of large end of the size range in T. terrestris, (Simpson, 1945). There is no difference in the dental mor-
except for craniocaudal diameter of the proximal portion of phology or size between T. terrestris and T. mesopotamicus.
316 Holanda et al.
Table 4. Measurements in millimetres from the second (M2) and third (M3) upper molar of Tapirus. Abbreviations: L: length; AW: maximum width of
anterior loph; PW: maximum width of posterior loph; where N > 1, first line is mean ± standard deviation (SD), second line is observed range (OR), and
third line is number of observations (N). Data of T. cristatellus from Winge (1906) and the cave provenance.

M2 M3
L AW PW L AW PW

T. terrestris 22.85 ± 1.42 25.85 ± 1.59 22.44 ± 1.31 22.88 ± 1.57 26.13 ±1.44 21.44 ± 1.27
19.1 -25.36 22.84 - 30.9 19 - 25.7 20 - 26 23.75 - 29.7 20 - 25.5
(48) (48) (48) (26) (26) (26)
MCP-4290PV 23.26 28.14 24.78 23.14 27.24 22.38
T. cristatellus Escrivania 1 26 28.75 - 26 28.75 -
T. cristatellus Escrivania 5 26.5 29 - - - -
T. mesopotamicus 23.74 28.74 25.3 24.3 28.1 22.54
T. indicus 27.85 ± 1.34 30.45 ± 0.77 26.02 ± 0.35 25.7 29.7 23
26 – 29.2 29.4 – 31.2 25.6 – 26.4
(4) (4) (4)

DISCUSSION AND CONCLUSIONS villages and cartographic toponyms, such as “Poço das
Antas” and “Anta Gorda”, indicate the occurrence of this
The material described herein confirms the presence species in the central region of Rio Grande do Sul before
of Tapirus terrestris in the late Pleistocene of Rio Grande do colonization. Ihering (1892) proposed that 32ºS latitude
Sul state. Currently, T. terrestris occurs in Rio Grande do Sul (Figure 1), which he called the “line of Paca”, was the
state, in some remnants of the Atlantic Forest in the Upper southern distribution limit of some species, including T.
Uruguay River, specifically at Parque Estadual do Turvo terrestris, which was the same as the limit of virgin forest
(Kasper et al., 2007), in the northwestern part of the state. at that time.
Historical records are known from the Taquari-Antas River In either case, regarding the southern area of Rio
(Ferri, 1991), but with no localities specified. In addition, Grande do Sul state, there are hitherto neither historical nor
Ihering (1892) and Rambo (2000) report the occurrence of recent records of this species. However, the fossil record
this species on the slopes of Serra Geral, Araucaria Plateau. indicates that during the late Pleistocene in Rio Grande do
Recently, Rosa and Jacobus (2009) recorded T. terrestris Sul, as well as in Argentina and Uruguay (Tonni, 1992;
from archaeological sites dated between ± 2,000 and 8,790 Ubilla, 1996), this species had a more southern distribution,
years B.P. at Maquiné, Santo Antônio da Patrullha, Torres, perhaps reflecting of a better climate and more southerly
Porto Alegre, and Candelária Municipalities, in the central distribution of forest that ceased at the end of Pleistocene
and east regions of the state. However, popular names of time and lead to its local extinction.

Table 5. Measurements from the metatarsal and metacarpal of Tapirus. Abbreviations: L: length; MDP: mediolateral diameter of proximal portion; CDP:
craniocaudal diameter of proximal portion; MDB: mediolateral diameter of the body; MDD: mediolateral diameter of distal portion; where N > 1, first
line is mean ± standard deviation (SD), second line is observed range (OR), and third line is number of observations (N). Data of T. cristatellus from
Winge (1906) and the cave provenance.

L MDP CDP MDB MDD

Metacarpal II
T. terrestris 103.57 ± 3.38 19.68 ± 2.27 17.66 ± 0.98 18.57 ± 1.95 25 ± 2.67
99 – 110 16 - 22.74 16.3 - 19.52 15.3 - 21.2 21 - 28.4
(11) (11) (11) (11) (11)
MCN-PV-7071 102.8 18 24.6 18 22.2
T. cristatellus Escrivania 5 -- 27 -- -- --

Metatarsal III
T. terrestris -- 28.98 ± 1.33 26.76 ± 1.45 -- --
27.2 – 31 24.3 – 29
(13) (13)
MCN-PV-10069 -- 32.4 27.6 -- --
T. cristatellus Escrivania 1 -- 35 -- -- --
T. cristatellus Escrivania 5 -- 37 -- -- --
 Tapirus from the Pleistocene of southern Brazil 317

ACKNOWLEDGEMENTS Cartelle, C.,1999, Pleistocene Mammals of the Cerrado and Caatinga of

Brazil, in Eisenberg, J. F, Redford, K. H. (eds.) Mammals of the
Neotropicals: The Central Neotropicals, Vol. 3: University of
The authors are greatly indebted to R. Cassab (DNPM), Chicago Press, Chicago, 27-46.
M.C. Malabarba (PUCRS) and D. D. Rego (MNRJ) for ac- Cattoi, N., 1951, El status de Tapirus dupuyi (C. Amegh.): Comunicaciones
cess to the specimens studied; to C.C. Cardoso (MHN/ del Instituto Nacional de Investigacion de las Ciencias Naturales e
UFMG), C. Cartelle (PUCMinas), D. Flores (MACN), Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”,
2 (8), 103-112.
M. Martins (MCN), M. Merino (MLP), J.A. de Oliveira Cattoi, N., 1957, Uma especie extinguida de Tapirus Brisson (T. rioplatensis
(MNRJ), and M. de Vivo (MZUSP), for access to compara- nov. sp.), Ameghiniana, 1 (3), 15 – 21.
tive material under their care; to L. Rota who collected many Colbert, M.W., 2005, The facial skeleton of the early Oligocene Colodon
of the studied fossils; to J.C. Cisneros for language revision; (Perissodactyla, Tapiroidea): Palaeontologia Eletronica, 8 (1),
12A-27, 600kb.
to L. F. Lopes (UFRGS) for photography of Figure 4; and Da Rosa, A. A. S., 2003, Preliminary correlation of fluvial deposits at the
to Conselho Nacional de Desenvolvimento Científico e extreme west of Rio Grande do Sul State, Southern Brazil, in 3th
Tecnológico (CNPq) for financial support (CNPq/Universal Latinamerican Congress of Sedimentology, Belém: Abstracts,
474485/2008-0, CNPq/Prosul 490299/2008-3 and E.C. p. 243-245.
Desmarest, A.G., 1819, Nouveau dictionnaire d’histoire naturelle. Volume
Holanda’s fellowship). 32: Chez Deterville, Paris.
DeSantis, L.R.G., MacFadden, B., 2007, Identifying forested environments
in Deep Time using fossil tapirs: evidence from evolutionary
APPENDIX morphology and stable isotopes: Courier Forschunsinstitut
Senckenberg, 258, 147-157.
Eisenmann, V., Guérin, C., 1992, Tapirus priscus Kaup from the Upper
Material used for anatomical comparisons: T. ter- Miocene of Western Europe: palaeontology, biostratigraphy, and
restris: MLP 01, 754, 755, 1349, 1402, 1681, 4IV0013; palaeoecology: Paleontologia I Evolució, 24-25, 113-122.
MACN 7.6, 31211, 33276, 50559; MCN 1315, 2532, 2750, Ferrero, B.S., Noriega, J. I., 2007, A new upper Pleistocene tapir from
Argentina: remarks on the phylogenetics and diversification of
2848; MN18, 600, 1605, 53701, 57062, 57067, 64437, neotropical Tapiridae: Journal of Vertebrate Paleontology, 27,
64572, 64652, 70698, 71599; MZUSP 106, 3232, 3266, 504-511.
3268, 3269, 3727, 3728, 3758, 5701, 6139, 6140, 6575, Ferrero, B.S., Brandoni, J. I., Noriega, J. I., Carlini, A. A., 2007, Mamíferos
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29085, 20035, 20037, 10715, 20034, 31983; UNIR 17, 20, “Bernardino Rivadavia”, 9, 109-117.
68, 83. T. indicus: MACN 129, 2553, 4347, 29926, 30351. Ferri, G.,1991, História do Rio Taquari – Antas: Encantado, Grafen Gráfica
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