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The Brazilian Intertropical Fauna from 60
to About 10 ka B.P.: Taxonomy, Dating,
Diet, and Paleoenvironments
Mário AndréTrinidade Dantas and Mario Alberto Cozzuol
Abstract This chapter reviews information about the extinct fauna that lived in the
Brazilian Intertropical Region (BIR) between 64 and 10 ka B.P. Data from the
available literature regarding dating (
14
C, ESR, U-series) and paleodiet recon-
struction (δ
13
C) for some of taxa of the BIR are herein presented. Furthermore,
paleoenvironmental reconstructions of two climatic moments are presented, one at
64 ka, and another between 27 and 10 ka B.P.
Keywords Brazilian Intertropical Region Pleistocene megafauna Datings
Feeding ecology
1 Introduction
The Brazilian Intertropical Region (BIR; Fig. 1) has been proposed and defined by
Cartelle (1999) as a zoogeographical domain, based on the occurrence of endemic
species from the Brazilian states of Goiás (GO), Minas Gerais (MG), Rio de Janeiro
(RJ), Espírito Santo (ES), Bahia (BA), Sergipe (SE), Alagoas (AL), Pernambuco
(PE), Rio Grande do Norte (RN), Paraíba (PB), Ceará(CE), and Piauí(PI).
Mammal fossils from this region are most commonly found within “tanks”(tem-
porary ponds), naturally formed by the accumulation of rain water (e.g., Araújo
et al. 2013), or inside caves (e.g., Hubbe and Auler 2012). Fossils of medium (10–
100 kg of biomass), large (100–1000 kg), and giant (more than 1000 kg) sized
mammals, have been found in both areas, whereas small-sized mammals (less than
M.A.T. Dantas (&)
Instituto Multidisciplinar em Saúde, Universidade Federal da Bahia—Campus Anísio
Teixeira, Rua Rio de Contas, 58, Candeias, Vitória da Conquista, BA 45029-094, Brazil
e-mail: matdantas@yahoo.com.br
M.A. Cozzuol
Laboratório de Paleozoologia, Departamento de Biologia Geral, Universidade Federal
de Minas Gerais, Av. Antônio Carlos, 6627, Belo Horizonte, MG 31270-010, Brazil
e-mail: mario.cozzuol@gmail.com
©Springer International Publishing Switzerland 2016
G.M. Gasparini et al. (eds.), Marine Isotope Stage 3 in Southern
South America, 60 ka B.P.–30 ka B.P., Springer Earth System Sciences,
DOI 10.1007/978-3-319-40000-6_12
207
10 kg) have mainly been found in caves. These differences can be explained by the
type of fossilization taking place in each of these deposition environments.
Cartelle (1999) listed a variety of taxa occurring in the BIR, presenting details
about findings from caves in the Minas Gerais and Bahia states. However, despite
the well-known taxonomic knowledge of this area, dating and feeding ecology
information about the taxa that lived in the BIR is scarce.
Thus, this chapter reviews the available literature in order to report: (i) an update
and refinement of the information on mammal taxa occurring within BIR; (ii) a
review of dating and feeding ecology information about these taxa; and (iii) com-
mentaries, whenever possible, about the paleoenvironments in which they lived.
2 Materials and Methods
The data analyzed in this review were obtained from: (i) 08 published feeding
paleoecology studies, which used isotopic carbon analysis (δ
13
C) performed on
hydroxyapatite and collagen in enamel, dentine or bone (Table 2); and (ii) 13
published dating studies, performed with the Electron Spin Resonance, Carbon 14
and U-series techniques (Table 2).
Fig. 1 Map showing the Brazilian Intertropical Region (BIR; represented by the dotted line;
sensu Cartelle 1999)
208 M.A.T. Dantas and M.A. Cozzuol
Table 1 Pleistocene megafauna of the Brazilian Intertropical Region (BIR)
Taxa BA SE AL PE PB RN CE PI GO MG RJ ES
PILOSA
Megatheriidae
Eremotherium laurillardi
(Lund, 1842)
xxxxxxxxxx xx
Mylodontidae
Mylodontinae
indeterminado
x
Glossotherium sp. x x x
Glossotherium lettsomi
(Owen, 1840)
x
Catonyx cuvieri (Lund,
1839)
xxx x xx x
Valgipes bucklandi
(Lund, 1839)
xxxx
Ocnotherium giganteum
(Lund, 1839)
xxx
Mylodonopsis ibseni
Cartelle, 1991
xx x
Megalonychidae x
Ahytherium aureum
Cartelle, De Iuliis and
Pujos, 2008
x
Australonyx aquae De
Iuliis, Pujos and Cartelle,
2009
x
Nothrotheriidae
Nothrotherium
maquinense Lydekker,
1889
xxxx
CINGULATA
Glyptodontidae
Glyptotherium sp. xx xxxxxxx
Panochthus greslebini
Castellanos, 1941
xxxxxx
Parapanocthus
jaguaribensis (Moreira,
1965)
xxxx
CINGULATA
Glyptodontidae
Hoplophorus euphractus
Lund, 1839
xxxxx
(continued)
The Brazilian Intertropical Fauna from 60 to About 10 ka B.P. …209
Table 1 (continued)
Taxa BA SE AL PE PB RN CE PI GO MG RJ ES
Dasypodidae
Pampatherium sp. x
Pampatherium humboldti
Ameghino, 1875
xx xxx
Homelsina paulacoutoi
(Guerra and Mahecha,
1984)
xxxxx
Pachyarmatherium
brasiliense Porpino,
Berqvist and Fernicola,
2009
xx x x ? x ?
CARNÍVORA
Felidae x
Smilodon populator
Lund, 1842
xx xxxxxxx
Ursidae
Arctotherium wingei
Ameghino, 1902
xxxx
Canidae
Protocyon troglodytes
(Lund, 1838)
xxxxx
PROBOSCIDEA
Gomphotheriidae
Notiomastodon platensis
(Ameghino, 1888)
xxxxxxxxxx xx
NOTOUNGULATA
Toxodontidae x
Toxodon platensis Owen,
1840
xxxx xxxx x x
Piauhytherium capivarae
Guérin and Faure, 2013
?x x
PERISSODACTYLA
Equidae
Equus (Amerhippus)
neogaeus Lund, 1840
xxxxxxxxxx
Hippidion principale
(Lund, 1846)
xxxxxxx
ARTIODACTYLA
Camelidae
Palaeolama major Liais,
1872
xx xxxxxxx
(continued)
210 M.A.T. Dantas and M.A. Cozzuol
The current taxonomic arrangements proposed for the South American
Gomphotheriidae are herein followed. Thus, the paleoecological data presented by
Sánchez et al. (2004) for Stegomastodon waringi (Holand 1920) will be attributed
to Notiomastodon platensis postulated by Mothéet al. (2012).
In the analyses of δ
13
C data from hydroxyapatite and collagen in bone or dentine
(e.g. MacFadden et al. 1999; Drefahl 2010; Dantas et al. 2013a; França et al. 2014),
samples were chemically pretreated to eliminate the potential effects of diagenesis
(secondary carbonate contamination), using the protocol described elsewhere.
The interpretation of the diet was based on the fact that most existing plants,
ranging from trees and woody shrubs to grasses found on prairies and steppes at high
altitudes or latitudes, use the Calvin–Benson (C
3
) photosynthetic cycle. These plants
present average values of δ
13
Cof−27 ‰. By contrast, the few terrestrial plants that
use the Hatch-Slack (C
4
) photosynthetic route are primarily tropical and subtropical
grasses (Ehleringer et al. 1991; Cerling 1992). These species are typically found in
open areas in warm regions subject to hydrological stress, and are able to tolerate low
concentrations of CO
2
. In general, C
4
plants have higher δ
13
C values, averaging
−13 ‰(MacFadden et al. 1999; MacFadden 2005). Those plants that photosyn-
thesize using Crassulacean Acid Metabolism (CAM), such as the succulents, present
intermediate δ
13
C values (MacFadden et al. 1999; MacFadden 2005).
Studies of modern medium- to large-sized herbivorous mammals recorded an
enrichment in δ
13
C values between 12 and 14 ‰(13 ‰on average) in comparison
with the values recorded for the ingested vegetation (Sánchez et al. 2004). Given
this, δ
13
C values lower than −10 ‰are typical of animals with a diet consisting
exclusively of C
3
plants, while δ
13
C values higher than −1‰are consistent with a
diet based on C
4
plants. Intermediate δ
13
C values (between −10 and −1‰) indicate
a mixed diet of C
3
and C
4
plants (MacFadden et al. 1999; MacFadden 2005).
Table 1 (continued)
Taxa BA SE AL PE PB RN CE PI GO MG RJ ES
Cervidae x
Mazama gouazoubira
(Fischer, 1814)
xxxxxxx
Ozotoceros bezoarticus
(Linneus, 1758)
xxxxx
LIPTOTERNA
Macraucheniidae
Xenorhinotherium
bahiense Cartelle and
Lessa, 1988
xxxxx xxx
Labels: confirmed presence: ‘x’; unconfirmed presence: ‘?’. Location: BA Bahia; SE Sergipe; AL
Alagoas; PE Pernambuco; PB Paraíba; RN Rio Grande do Norte; CE Ceará;PI Piauí;GO Goiás;
MG Minas Gerais; RJ Rio de Janeiro; ES Espírito Santo
The Brazilian Intertropical Fauna from 60 to About 10 ka B.P. …211
3 Results
3.1 Taxonomy and Feeding Paleoecology
Paleoecological knowledge about the Pleistocene megafauna that lived in the BIR is
mainly based on interpretations made by a few authors (e.g., Cartelle 1999). Data from
ecomorphological studies have not been published yet (only as gray literature), and
the few published data from carbon isotope analysis are restricted to herbivore taxa
(e.g., Viana et al. 2011; Dantas et al. 2013a; França et al. 2014). The taxa registered in
the BIR, as well as interpretations about their feeding ecology, are presented below.
These interpretations, whenever possible, are based on analyses of fossils from the
BIR, or otherwise, from other regions. The extinct medium- and small-sized herbi-
vores were excluded from this analysis (e.g., primates, rodents, and marsupials), given
the morphological similarities with the living species within these groups, what is
helpful in describing their ecology. The same reasoning was applied to the other living
mammal taxa, which have a generally well-described ecology (Reis et al. 2011).
3.2 Order Carnivora
Fossils of a variety of living and extinct carnivore species have been discovered in
“tanks”and caves from the BIR. Extinct taxa include representatives of the three
families, Protocyon troglodytes (Lund 1838) (Canidae), Smilodon populator (Lund
1842) (Felidae), and Arctotherium wingei (Ameghino 1902) (Ursidae), each
occupying specific niches (Table 1).
P. troglodytes was a hypercarnivore species (i.e., diet composed of 70 % meat)
with a body mass estimated from 16 to 37 kg. It is believed to have inhabited open
fields, where it hunted in groups, feeding on medium-sized herbivores, such as
species of the families Cervidae, Tayassuidae, Camelidae, and Equidae, and
small-sized terrestrial sloths (Cartelle and Langguth 1999; Prevosti et al. 2005).
According to evidence collected by Prevosti and Vizcaíno (2006), the
saber-toothed cat S. populator was probably a solitary species, with a body mass
ranging from 220 to 360 kg. It has likely been specialized to predate on large-sized
prey, such as the giant sloths and gomphothere species.
A. wingei was a medium-sized bear adapted to open areas and dry climates, with
a body mass estimated in 43–107 kg (Soibelzon and Tarantini 2009). It was
probably an omnivorous species, tending to herbivorous habits, which fed on plant
soft tissues (Trajano and Ferrarezzi 1994; Soibelzon and Schubert 2011).
3.3 Order Pilosa
Sloths (extinct and living) and anteaters (Gaudin 2004) belong to this order. Pilosa
(excepting the toothless anteaters) are characterized by a high degree of dental
212 M.A.T. Dantas and M.A. Cozzuol
reduction and teeth that lack the enamel layer, classified as hypsodont and pris-
matic. The teeth of these forms of reduced and simplified dentition are generally
known as molariforms, even when some of them are projected toward the front, in
the position of the canines or the incisors (Paula Couto 1979). Forms of sexual
dimorphism, in which the females are smaller than males, have been proposed for
some Pleistocene taxa belonging to the Megatheriidae (Cartelle 1992) and
Mylodontidae (Abuhid 1991 apud Cartelle 1999; McDonald 2006) families.
Phylogenetic analyses support the existence of nine species and four families of
Pilosa during the Pleistocene inhabiting the BIR: Eremotherium laurillardi
(Megatheriidae, Megatheriinae); Mylodonopsis ibseni,Glossotherium lettsomi,
Ocnotherium giganteum (Mylodontidae, Mylodontinae); Valgipes bucklandi,
Catonyx cuvieri (Mylodontidae, Scelidotheriinae); Nothrotherium maquinense
(Nothrotheriidae, Nothrotheriinae); Ahytherium aureum,Australonyx aquae
(Megalonychidae) (Cartelle 1991; Cartelle and De Iuliis 1995; Cartelle 1999;
Gaudin 2004; Cartelle et al. 2008,2009; De Iuliis et al. 2009).
Although there are only a few studies about the feeding paleoecology of species
inhabiting the BIR, the analysis of carbon isotope ratios for some of these animals
yielded important results.
Among the terrestrial sloths living in this region, E. laurillardi had the largest
body size, with some of the specimens reaching six meters in length. Its body mass
was estimated in four tons, similar to that proposed for Megatherium americanum
Cuvier, 1796 (Fariña et al. 1998). The masticatory apparatus of Eremotherium and
Megatherium is morphologically similar, indicating resembling biomechanics.
These sloths had a great capacity for oral processing of food, which suggests low
digestive efficiency. They had likely a powerful bite, allowing the processing of soft
and fibrous types of food, and suggesting a diet composed of leaves from trees and
shrubs, along with fruits (Bargo et al. 2006a; Guimarães et al. 2008).
Carbon isotope analyses for E. laurillardi found in the Rio Grande do Norte,
Alagoas, Sergipe, and Bahia states indicate that, in the BIR, these animals had a diet
based on grass and herbaceous plants (i.e., C
4
plants; δ
13
C = 0.3 and 0.91;
Table 2), or a mixed diet, feeding also on leaves and fruits, of trees and shrubs (C
3
plants, δ
13
C=−9.20 to −2.06; Table 2). These species inhabited open areas or
forest edges.
Fariña(1996) proposed that E. laurillardi might have also fed on meat, which
could classify them as opportunistic omnivores, however evidence to support this
hypothesis is still to be found.
According to Bargo et al. (2006a), the Mylodontidae, as opposed to the
Megatheriidae, did not have a high capacity for oral processing of food; neither had
they a strong bite power. Bargo et al. (2006a,b) attributed grazer habits to the
Mylodontinae giant sloths Glossotherium robustum and Lestodon armatus, which
would have diets based on grass and herbaceous plants. The estimated body mass of
these species is about 1200–2500 kg, respectively (Fariña et al. 1998).
Two species with morphologies of the masticatory apparatus similar to the ones
above mentioned were found in the BIR. G. lettsomi has been proposed to be a
synonym of G. robustum (Esteban 1996 apud Fernicola et al. 2009), and O.
The Brazilian Intertropical Fauna from 60 to About 10 ka B.P. …213
giganteum, which according to Cartelle (1999) is morphologically similar to
Lestodon. Therefore, it is reasonable to expect that, given their latitudinal locations,
these species would also feed on C
4
gramineae species.
Cartelle (1991) described M. ibseni and stated in the diagnosis that it is mor-
phologically close to Mylodon darwini. Additionally, Bargo et al. (2006a,b)
assigned a generalist diet to M. darwini and this diet is attributed tentatively to M.
ibseni in this paper.
In the phylogenetic analysis of Scelidotheriinae proposed by Gaudin (2004) and
Cartelle et al. (2009), C. cuvieri and V. bucklandi are considered morphologically
close to Scelidotherium leptocephalum. This may suggest that C. cuvieri and V.
bucklandi have shown high hypsodonty levels due to the large amount of dust
particles of the soil they consumed together with the food, and also to the observed
adaptations to burrowing habits. Body masses were estimated in about 500 kg
(Fariña et al. 1998; Bargo et al. 2000; Vizcaíno et al. 2001). These taxa were likely
browsers, feeding mainly on plant buds, fruits, or roots (Bargo 2001 apud Bargo
et al. 2006b). Data from Pereira et al. (2013), showing values of δ
13
C=−10.17 ‰
for V. bucklandi, further support these assumptions (Table 2).
The only representative of the family Nothrotheriidae in the BIR is N. maqui-
nense. Cartelle (1999) proposed this species should occupy a similar niche to the
current arboreal sloths, feeding from the leaves found on tree tops.
Among the Megalonychidae, there are two Pleistocene species currently rec-
ognized in the BIR, A. aureum and A. aquae. According to De Iuliis et al. (2009), A.
aureum would be more closely related to the North American taxa (e.g.,
Megalonyx), whereas A. aquae would be to species from the Antilles (e.g.,
Megalocnus). These species are believed to have been browsers, such as Megalonyx
(McDonald et al. 2001), since they share similar cranial morphologies, and have the
same dental formula M4/m3, with triangular molariforms and “incisor”teeth.
3.4 Order Cingulata
This order is composed of xenarthrans with a carapace of bony plates, which covers
their back, sides, head top, and tail. They featured a higher number of teeth than
sloths, a minimum of 28 hypsodont, rootless molariforms (Paula Couto 1979).
Fossils from the BIR comprise living and extinct taxa, the latter highlighted by the
giant armadillos and glyptodont.
Glyptodonts differ from giant species of armadillos (and the remaining
Dasypodidae armadillos) by the absence of movement in the plates of their cara-
pace, showing vertebrae fusions, and by the presence of trilobed teeth (Hoffstetter
1958; Paula Couto 1979). Two giant species of armadillos are known for the BIR,
Pampatherium humboldti Ameghino, 1875 and Homelsina paulacoutoi (Guerra
and Mahecha 1984), both considered as grazers of harsh vegetation (Vizcaíno
2009). Grazer glyptodonts (Vizcaíno 2009) with body masses varying from 1000 to
2000 kg (Fariña et al. 1998) were represented by the following taxa: Panochthus
214 M.A.T. Dantas and M.A. Cozzuol
Table 2 Review of the available dating data from 60 to *10 ka (Carbon 14—
14
C; Electron Spin
Resonance—ESR; Thorium/Uranium—Th/U) and paleodiet reconstructions (Carbon isotopes
—δ
13
C; b—bioapatite; c—collagen) for some taxa from the Brazilian Intertropical Region—BIR
Taxa Sample Material Latitude
13
C (CO
3
)
‰
VPDB
Dating
technique
Reference
V. bucklandi UGAMS
11763
Bone 05° 49′−10.17
(b)
–Pereira et al.
(2013)
T. platensis UGAMS
09442
Enamel 05° 52′−1.32
(b)
10.730 ±30
(
14
C)
Dantas et al.
(2013a)
E. laurillardi UGAMS
09436
Dentine 05° 57′−5.22
(b)
–Dantas et al.
(2013a)
N. platensis UGAMS
09440
Enamel 05° 57′0.44
(b)
16.150 ±40
(
14
C)
Dantas et al.
(2013a)
E. laurillardi UGAMS
09435
Dentine 06° 15′0.50
(b)
15.490 ±40
(
14
C)
Dantas et al.
(2013a)
N. platensis Unnumbered Enamel 07° 11′–39 ±7 (ESR) Kinoshita
et al. (2005)
N. platensis Unnumbered Enamel 07° 11′–30 ±5 (ESR) Kinoshita
et al. (2005)
X. bahiense Unnumbered Enamel 07° 11′–39 ±7 (ESR) Kinoshita
et al. (2005)
N. platensis Unnumbered Enamel 07° 45′–22 ±3 (ESR) Kinoshita
et al. (2013)
T. platensis Unnumbered Enamel 07° 45′–26 ±4 (ESR) Kinoshita
et al. (2013)
N. platensis Unnumbered Enamel 08° 14′–60 ±9 (ESR) Kinoshita
et al. (2008)
N. platensis Unnumbered Enamel 08° 14′–63 ±8 (ESR) Kinoshita
et al. (2008)
E. laurillardi SM-1 Dentine 09° 22′0.30
(b)
–Viana et al.
(2011)
N. platensis SM-3 Enamel 09° 22′0.00
(b)
–Viana et al.
(2011)
N. platensis Unnumbered Enamel 09° 22′–39.8 ±1 (ESR) Oliveira et al.
(2010b)
N. platensis Unnumbered Enamel 09° 22′–10 ±0.5 (ESR) Oliveira et al.
(2010b
T. platensis SM-5 Enamel 09° 22′−4.10
(b)
–Viana et al.
(2011)
P. major Unnumbered Enamel 09° 46′–38 (ESR) Dantas et al.
(2011)
T. platensis Unnumbered Enamel 09° 46′–50 (ESR) Dantas et al.
(2011)
E. laurillardi UGAMS
09431
Dentine 09° 55′−6.65
(b)
–Dantas et al.
(2013a)
E. laurillardi UGAMS
09432
Dentine 09° 55′−3.85
(b)
22.440 ±50
(
14
C)
Dantas et al.
(2013a)
E. laurillardi UGAMS
09433
Dentine 09° 55′−2.45
(b)
–Dantas et al.
(2013a)
(continued)
The Brazilian Intertropical Fauna from 60 to About 10 ka B.P. …215
Table 2 (continued)
Taxa Sample Material Latitude
13
C (CO
3
)
‰
VPDB
Dating
technique
Reference
E. laurillardi UGAMS
13539
Dentine 09° 55′−7.70
(b)
10.990 ±30
(
14
C)
França et al.
(2014)
E. laurillardi UGAMS
13540
Dentine 09° 55′−3.30
(b)
11.010 ±30
(
14
C)
França et al.
(2014)
E. laurillardi UGAMS
13541
Dentine 09° 55′−6.00
(b)
9.720 ±30
(
14
C)
França et al.
(2014)
E. laurillardi UGAMS
13542
Dentine 09° 55′−3.30
(b)
9.730 ±30
(
14
C)
França et al.
(2014)
E. laurillardi UGAMS
13543
Dentine 09° 55′−4.70
(b)
11.580 ±30
(
14
C)
França et al.
(2014)
N. platensis UGAMS
09437
Dentine 09° 55′0.76
(b)
–Dantas et al.
(2013a)
N. platensis UGAMS
09438
Enamel 09° 55′−1.04
(b)
13.980 ±40
(
14
C)
Dantas et al.
(2013a)
N. platensis UGAMS
13535
Enamel 09° 55′−0.40
(b)
13.380 ±35
(
14
C)
França et al.
(2014)
N. platensis UGAMS
13536
Enamel 09° 55′−0.20
(b)
16.370 ±40
(
14
C)
França et al.
(2014)
N. platensis UGAMS
13537
Enamel 09° 55′−1.10
(b)
10.440 ±30
(
14
C)
França et al.
(2014)
N. platensis UGAMS
13538
Enamel 09° 55′1.30
(b)
13.760 ±35
(
14
C)
França et al.
(2014)
N. platensis Unnumbered Enamel 09° 55′–42 (ESR) Dantas et al.
(2011)
N. platensis Unnumbered Enamel 09° 55′–27 ±3 (ESR) Dantas et al.
(2013b)
T. platensis UGAMS
09446
Enamel 09° 55′−3.68
(b)
10.050 ±30
(14C)
Dantas et al.
(2013a)
E. laurillardi UGAMS
09434
Dentine 10° 00′−3.25
(b)
–Dantas et al.
(2013a)
N. platensis Unnumbered Enamel 10° 00′–50 (ESR) Dantas et al.
(2011)
T. platensis Unnumbered Enamel 10° 00′–50 (ESR) Dantas et al.
(2011)
N. platensis UGAMS
09439
Enamel 10° 05′−1.86
(b)
17.910 ±50
(14C)
Dantas et al.
(2013a)
N. platensis UGAMS
09441
Enamel 10° 17′−0.49
(b)
–Dantas et al.
(2013a)
T. platensis UGAMS
09443
Enamel 10° 17′−1.08
(b)
–Dantas et al.
(2013a)
T. platensis UGAMS
09444
Dentine 10° 17′−1.00 –Dantas et al.
(2013a)
N.
maquinense
Unnumbered Calcite 10° 18′–15.425 ±491
(Th/U)
Auler et al.
(2006)
N.
maquinense
Unnumbered Calcite 10° 18′–15.031 ±375
(Th/U)
Auler et al.
(2006)
(continued)
216 M.A.T. Dantas and M.A. Cozzuol
Table 2 (continued)
Taxa Sample Material Latitude
13
C (CO
3
)
‰
VPDB
Dating
technique
Reference
T. platensis U-96-150 Enamel 10° 21′−5.50
(b)
–MacFadden
(2005)
E. laurillardi UGAMS
06136
Bone 10° 42′−18.20
(c)
15.770 ±40
(14C)
Drefahl
(2010)
E. (A.)
neogaeus
Unnumbered Enamel 10° 55′1.10
(b)
–MacFadden
et al. (1999)
E. (A.)
neogaeus
Unnumbered Enamel 10° 55′1.70
(b)
–MacFadden
et al. (1999)
N. platensis Unnumbered Enamel 10° 55′−8.20
(b)
–Sánchez et al.
(2004)
N. platensis Unnumbered Enamel 10° 55′−5.00
(b)
–Sánchez et al.
(2004)
T. platensis U-96-148 Enamel 10° 55′−12.60
(b)
–MacFadden
(2005)
T. platensis U-96-149 Enamel 10° 55′−7.70
(b)
–MacFadden
(2005)
E. laurillardi Unnumbered Calcite 10° 58′–15.000 ±500
(Th/U)
Auler et al.
(2006)
E. laurillardi Unnumbered Calcite 10° 58′–16.100 ±3.900
(Th/U)
Auler et al.
(2006)
E. laurillardi Unnumbered Calcite 10° 58′–15.800 ±2.000
(Th/U)
Auler et al.
(2006)
N. platensis Unnumbered Enamel 11° 32′–50 ±10 (ESR) Ribeiro et al.
(2013)
T. platensis Unnumbered Enamel 11° 32′–43 ±8 (ESR) Ribeiro et al.
(2013)
E. (A.)
neogaeus
CM 11032 Enamel 12° 00′−0.60
(b)
–MacFadden
et al. (1999)
T. platensis UGAMS
09445
Enamel 14° 46′−13.24
(b)
10.970 ±30
(14C)
Dantas et al.
(2013a)
N. platensis Unnumbered Enamel 19° 35′–64 ±5 (ESR) dos Avilla
et al. (2013)
C. cuvieri Unnumbered Bone 19° 37′–14.030 ±50
(14C)
Neves and
Piló(2003)
C. cuvieri Unnumbered Bone 19° 37′–13.920 ±50
(14C)
Neves and
Piló(2003)
C. cuvieri Unnumbered Bone 19° 37′–9.960 ±40
(14C)
Neves and
Piló(2003)
C. cuvieri Unnumbered Calcite 19° 37′–27.1 ±3.400
(Th/U)
Auler et al.
(2006)
E. (A.)
neogaeus
Unnumbered Bone 19° 37′–16.900 ±70
(14C)
Neves and
Piló(2003)
E. (A.)
neogaeus
Unnumbered Bone 19° 37′–16.250 ±60
(14C)
Neves and
Piló(2003)
E. (A.)
neogaeus
Unnumbered Bone 19° 37′–16.180 ±70
(14C)
Neves and
Piló(2003)
(continued)
The Brazilian Intertropical Fauna from 60 to About 10 ka B.P. …217
greslebini Castellanos, 1941, Panocthus jaguaribensis (Moreira 1965),
Hoplophorus euphractus Lund (1839), and Glyptotherium sp. Oliveira et al. (2010a
designated as Glyptotherium sp. for all the material previously considered as part of
the Glyptodon genus of this region.
Additionally, Pachyarmatherium brasilense Porpino, Bergqvist and Fernicola
(2009), which is a cingulate showing characteristics from both armadillos and
glyptodonts, has also been recorded in the BIR. Downing and White (1995) assigned
myrmecophagous habits to Pachyarmatherium leiseyi (Downing and White 1995),
and it is herein believed also true for the Brazilian species P. brasilense.
3.5 Order Proboscidea
Only one species of Proboscidea is currently known for the BIR, N. platensis
(Ameghino 1888). Recent studies suggest that this species lived in groups (Mothé
et al. 2010), likely formed by adult females and their youngsters, and possibly other
young individuals, in a similar structure to what is currently observed for living
elephant populations. This species had a body mass of about four tons, and their
diet consisted of grasses and shrubs (C
3
and C
4
plants; Table 2), being considered
as generalists (Fariña et al. 1998;Sánchez et al. 2004; Asevedo et al. 2012).
3.6 Order Notoungulata
Two taxa of Notoungulata are recorded in the BIR: Toxodon platensis (Owen 1840)
and Piauhytherium capivarae (Guerin and Faure 2013). Both species were grazers
(Cartelle 1999) and are believed to have shared similar diets and body masses of
about 1100 kg (Fariña et al. 1998). However, MacFadden (2005) stated that tox-
odonts presented a large variability in their diet, depending on the habitat. In the
BIR, these species may have had more exclusive diets, mainly based on grasses and
herbaceous plants, or mixed diets involving C
3
and C
4
plants, or even diets
exclusively based on C
3
plants (Table 2).
Table 2 (continued)
Taxa Sample Material Latitude
13
C (CO
3
)
‰
VPDB
Dating
technique
Reference
H.
euphractus
Unnumbered Calcite 19° 37′–14.849 ±711
(Th/U)
Auler et al.
(2006)
S. populator Unnumbered Bone 19° 37′–9.130 ±150
(14C)
Neves and
Piló(2003)
218 M.A.T. Dantas and M.A. Cozzuol
3.7 Order Perissodactyla
Cartelle (1999) described two species of Perissodactyla occurring in the BIR, Equus
(Amerhippus)neogaeus (Lund 1840) and Hippidion principale (Lund 1846), gen-
erally found in association, in fossiliferous outcrops (i.e., “tanks”and caves; Table 1).
Both species are considered grazers with body masses of about 300 kg (Fariña
et al. 1998). However, data from carbon isotope studies, although scarce, have
shown that in the low latitudes, these species likely fed predominantly on C
4
grasses,
such as it has been observed in the state of Bahia (12° S, latitude) in Equus
(Amerhippus)neogaeus [δ
13
C from −0.6 to 1.7 ‰; MacFadden et al. (1999)].
However, in the Argentine pampas (35° S, latitude), studies have shown mixed diets,
composed of C
3
and C
4
grasses, tending to a C
3
grass predomination, for H. prin-
cipale (δ
13
C from −12.05 to −8.08 ‰) and Equus (Amerhippus)neogaeus (δ
13
C
from −11.46 to −7.21 ‰) (MacFadden et al. 1999;Sánchez et al. 2006), which
indicates they likely inhabited open areas. Nevertheless, Bernardes et al. (2013)
suggested that these species could coexist with low trophic superposition levels,
because H. principale might have been more selective for softer plant tissues.
3.8 Order Artiodactyla
Two fossil species of Camelidae are known for the BIR: Palaeolama major Liais,
1872 and Palaeolama sp. (Marcolino et al. 2012). Palaeolama (Hemiauchenia)
niedai (Guérin and Faure 1999) is believed to be a junior synonym of P. major
(Scherer 2009). Marcolino et al. (2012) reviewed the diet of this taxon, and also
presented new data regarding the analysis of mummified coprolites found in
association with a skeleton of Paleolama major, suggesting a diet composed of
shrubs (C
3
plants). According to these authors, this taxon likely lived in open areas.
3.9 Order Litopterna
According to Cartelle (1999), only one litoptern species is known for the BIR:
Xenorhinotherium bahiense (Cartelle and Lessa 1988). However, Guérin and Faure
(2004) believed this species does not differ from Macrauchenia patachonica,
assigning fossils found in the Piauístate to the latter. Given that this discussion is
beyond the scope of this study, the systematic proposal of Cartelle (1999) will be
considered here.
Cartelle and Lessa (1988) assigned to X. bahiense a diet based on grasses and
herbaceous plants, what is supported by previous research about the Pleistocene
flora in the region where the type specimen was found (i.e., state of Bahia), and
because the specimen of the BIR was found in an aggregation with other species of
grazing megamammals. Fariña et al. (1998) assigned to M. patachonica a body
mass of about 1000 kg, which we also tentatively suggest for X. bahiense.
The Brazilian Intertropical Fauna from 60 to About 10 ka B.P. …219
3.10 Dating Review
The South American time scale for the Pleistocene was established on the
Argentine pampas region, where four land mammal ages were recognized for this
epoch: the “Ensenadan,”“Bonaerian,”“Lujanian,”and “Platan”stages (Cione and
Tonni 1999). The fauna from the BIR is generally assigned to the Lujanian stage
(Cartelle 1999). However, available studies presenting numerical data, despite
punctual, show that this fauna has been present in the area for a longer period, from,
at least, 350–9 ka. This suggests that this fauna was coeval with the Bonaerian,
Lujanian, and the beginning of the Platan land mammal ages.
It is recognized therefore that this fauna has lived in the BIR during the various
climatic changes occurred during the Pleistocene/Holocene. The inferred ecology
for these animals suggests they were adapted to open environments, such as those
found today for the Caatinga and Cerrado biomes. Additionally, it is likely that
during drier periods, they would have extended their distribution, whereas the
opposite seems to have taken place during the more humid periods, as suggested by
Cione et al. (2007).
Between 60 and about 10 ka, two distinct climatic moments are recognized for
the BIR. Data from 93 to 47 ka, collected from stalagmites using δ
18
O (Wang et al.
2004; de Barreto 2010), indicate a prolonged warmer and drier period, with short
intervals of higher moisture. During this period, it is likely that the areas of the
Caatinga and Cerrado would have expanded geographically and thus, connected to
each other. Evidence indicates that the Caatinga already existed in this region 42 ka
(De Oliveira et al. 1999; Behling et al. 2000), along with some of the megafauna
taxa, such as P. humboldti,H. euphractus,N. platensis, and, probably, also C.
cuvieri (Table 2), which lived during this period.
Palynological and δ
18
O data available for the period from 40 to 10 ka are not
continuous, although they seem to indicate a long period of wetness and predom-
ination of forests, possibly forming a connection between the Amazon and Atlantic
Forests (Behling et al. 2000; Auler and Smart 2001; Sifeddine et al. 2003;de
Barreto 2010). For this period, there are records in the BIR of E. laurillardi,C.
cuvieri,Nothrotherium,T. platensis,N. platensis, and S. populator (Table 2).
4 Discussion
4.1 Paleoenvironmental Reconstruction of the BIR at About
64 ka
Data about the feeding ecology of the megafauna from about 60 ka in the southern
BIR are restricted to taphonomic and paleoecological studies of N. platensis, from
the fossiliferous outcrop “Águas do Araxá,”in Minas Gerais. These gomphotheres
inhabited a dry environment, with well-defined seasons, at about 64 ka, (Avilla
220 M.A.T. Dantas and M.A. Cozzuol
et al. 2013). Fragments of conifers and grasses were found associated to teeth
remains of this species, suggesting a colder and drier season; whereas barite min-
erals encountered in their bones are recognized as indicators of warmer seasons
(Avilla et al. 2013; Dominato 2013). Palynological data from locations above
900 m a.s.l., near Araxá, corroborate this climatic environmental pattern.
4.2 Paleoenvironmental Reconstruction for BIR at About
10–27 ka
Published isotopic carbon ratio (δ
13
C) data are available for this period in the BIR,
in relation to the following taxa: E. laurillardi (Lund 1842); V. bucklandi (Lund
1839); N. platensis (Ameghino 1888); T. platensis (Owen 1840); and Equus
(Amerhippus)neogaeus (Lund 1840). Although the available studies are few, δ
13
C
analyses suggest E. laurillardi had a mixed diet, consuming large amounts of C
3
plants, exploring the edges of forests, and feeding on fruits and herbaceous species
across the BIR (Table 2; Fig. 2).
N. platensis and T. platensis were grazers between latitudes 5° 49′S and 6° 15′S;
whereas V. bucklandi lived near the edges of forests, feeding on C
3
plants (Table 2;
Fig. 2). Between the latitudes 9° 22′S and 10° 17′S, N. platensis is known to have a
diet exclusively based on C
4
grasses, whereas T. platensis had a mixed diet, although
apparently favoring C
4
plants (Table 2; Fig. 2). Finally, between latitudes 10° 21′S
and 14° 46′S, E. neogeus was a grazer, while T. platensis and N. platensis had mixed
diets, tending to higher consumption of herbaceous C
3
plants (Table 2; Fig. 2).
These results suggest that about 27–11 ka, between the latitudes 14° S and 5° S,
the BIR presented a gradual environmental change, from more open (where grasses
and herbaceous plants predominated) to more forested ones. A recent
Fig. 2 Tooth enamel carbon isotope ratio (δ
13
C) values for five species in Brazilian Intertropical
Region
The Brazilian Intertropical Fauna from 60 to About 10 ka B.P. …221
biogeographical proposal for N. platensis suggests this species was adapted to dry
seasonal forest environments (i.e., Caatinga, Cerrado; Dantas et al. 2013b), what
may be considered as indicative that similar adaptations also happened to the
remaining species of the megafauna of the region.
5 Final Remarks
This chapter represents the first step improving our knowledge about the paleoe-
cology of the megafauna and climatic environmental patterns occurring during the
Pleistocene in the BIR. Much, however, is still to be done, and it is believed that
further research in this area is highly promising.
Acknowledgments To Flavia Franchini (Memorial University of Newfoundland) for the English
review of the manuscript. To the anonymous reviewers which corrections and suggestions
improved the quality of this manuscript.
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