Quaternary mammals from central Brazil (Serra da Bodoquena, Mato Grosso do Sul) and comments on paleobiogeography and paleoenvironments

Abstract

Studies regarding Quaternary mammals from the Serra da Bodoquena (South-western Brazil, state of Mato Grosso do Sul) are scarce. In the region, the Fadas Cave has been an important paleontological site. Remnants of Pleistocene fauna were collected along a river channel inside the cave. It is the first record of Arctotherium, Scelidotheriinae and Nothrotheriinae in the Serra da Bodoquena. Smilodon populator, Eremotherium laurillardi, Gomphotheriidae and Glyptodontinae, already known for the region, were also identified. These new occurrences allow us to redefine the paleozoogeographical distribution of some of the aforementioned taxa. Herein we propose the expansion of the range of the Brazilian Intertropical Region, which should include Mato Grosso do Sul and São Paulo. The state of Rio Grande do Sul keeps a greater paleozoogeographical affinity to others temperate zones from Argentina and Uruguay. Taxa associated to open environments indicate the existence of large areas of savannas during the late Pleistocene in the Serra da Bodoquena.

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31
Rev. bras. paleontol. 20(1):31-44, Janeiro/Abril 2017
© 2017 by the Sociedade Brasileira de Paleontologia
doi: 10.4072/rbp.2017.1.03
ABSTRACT – Studies regarding Quaternary mammals from the Serra da Bodoquena (South-western Brazil, state of
Mato Grosso do Sul) are scarce. In the region, the Fadas Cave has been an important paleontological site. Remnants of
Pleistocene fauna were collected along a river channel inside the cave. It is the rst record of Arctotherium, Scelidotheriinae
and Nothrotheriinae in the Serra da Bodoquena. Smilodon populator, Eremotherium laurillardi, Gomphotheriidae and
Glyptodontinae, already known for the region, were also identied. These new occurrences allow us to redene the
paleozoogeographical distribution of some of the aforementioned taxa. Herein we propose the expansion of the range of the
Brazilian Intertropical Region, which should include Mato Grosso do Sul and São Paulo. The state of Rio Grande do Sul
keeps a greater paleozoogeographical anity to others temperate zones from Argentina and Uruguay. Taxa associated to
open environments indicate the existence of large areas of savannas during the late Pleistocene in the Serra da Bodoquena.
Key words: Brazilian Intertropical Region, Pampean Region, paleozoogeographical zones, Fadas Cave, Quaternary mammals,
Serra da Bodoquena.
RESUMO – Os estudos sobre mamíferos quaternários da Serra da Bodoquena (Mato Grosso do Sul) são escassos. Na região,
a Gruta das Fadas tem sido um importante sítio paleontológico. Os restos da fauna do Pleistoceno foram coletados ao longo
de um canal do rio dentro da caverna. É o primeiro registro de Arctotherium, Scelidotheriinae e Nothrotheriinae na Serra da
Bodoquena. Smilodon populator, Eremotherium laurillardi, Gomphotheriidae e Glyptodontinae, já conhecidos pela região,
também foram identicados. Estas novas ocorrências nos permitem redenir a distribuição paleozoogeográca de alguns dos
táxons acima mencionados. Aqui propomos a ampliação da faixa da Região Intertropical Brasileira, que deve incluir Mato
Grosso do Sul e São Paulo. O Estado do Rio Grande do Sul mantém uma maior anidade paleozoogeográca com outras
zonas temperadas da Argentina e do Uruguai. Táxons associados a ambientes abertos indicam a existência de grandes áreas
de savanas durante o Pleistoceno nal na Serra da Bodoquena.
Palavras-chave: Região Intertropical Brasileira, Região Pampeana, zonas paleozoogeográcas, Gruta das Fadas, mamíferos
quaternários, Serra da Bodoquena.
QUATERNARY MAMMALS FROM CENTRAL BRAZIL (SERRA DA
BODOQUENA, MATO GROSSO DO SUL) AND COMMENTS ON
PALEOBIOGEOGRAPHY AND PALEOENVIRONMENTS
ALESSANDRO MARQUES DE OLIVEIRA
Laboratório de Sistemática e Diversidade, Universidade Federal do ABC, Avenida dos Estados, 5001,
Bairro Bangu, 09210-580, Santo André, SP, Brasil. biolessandro@gmail.com
BRUNO BECKER-KERBER
Departamento de Ecologia e Biologia Evolutiva, Universidade Federal de São Carlos, Washington Luiz 325 km,
13565-905, São Carlos, SP, Brasil. bruno.becker92@gmail.com
LÍVIA MEDEIROS CORDEIRO, RODRIGO BORGHEZAN
Centro de Ciências Biológicas e da Saúde, Universidade Federal do Mato Grosso do Sul, 79070900,
Campo Grande, MS, Brasil. liv.biosubt@gmail.com, rodrigo_borghezan@hotmail.com
LEONARDO SANTOS AVILLA
Laboratório de Mastozoologia, Universidade Federal do Estado do Rio de Janeiro, Av. Pasteur, 458, Urca,
22290-255, Rio de Janeiro, RJ, Brasil. leonardo.avilla@gmail.com
MÍRIAN LIZA ALVES FORANCELLI PACHECO
Departamento de Biologia, Universidade Federal de São Carlos, Campus Sorocaba, Rodovia João Leme dos Santos,
km 104, Parque Reserva Fazenda Imperial, 18052-780, Sorocaba, SP, Brasil. forancelli.ufscar@gmail.com
CHARLES MORPHY D. SANTOS
Laboratório de Sistemática e Diversidade, Universidade Federal do ABC, Avenida dos Estados, 5001,
Bairro Bangu, 09210-580, Santo André, SP, Brasil. charlesmorphy@gmail.com

REVISTA BRASILEIRA DE PALEONTOLOGIA, 20(1), 2017
32
INTRODUCTION
Despite being constantly debated, there are several
aspects about South American Pleistocene megafauna (giant
mammal fossils with estimated biomass above 1000 kg);
especially regarding paleoecology and paleobiogeography,
that still need investigation. Frequently, paleoecological
and paleoenvironmental approaches take into account
the feeding habits based on isotopical and morphological
analyses (Bargo & Vizcaíno, 2008; Dantas et al., 2013a). In
Brazil, most of these studies have focused on fossils from
North and Northeast regions (Dantas et al., 2013a; França
et al., 2014). Concerning biogeographical aspects, there
are no attempts to correlate or to establish biogeographical
affinities among taxa from South-western Brazil (e.g. Mato
Grosso and Mato Grosso do Sul) and those from others
South American paleozoogeographical zones. Nevertheless,
understanding the paleoecological and paleobiogeographical
aspects of Quaternary giant mammals can elucidate several
questions about their environment and extinction.
In the State of Mato Grosso do Sul (MS), paleontological
studies regarding Quaternary mammals are rare (Scheffler
et al., 2010). The first mention to Pleistocene mammals in
the region was made by Oliveira (1915), based on material
collected in Lagoa Seca, 12 km south of the city of Corumbá.
In the last 40 years, additional reports on Pleistocene mammal
fossils from Mato Grosso do Sul were made by Cunha (1975,
1981), Auler & Boller (1992), Salles et al. (2006), Boggiani et
al. (2009), Perini et al. (2009) and Oliveira (2013). However,
few of the aforementioned studies deal with biogeographical,
paleoecological or paleoenvironmental implications, being
restricted to the register of new fossil occurrences.
The present study deals with material from the Serra da
Bodoquena, a sustained limestone plateau of the Corumbá
Group (Ediacaran), located in the western portion of Mato
Grosso do Sul (Figure 1). Caves and sinkholes in the karst
plateau of Bodoquena are Quaternary deposits rich in
remnants of Pleistocene megafauna and other mammals
(Salles et al., 2006; Scheffler et al., 2010; Oliveira, 2013).
Most of the caves occur in dolostones of residual hills in the
depression of the Miranda River (Kholer et al., 1998).
The Fadas Cave (Figure 1) (20°34’04”S, 56°46’28”W,
altitude 336 m) is located in the Campina Settlement, about
10 km from the city of Bodoquena (MS). The cavity has a
small entrance, 1.5 m wide and 0.6 m high, which opens into
a small chamber with breakdown blocks in the substrate. A
cluster of speleothems abruptly contacts the first chamber
to a second one, larger, ca. 60 m long and 10 m high at
its highest point. Large sandy-clay deposits constitute
the substrate. On the left side of the second chamber, two
passages lead to a lower conduit through which flows an
underground river.
In 2006, the first exploration of the Fadas Cave
revealed new Pleistocene mammals, such as ground
sloths, gomphotheres, glyptodonts, felines and a single
Ursidae specimen (Oliveira, 2013), which were found at
the bottom of the river conduit. These findings suggest the
paleogeographical and paleoecological importance of Serra
da Bodoquena regarding the South American Pleistocene
megafauna and others large mammals. The present paper aims
to describe these new records of Pleistocene mammal fossils
from Fadas Cave, and to comment on their biogeographical
significance, as well as to discuss the paleoenvironment of
the Serra da Bodoquena during the late Quaternary.
MATERIAL AND METHODS
The fossils were collected on the surface, in gravel banks,
and on the riverbed, using snorkeling in deeper sections (1.5 m
depth) of the river flowing in Fadas Cave. The specimens were
deposited at the Museu de Arqueologia of the Universidade
Federal de Mato Grosso do Sul (campus Campo Grande)
(labeled PBQ1-23, PBQ1-20, PBQ1-01, PBQ1-22, PBQ1-06,
PBQ1-04) and at the Laboratório de Sistemática e Diversidade
of the Universidade Federal do ABC (campus Santo André)
(labeled PBQ2-1, PBQ2-2, PBQ2-3, PBQ2-4).
In order to establish a clearer taxonomical correlation
among Mato Grosso do Sul and other South American
paleozoogeographical zones, we made a cluster analysis.
Such procedure took into account the occurrence of extinct
Quaternary mammals above 50 kg in the Mato Grosso do
Sul, Brazilian Intertropical region, São Paulo, Rio Grande do
Sul, Uruguay, Mesopotamian region, North central Argentina,
South of Bolivia, Paraguay and Pampean region. In Table 1,
we present a matrix of presence (1) and absence (0) of taxa
in South American paleozoogeographical zones, identified to
species level, based on Prado & Alberdi (1999), Bond et al.
(2001), Alberdi & Prado (2004), Carlini et al. (2004), Noriega
et al. (2004), Zurita et al. (2004, 2007, 2011), Soibelzon et al.
(2005, 2012), Salles et al. (2006), Pitana & Ribeiro (2007),
Scherer et al. (2007), Scheffler et al. (2010), Pitana (2011),
Miño-Boilini (2012), Silva et al. (2012), Zurita et al. (2012),
Lopes (2013), Díaz et al. (2014), Lobo et al. (2015), Varela
& Fariña (2016), McAfee (2016), Missagia et al. (2016), and
Figure 1. Localization of Fadas cave, Serra da Bodoquena, State of
Mato Grosso do Sul, Central Brazil.

33
OLIVEIRA ET AL. – QUATERNARY MAMMALS FROM CENTRAL BRAZIL
Table 1. Matrix of presence/absence of taxa in South American paleozoogeopgraphical zones. The list include only extinct Quaternary mammals
above 50 kg weigh. Abbreviations: SP, São Paulo; RS, Rio Grande do Sul; BIR, Brazilian Intertropical Region; PR, Pampean region; MS, Mato
Grosso do Sul; NCA/SB/P, North central Argentina, South of Bolivia and Paraguay; MR, Mesopotamian Region.
Taxa SP RS BIR PR MS NCA/SB/P MR
Arctotherium wingei 0110010
Arctotherium bonariense 0001000
Arctotherium vetustum 0001001
Arctotherium angustidens 0001010
Arctotherium tarijensis 0001010
Smilodon populator 1111111
Toxodon gracilis 0001001
Toxodon platensis 1111111
Cuvieronius hyodon 0000010
Notiomastodon platensis 1111111
Eutatus seguini 0001000
Holmesina paulacoutoi 011101 1
Holmesina majus 0110000
Pampatherium typum 0001010
Pampatherium humboldt 011010 0
Propraopus sulcatus 0110000
Propraopus magnus 1010000
Propraopus grandis 010101 1
Sclerocalyptus ornatus 0001010
Doedicurus clavicaudatus 0101000
Panochytus tuberculatus 010001 1
Neuryurus rudis 0100000
Glyptodon munizi 0001000
Glyptodon elongatus 0001010
Glyptodon reticulatus 110101 1
Glyptodon clavipes 0101000
Glyptodon perforatus 0000001
Glyptotherium cylindricum 0010000
Oncopus gracilis 1010000
Aytherium aureum 1010000
Eremotheirum laurillardi 111010 0
Mylodon darwini 0101001
Mylodonopsi ibseni 001010 0
Lestodon ortizianus 0000001
Lestodon armatus 110101 1
Glossotherium lettsomi 001110 0
Glossotherium robustum 0101010
Scelidodon tarijensis 0001010
Scelidotherium bravardi 0001000
Scelidotherium leptocephalum 010101 1
Valgipes buckland 0010000
Catonyx cuvieri 1111000
Megatherium americanum 010101 1
Nothrotherium maquinense 1010000
Equus (amerhipus) neogenus 0111111
Hippidium devillei 0011010
Hippidium principale 011110 0
Catagonus stenocephalus 0111010
Morenelaphus brachyceros 000001 1
Neolicaphrium recens 0101000
Macrauchenia patachonica 011101 1
Palaeolama major 001010 0
Hemiauchenia paradoxa 010101 1
Lama gracilis 0101001
Lama guanicoe 0111001

REVISTA BRASILEIRA DE PALEONTOLOGIA, 20(1), 2017
34
including the specimens herein identified. The cluster analysis
was performed on binary matrix employing the clustering
algorithm UPGMA (Average Linkage Clustering; Sokal &
Michener, 1958) and the Euclidean similarity index using
the software Past 3.14.
Taxonomical identification followed Cartelle (1992), De
Iuliis (1996), Cartelle & De Iullis (1995, 2006), Pereira et al.
(2012), for Eremotherium laurillardi, Brandoni & McDonald
(2015) and De Iullis et al.(2011) for Nothrotheriinae, Miño-
Boilini et al. (2014) for Scelidotherium, Rinderknecht
(2000), Zurita et al. (2009), Luna & Krapovickas (2011)
for Glyptodon, Paula Couto (1979) and Mothé et al. (2012)
for Gomphotheriidae, and Cartelle (1998) for Arctotherium.
The identification of Smilodon populator was made after
anatomical comparison with material housed at the Natural
History Museum of London (cast – specimen no labeled)
and at the National Museum of Natural History from Paris
(specimen TAR-52).
SYSTEMATIC PALEONTOLOGY
Order XENARTHRA Cope, 1889
Family MEGATHERIIDAE (Gray, 1821) Owen, 1842
Eremotherium Lund, 1842
Eremotherium laurillardi (Lund, 1842) Cartelle &
Bohórquez, 1982
(Figures 2A–C)
Material. PBQ1-23, right astragalus; PBQ2-1, left calcaneum;
PBQ2-2, fragment of right mandibular branch.
Description. The right mandibular branch (Figure 2C) is
fragmented at the beginning of the second molariform alveoli
(MF2). The specimen presents the last molariform (MF4)
(measuring 125 mm in length), the third alveoli molariform
(MF3), and part of the ascending ramus containing the
mandibular foramen. There is a shallow groove between
the ascending ramus and the last molariform, typical of
the genus Eremotherium (Cartelle, 1992; Cartelle & De
Iullis, 1995; Pereira et al., 2012). As described by Pereira
et al. (2012) for Eremotherium laurillardi, the MF4 of the
specimen studied here shows the mesial ridge wider and
mesodistally compressed when compared to the distal crest.
The pulp cavity occupies about two-thirds of the total size of
the tooth, a diagnostic character for E. laurillardi according
to Cartelle & De Iuliis (2006). The astragalus (Figure 2A) is
complete, with few fractures at the edges. Its proximodistal
and anteroposterior extensions, measuring 200 and 230 mm,
respectively, are within the range of variation presented among
Figure 2. A–C, Eremotherium laurillardi. A, right astragalus; B, right calcaneum; C, right mandibular branch. D, Gomphoteriidae, part of an
incisive tooth. E–F, Glyptodon, E, dorsal portion of the carapace, dorsal view; F, detail of osteoderm highlighted. Abbreviations: 1, medial
trochlea; 2, lateral trochlea; 3, articular surface to the navicular; 4, articular surface to the bula; 5, tuber calcis; 6, sustentacular facet; 7, sulcus
talis; 8, ectal facet; 9, distal crest; 10, medial crest; 11, mandibular foramen; 12, alveoli of third molariform; 13, lateral osteoderm; 14, pilose
foramen; 15, central gure. The white arrow indicates downgraded area of central gure. Scale bars: A–C = 50 mm; D–F = 40 mm.
A
1
2
4
9
13
3
5
6
7
8
10 11
12
14
15
B
D F E
C

35
OLIVEIRA ET AL. – QUATERNARY MAMMALS FROM CENTRAL BRAZIL
adult megatheriids (De Iuliis, 1996). The medial trochlea
is a semi-spherical odontoid process. Lateral trochlea is
anteroposteriorly flattened, discoid shaped. Both throcleas
form an angle of 90° to each other, indicating a total medium
lateral foot rotation. The articular surface to the navicular has
a circular concave shape. The calcaneum (Figure 2B) has large
sustentacular and ectal facets, each of which measuring 120
x 80 mm, slightly concave and separated by a 20 mm wide
sulcus talis. The calcaneal tuber is fragmented, with few signs
of abrasion on its edges.
Remarks. Usually, three species of Eremotherium were
recognized as valid species: E. laurillardi (from Brazil),
E. rusconii (from Central America and Northeastern South
America) and E. mirabile (from the United States of
America) (Porta, 1961; Edmund 1965; Bocquentin, 1979;
Polaco-Ramos, 1981). Later, Cartelle & De Iuliis (1995)
synonymized these three species as E. laurillardi. These
authors observed a wide variation in morphology and size
among the specimens of E. laurillardi, which may bias the
identification of Megatheriidae. Recently, Faure et al. (2014)
sustained the hypothesis of E. rusconii as a valid species
based on the description of two mandibles, which these
authors interpreted one as belonging to a young individual
of E. rusconii and the other to an adult of E. laurillardi.
However, after studying 944 molariform teeth, Cartelle
et al. (2015) showed that E. rusconii is more likely to be
a junior synonym and that the morphological differences
represent intraspecific variations of E. laurillardi. It is
possible that E. laurillardi was distributed throughout Brazil
during the Pleistocene (Cartelle & Bohorquez, 1982; De
Iuliis & Cartelle, 1999). E. laurillardi was first recorded to
Mato Grosso do Sul by Ayub et al. (1996). Carbon isotope
analyses have shown a generalist diet for E. laurillardi in the
Brazilian Intertropical Region, suggesting that the species
was associated with both open areas and forests (Dantas et
al., 2013a; França et al., 2014).
Family MYLODONTIDAE Gill, 1872
Subfamily SCELIDOTHERIINAE Ameghino, 1904
(Figures 3A–D)
Material. PBQ1-01, left humerus.
Description. Humerus (Figures 3A–D) 480 mm length, with
straight and semi-cylindrical shaft, the structure is shorter and
more robust compared to other mylodontids, such as Catonyx
cuvieri and Valgipes buckland. The deltoid and pectoral ridges
are not evident in cranial view and there is no intermediate
ridge. At the proximal epiphysis, the humeral head is less
prominent and limited by the greater and lesser tubercles; the
lateromedial diameter is approximately 160 mm. The distal
epiphysis is lateromedially expanded, about 225 mm in width,
and anteroposteriorly flattened. The entepicondylar foramen
is present, and its entepicondylar bar is horizontally oriented.
Remarks. Among the mylodontids recorded in Brazil,
Glossotherium, Lestodon and Catonyx cuvieri do not have
an entepicondylar foramen. Valgipes buckland presents the
entepicondylar bar obliquely arranged. The species Scelidodon
tarijensis, unknown from Brazil, has a more developed deltoid
crest in comparison to the specimen here described. The
morphology of our specimen is suggestive, but not conclusive:
identifying the material to species level demands some cranial
bone elements absent here. Nonetheless, positioning the
specimen in the subfamily Scelidotheriinae seems reliable.
Miño-Boilini et al. (2014) recognized the existence of two
species of this genus, S. leptocephalum and S. bravardi.
S. leptocephalum is restricted to the Bonaerense-Lujanense of
the Pampean Region and to the Lujanense of the Argentinian
provinces of Córdoba, Santa Fe, Corrientes, Salta and
Formosa. S. bravardi is known for Ensenadan Age deposits,
near the cities of Buenos Aires, La Plata and Mar del Plata, and
for the Pleistocene of Villa Larca, in the Argentinian province
of San Luis. The specimen here described is the first record
of Scelidotheriinae to Mato Grosso do Sul state. Species of
this subfamily were probably adapted to both dry/cold and
hot/humid environments (Cione & Tonni, 1995; Carlini et
al., 2004, 2008; Ferrero & Noriega, 2009; Tonni, 2009), with
great ecological tolerance (Miño-Boilini, 2012).
Family NOTHROTHERIIDAE Ameghino, 1920
Subfamily NOTHROTHERIINAE Ameghino, 1920
(Figures 3E–H)
Material. PBQ2-3, left femur; PBQ2-4, right femur.
Description. Both femurs (Figures 3E–H) are complete and
have 430 mm in length. They are wide and anteroposteriorly
flattened, with the distal portion lightly broader than the
proximal one. At the proximal epiphysis, the femoral head
is prominent but the neck is reduced and the fovea capitis
femoris is fully inserted in the caput. The greater trochanter
is prominent, with a discrete proximal projection. The lesser
trochanter is not prominent, being more medially projected in
PBQ2-4 than in PBQ2-3, forming a triangular protuberance.
A deep trochanteric fossa is present. Distally, the third
trochanter is connected to the lateral epicondyle forming a
continuous surface. At the distal epiphysis, the medial and
lateral condyles are asymmetric (80 and 60 mm respectively),
and separated by the intercondylar notch. The patellar trochlea
has a concave surface and it is not connected to the medial and
lateral condyles. The lateral epicondyle is anteriorly projected.
Remarks. Brandoni & McDonald (2015) described a
similar specimen of Nothrotheriinae from the Pleistocene
of Argentina (MACN Pv 14148 and MACN Pv 14149),
placing it as a Nothrotheriops unidentified species or as
Nothropus carcarensis (which was described based only
on a partial mandible). The specimens from Serra da
Bodoquena (Figure 3) have a set of characters that strongly
suggests their identification as a species of the genus
Nothrotheriops (e.g. shape and position of the greater
trochanter, development of the lesser trochanter, connection
between the third trochanter and the lateral epicondyle, and
the size). However, the lack of specific diagnostic characters
led us to identify the specimens PBQ2-3 and PBQ2-4 as an
undetermined Nothrotheriinae, based on the same features
listed by Brandoni & McDonald (2015). Nothrotheres

REVISTA BRASILEIRA DE PALEONTOLOGIA, 20(1), 2017
36
are known from the first half of the Miocene until the
Pleistocene (De Iullis et al., 2011). They have been placed
in the family Megalonychidae (McDonald & Muizon, 2002)
or among the Megatheriidae (Paula Couto, 1979; Patterson
et al., 1992). However, nothrotheres are considerably
distinct from Megatheriidae and Megalonychidae. Hence,
we place the Nothrotheriinae here identified in the family
Nothrotheriidae (cf. Gaudin & De Iullis, 1999; Muizon et
al., 2004).This is the first record of a nothrothere for Central
Brazil – Brazilian nothrotheres were previously known
only for the states of São Paulo, Minas Gerais, Bahia and
Ceará (Lund, 1839; Cartelle & Fonseca, 1983; Cartelle,
2000; Ghilardi et al., 2011). In South America, Quaternary
nothrotheres are included in two genera: Nothrotherium
(found mainly in Brazil) and Nothropus (Argentina).The
genus Nothrotheriops is unknown for South American
Quaternary deposits, being recorded only in North America
(Brandoni & McDonald, 2015). In general, species of
Nothrotheriinae were inhabitants of open environments,
browsing on a variety of xerophytic vegetation (Thompson
et al., 1980; Ghilardi et al., 2011). Although there are several
evidences that support an opportunistic browsing or mixed
feeding strategy for Nothrotheriops shastensis (Poinar et al.,
1998; Hofreiter et al., 2000), Green (2009) found a more
abrasive diet based on a dental microwear analysis, which
could be explained by the consume of grasses.
Family GLYPTODONTIDAE Gray, 1869
Subfamily GLYPTODONTINAE Burmeister, 1879
(Figures 2E–F)
Material. PBQ1-22, fragments of carapace.
Description. 33 osteoderms (Figures 2E–F) were recovered
and analyzed. Three are lateral cone-shaped osteoderms. The
other dermal plates have irregular, pentagonal or hexagonal
shape, with a circular central figure, surrounded by poorly
Figure 3. A–D, Scelidotheriinae, left humerus in anterior (A, C) and posterior (B, D) views. E–H, Nothrotheriinae, left femur in posterior (E, G)
and anterior (F) views; right femur in anterior view (H). Abbreviations: 1, humeral head; 2, lesser tubercle; 3, greater tubercle; 4, pectoral crest;
5, entepicondylar foramen; 6, medial epicondyle; 7, trochlea; 8, deltoid crest; 9, femoral head; 10, lesser trochanter; 11, major trochanter; 12,
third trochanter; 13, medial epicondyle; 14, lateral epicondyle; 15, patella surface; 16, trochanteric fossa; 17, medial condyle; 18, intercondylar
fossa; 19, lateral condyle. Arrow indicates entepicondylar bar. Scale bars = 40 mm.
A
12
3
4
5
13 13
66
14 14
715
16
17
18
19
412 12
11
10 10
19
C
B
D
E
G
F
H
8

37
OLIVEIRA ET AL. – QUATERNARY MAMMALS FROM CENTRAL BRAZIL
defined peripheral figures that are much smaller than the
central one. Such peripheral figures vary in number from
7 to 8 (the number of peripheral figures varies from 6 to 8
in Glyptodon and from 7 to 13 in Glyptotherium, and the
overlap often leads to misidentification) and are separated by
a piliferous foramen, which also varies in number from 7 to
8. The sulci separating the peripheral figures, as well as the
central sulcus, are very shallow, almost imperceptible. The
central figure appears circular and higher in relation to the
peripheral, but due to the degree of abrasion of the material, it
is difficult to observe such a feature. In the central area of the
central figure, there is a slight downgrade, typical of juvenile
specimens of glyptodonts, which includes Glyptodon (Luna
& Krapovickas, 2011; Gillette et al., 2016). The diameter of
each osteoderm varies from 30 to 40 mm.
Remarks. At the Pliocene/Pleistocene boundary,
glyptodonts were widely distributed in South America,
from Argentina to Venezuela. Glyptodontidae is represented
by approximately 67 genera (Fernicola, 2008), with
Glyptodon being the most documented in Brazil. Previous
authors suggested that the Brazilian Intertropical Region
was inhabited by Glyptotherium, while Glyptodon would
have been restricted to Southern Brazil (Oliveira et al.,
2010). In the state of the Mato Grosso do Sul, the genus
Glyptodon was previously recorded by Salles et al. (2006)
and Ayub et al. (1996) at the city of Jardim. In general,
the glyptodonts are considered grazers adapted to open
environments (Bargo et al., 2006).
Order PROBOSCIDEA Illiger, 1811
Family GOMPHOTHERIIDAE Hay, 1922
(Figure 2D)
Material. PBQ1-06, incisor fragment of juvenile individual.
Description. Incisor fragmented (Figure 2D) in the middle
portion, 200 mm length, with longitudinal bands of enamel.
Remarks. Gomphotheres are well documented in South
America (Argentina, Brazil, Chile, Colombia, Ecuador,
Paraguay, Peru, Uruguay and Venezuela) from Ensenadense
to Lujanense (Prado et al., 2011), with a single species found
in Brazil, Notiomastodon platensis (Mothé et al., 2012). Due
to the absence of diagnostic features on the studied specimen,
we have decided to not assign a specified taxonomical
classification beyond Gomphotheriidae. Based on stable
isotopes analysis, Dantas et al. (2013a) attributed a generalist
diet to Brazilian gomphotheres (N. platensis). This would
indicate that the species had a high ecological flexibility,
being able to live in open areas as well as in forests. However,
França et al. (2014) stated that N. platensis could not modify
its diet in a short period of time, which would expose them to
local extinction or force them to migrate if the environment
abruptly changed.
Order CARNIVORA Bowdich, 1821
Family FELIDAE Gray, 1821
Smilodon Lund, 1842
Smilodon populator Lund, 1842
(Figures 4A–D)
Material. PBQ1-21, right humerus with part of the epiphyses
fragmented.
Description. Right humerus (Figures 4A–D), 400 mm length.
At the proximal epiphysis, portions of the greater tubercle,
lesser tubercle and the humeral head are fragmented. The shaft
is sub-triangular in cross section, with distal portion thinner
than the proximal one. The deltoid crest is poorly developed
compared to extant big cats (Panthera tigris and Panthera
leo). Distal epiphysis has medial-lateral extent of 125 mm,
and is fragmented in the medial epicondyle. Entepicondylar
foramen is present and entepicondylar bar is obliquely
oriented. In anterior view, the radial fossa is shallow, and
the capitulum and the mediodistally directed trochlea are
highly developed. In posterior view, the olecranon fossa is
deep and the lateral supracondylar ridge is well developed.
The aforementioned features are diagnostic for Smilodon
populator, the single saber toothed-tiger recorded to Brazil.
Remarks. Smilodon populator had occurred in the Pampean
Region (PR) in Argentina, Southern Chile (Prieto et al., 2010),
Peru (Shockey et al., 2009), Uruguay, Ecuador, Brazil, Bolivia
and Venezuela (Rincón, 2006). In Brazil, fossils of this feline
have been found in the states of Minas Gerais (Piló & Neves,
2003), Paraíba (Correa et al., 2012), Ceará (Gomide et al.,
1987; Bergqvist et al., 1997), Goiás (Gomide et al., 1987),
Bahia (Lessa et al., 1998), Rio Grande do Sul (Lessa et al.,
1998), Rio Grande do Norte (Lessa et al., 1998; Porpino et
al., 2004), Piauí (Guérin et al., 1996; Faure & Guérin, 2014),
Pernambuco (Silva et al., 2003), Sergipe (Dantas, 2012), São
Paulo (Castro & Langer, 2008; Ghilardi et al., 2011) and Mato
Grosso do Sul (Salles et al., 2006; Perini et al., 2009). The
classification of the genus Smilodon is controversial. Some
authors claim there was a single species in the Americas (Paula
Couto, 1979; Berta, 1987; Lessa et al., 1998; Cartelle & De
Iuliis, 2006), whereas others recognize two species (Kurtén &
Werdelin, 1990; Christiansen, 2012). Bocherens et al. (2016)
suggested that S.populator predominantly hunted prey in
open environments, behaving as a social predator. However,
ecomorphological studies have indicated that these great
felines were also adapted to forests (Meloro et al., 2013).
Family URSIDAE Gray, 1825
Arctotherium Bravard, 1857
(Figures 4E–H)
Material. PBQ1-04, distal epiphysis of left humerus with
part of a shaft.
Description. Distal epiphysis of left humerus (Figures
4E–H), 200 mm length. The humeral shaft is cylindrical.
The distal epiphysis is wide with 120 mm medial-lateral
extent. The medial condyle is greater than the lateral one,
mediodistally directed. Entepicondylar foramen is arranged
on the medial face; entepicondylar bar is obliquely arranged
and the trochlear fossa is quite broad. The abrasion marks on

REVISTA BRASILEIRA DE PALEONTOLOGIA, 20(1), 2017
38
the edges are minimal. All these descriptions are compatible
with the specimen IGC-10 attributed to Arctotherium by
Cartelle (1998).
Remarks. The Ursidae subfamily Tremarctinae was
distributed exclusively in the Americas. It probably originated
in Central Western North America, reaching South America
during the so called“Great American Biotic Interchange”,
from the Later Miocene to the Holocene (Soibelzon et al.,
2005). In a systematic review of the Tremarctinae, Soibelzon
(2004) recognized a single genus (Arctotherium) and five
species for this group in South America (A. augustinenses,
A. wingei, A. vetustum, A. bonariense and A. tarijense).
Arctotherium was recorded in Brazil in the states of Minas
Gerais, Ceará, Bahia, Rio Grande do Norte and possibly Rio
Grande do Sul (Trajano & Ferrarezi, 1994; Cartelle, 1998;
Lessa et al., 1998; Soibelzon et al., 2005; Porpino et al.,
2004; Pereira et al., 2012). With the occurrences registered
in the present paper a new geographical distribution for
Arctotherium in South America is configured, now including
the Brazilian territory of Mato Grosso do Sul. A primarily
herbivore diet has been attributed to A. wingei (Figueirido
& Soibelzon, 2009). According to Soibelzon & Schubert
(2011), large guilds of big carnivores would have increased in
diversity after the “Great American Biotic Interchange”, and
smaller species of Arctotherium could have changed their diets
to herbivory or omnivory in response to competitive pressures
with large carnivores. However, our specimen is bigger than
other Arctotherium specimens previously reported to Brazil.
Some authors claim that Arctotherium was associated with
arid and semi-arid paleoenvironments (Trajano & Ferrarezzi,
1994) and even to cold environments resultant from glacial
events during the Late Pleistocene (Lopez et al., 2008). If
this statement is correct, the distribution of Arctotherium
may indicate that the Serra da Bodoquena was more arid in
that period.
CLUSTER ANALYSIS
The greatest Euclidean Distance Index (EDI) (6,0827625)
obtained is between Brazilian Intertropical Region (BIR)
and Pampean Region (PR). The lesser EDI (3,6055513) is
observed between Mato Grosso do Sul (MS) and São Paulo
(SP), and between Rio Grande do Sul (RS) and Uruguay (Uru)
(Table 2). The state of the MS, SP and BIR form a group,
Figure 4. A–D, Smilodon populator, right humerus in posterior (A, C) and anterior (B, D) views. E–H, Arctotherium sp., distal epiphysis of left
humerus in anterior (F, H) and posterior (E, G) views. Abbreviations: 1, lateral epicondyle; 2, capitulum; 3, trochlea; 4, medial epicondyle; 5,
entepicondylar foramen; 6, entepicondylar bar; 7, olecranon fossa; 8, shaft; 9, lateral supracondylar crest; 10, deltoid crest; 11, humeral head;
12, greater tubercle. Scale bars = 40 mm.
A
E F G H
BC D
7
7
1
1
2
2
3
3
3
4 4 5
5
6
6
8
8
9
10
9

39
OLIVEIRA ET AL. – QUATERNARY MAMMALS FROM CENTRAL BRAZIL
Figure 5. Cluster analysis indicating greater taxonomical a󰀩nity
among Mato Grosso do Sul (MS), São Paulo (SP) and the Brazilian
Intertropical Region (BIR). The state of Rio Grande do Sul (RS) is
more similar to Uruguay (Uru). The group composed by RS and Uru,
Mesopotamian Region (MR), north central Argentina, South of Bolivia
and Paraguay (NCA/SB/P) share most of their taxa from Pampean
Region (PR).
whereas RS, Uruguay, North central Argentina, South of
Bolivia and Paraguay (NCA/SB/P) and Mesopotamian Region
(MR) share greater taxonomical affinity to PR, forming a
second group (Figure 5).
PALEOBIOGEOGRAPHY AND
PALEOENVIRONMENT
Cartelle (1999) defined the Brazilian Intertropical
region (BIR) as a paleozoogeographical region especially
characterized by the presence of large mammals in savannahs
ca. 20,000–10,000 years BP. BIR included the states of Goiás,
Minas Gerais, Rio de Janeiro, Espírito Santo, Bahia, Sergipe,
Alagoas, Pernambuco, Rio Grande do Norte, Paraiba, Ceará
and Piauí. The Pampean region (PR) is located in Argentina
between 30°–39°S and 57°–66°W, with current climate
ranging from temperate to humid subtropical (Prado &
Alberdi, 2010). According to Tonni & Figalgo (1978), during
Pleistocene the climatic conditions alternated between arid
or semi-arid and humid. The predominance of dry and cold
climate in the late Pleistocene is suggested by the presence of
fossil pollen especially from the Brassicaceae (Quattrocchio et
al., 2008), and of fossil of extant mammals from Patagonian
and Central sub-regions, associated to arid or semi-arid
and cold weathers (Microcavia australis and Pediolagus
salinicola) (Tonni, 1981, 1985; Tonni et al., 1992; Prado &
Alberdi, 2010).
Based on fossils from the Upper Ribeira speleological
deposits, Ghilardi et al. (2011) considered the states of São
Paulo and Southern Brazil as the transition area between
the BIR and PR. According to them, this transition area
encompassed a combination of megafauna species recurrent
in both BIR and PR, with greater taxonomical affinity to BIR.
Carlini et al. (2004) considered four paleozoogeographical
zones in Southern portion of South America (Figure 6): (i)
North central Argentina, South of Bolivia and Paraguay
(NCA/SB/P); (ii) Mesopotamian region (MR); (iii) Rio
Grande do Sul and Uruguay (RS/Uru); (iv) Pampean region.
However, these authors noted high taxonomical similarity
between NCA/SB/P and PR; whereas the MR and RS/Uru
would share taxa from both PR and BIR. The taxonomical
affinities between Paraguay and PR have been noticed by
Carlini & Tonni (2000), Souberlich & Uchoa (2013) and Diaz
et al. (2014) as well.
The cluster analysis (Figure 5) performed herein shows
that Mato Grosso do Sul has greater taxonomical affinity to
São Paulo, and both share the most of taxa from BIR. This
result corroborates Ghilardi et al. (2011) who proposed a
greater taxonomical similarity between the state of São Paulo
and BIR, relative to PR. Therefore it is suggested herein an
expansion of BIR, which should include Mato Grosso do Sul
(Figure 6). Additionally our results confirm partially Carlini et
al. (2004), who hypothesized greater taxonomical affinity: (i)
between Rio Grande do Sul and Uruguay (composing a single
area); (ii) between the Mesopotamian region and that area
constituted by Rio Grande do Sul and Uruguay. However, the
EDI is lesser between NCA/BS/P and Uruguay; and between
NCA/SB/P and MR. Therefore, the hypothesis of greater
taxonomical affinity between NCA/SB/P and PR is not clear.
Table 2. Euclidian Distance Index among paleozoogeographical zones in South America.
SP RS BIR PR MS NCA/SB/P MR Uru
SP 0 5,099019 4,242640 5,916079 3,605551 5,099019 4,690415 4,582575
RS 5,099019 0 4,898979 4,582575 5 4,690415 4,242640 3,605551
BIR 4,242640 4,898979 0 6,082762 3,872983 5,656854 5,656854 5,567764
PR 5,916079 4,582575 6,082762 0 5,830951 4,358898 4,795831 4,690415
MS 3,605551 5 3,872983 5,830951 0 5,196152 4,795831 4,690415
NCA/SB/P 5,099019 4,690415 5,656854 4,358898 5,196152 0 4,242640 4,123105
MR 4,690415 4,242640 5,656854 4,795831 4,795831 4,242640 0 3,872983
Uru 4,582575 3,605551 5,567764 4,690415 4,690415 4,123105 3,872983 0

REVISTA BRASILEIRA DE PALEONTOLOGIA, 20(1), 2017
40
As Prado et al. (1987) claimed, at the late Pleistocene, the
PR probably had much lower winter temperatures than today’s
average, which led to the dispersion of some species towards
the North. In this context, it is likely that species from PR
(e.g. Scelidotherium, Megatherium, Lestodon armatus and
Glyptodon) dispersed to warmer areas, reaching Paraguay,
South of Bolivia, North Argentina, Southern Brazil, São Paulo
and Mato Grosso do Sul.
As mentioned by Ghilardi et al. (2011), the presence of
taxa from temperate region in the state of São Paulo can
indicate that this region and Southern Brazil composed a
transitional zone between temperate and intertropical regions.
In this context, some authors have mentioned, doubtfully, the
presence of Megatherium and Glyptodon in Mato Grosso
do Sul (Scheffler, 2006; Salles et al., 2006). If this assertion
is confirmed, Mato Grosso do Sul could be included in the
transitional zone.
The occurrence of taxa associated with different
paleoenvironmental contexts led us to hypothesize that Serra
da Bodoquena would be marked by large areas of grasslands
and forest fragments or spots, and lower temperatures than
the current ones (between 20° and 22° Celsius in average)
during the late Pleistocene. This assumption can be compared
to other paleoclimate data from Central Brazil, which indicates
drier environmental conditions from 22,000–11,500 years
BP, followed by an increase in humidity in the last 7,000
years (Salgado-Labouriau et al., 1997; Barberi et al., 2000).
Drier conditions during the late Pleistocene have also been
inferred for regions adjacent to the Serra da Bodoquena, like
the Pantanal, specifically in the alluvial fan of the Taquari
River (Soares et al., 2003).
CONCLUSIONS
The present paper is the first attempt to correlate the
Pleistocene fauna from Serra da Bodoquena to other South
American paleozoogeographical regions. Our data strongly
suggest that species of the genus Arctotherium, a third species
of Scelidotheriinae unidentified and a second species of the
Nothrotheriinae was distributed throughout the Western
portion of the State of Mato Grosso do Sul during the
Pleistocene.
The result of cluster analysis reinforces the hypothesis
that the State of Mato Grosso do Sul took part of Brazilian
Intertropical region. Likely the State of São Paulo served
as a transitional zone contacting Northern, Northeastern
and Southern regions of South America. However, the
same it is not clear for Mato Grosso do Sul, which the
presence of taxa from Pampean region need to be confirmed.
The state of Rio Grande do Sul keeps greater similarity
to North central Argentina, South of Bolivia, Paraguay,
Mesopotamian region, Pampean region and Uruguay than
to Brazilian Intertropical region. Additionally, a greater
paleozoogeographical affinity between Rio Grande do Sul
and Uruguay is corroborated.
Paleoenvironmental inferences from the identified taxa
indicate drier scenarios with larger areas of open vegetation
in the Pleistocene/Holocene transition when compared to
the current characterization of Serra da Bodoquena. Future
studies, including the analysis of stable isotopes and absolute
dating, should be made in order to offer a more complete
picture of the paleoenvironmental aspects of this region.
ACKNOWLEDGEMENTS
We are grateful to Instituto Chico Mendes de Conservação
da Biodiversidade (ICMBio), to M. Domingues, to G.R.
Martins (Museu de Arqueologia, Universidade Federal
de Mato Grosso do Sul). We also thank Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (CAPES)
and Conselho Nacional de Desenvolvimento Científico e
Tecnológico (CNPq) for financial support. Finally we thank
the reviewers for criticisms and suggestions in an early version
of this manuscript.
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