Journal of South American Earth Sciences 96 (2019) 102362

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Journal of South American Earth Sciences

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Late Pleistocene meso-megamammals from Anagé, Bahia, Brazil: Taxonomy T

and isotopic paleoecology (δ13C)
Juliana de Almeida da Silvaa,∗∗, Luciano Artemio Lealb, Alexander Cherkinskyc,
Mário André Trindade Dantasa,d,∗
a
Programa de Pós-graduação em Genética, Biodiversidade e Conservação, Universidade Estadual do Sudoeste da Bahia – Campus Jequié, Jequié, BA, Brazil
b
Laboratório de Geociências, Universidade Estadual do Sudoeste da Bahia – Campus Jequié, Jequié, BA, Brazil
c
Center for Applied Isotope Studies, University of Georgia, Athens, GA, 30602, USA
d
Laboratório de Ecologia e Geociências, Universidade Federal da Bahia (IMS/CAT), Vitória da Conquista, BA, Brazil

A R T I C LE I N FO A B S T R A C T

Keywords: The fossils of Pleistocene megafauna are commonly found in tanks and cave deposits in the State of Bahia,
Brazilian intertropical region northeastern Brazil. In general, studies on these materials have focused mainly on taxonomic aspects, but more
Tanks recently, there has been an increase in paleoecological studies based on stable isotopes. The present paper
Isotopic paleoecology describes fossils recovered from a tank deposit in Lagoa de Pedra (Anagé municipality, Bahia State). The material
Megafauna
was identified as: Eremotherium laurillardi, Panochthus sp., Holmesina paulacoutoi, Palaeolama major,
Notiomastodon platensis, and Toxodontinae indet. Stable isotope analyses were performed in order to estimate
their diet and the paleoenvironment in which these taxa lived. The results indicate the occurrence of two guilds:
grazers (Panochthus sp.) and mixed-feeders (H. paulacoutoi, N. platensis and E. laurillardi). These animals lived in
a more open environment where N. platensis possibly had a major role in the community organization.

1. Introduction
Fossils of Pleistocene mesoherbivores (body mass between 100 and
750 kg) and megaherbivores (body mass > 800 kg) are commonly
found in “tank” and cave deposits (e.g.: Cartelle, 1992; Lobo et al.,
2015) in the State of Bahia, northeastern Brazil (Coe et al. 1976). The
tanks are depressions naturally excavated in Neo-Mesoproterozoic
crystalline rocks that contain the remains of animals and plants that
were carried into the tanks during rainy seasons. These fossil records
from Bahia were already found in Toca dos Ossos (Ourolândia muni-
cipality; Cartelle, 1992), Toca da Boa Vista (Campo Formoso munici-
pality; Cartelle, 1994), and Toca das Onças (Jacobina municipality;
Cartelle and De Iullis, 2006) caves, and in the tanks of Fazenda Suse II
(Vitória da Conquista municipality; Dantas and Tasso, 2007); Fazenda
Caraíba (Coronel João Sá municipality; Dantas and Zucon, 2007), Fa-
zenda Tuquinha (Palmas de Monte Alto municipality; Dantas et al.,
2008), Lagoa do Rumo (Baixa Grande municipality; Ribeiro and
Carvalho, 2009), Povoado Sítio (Quinjingue municipality; Bertoni-
Machado et al., 2011), Sítio Novo (Matina municipality; Lobo
et al.,2015), and Guanambi municipality (Scherer et al., 2017).
The studies on the fossil material recovered from the aforemen-
tioned localities have focused mainly on taxonomic identification.
However, more recent studies (e.g.: Dantas et al., 2017 and references
therein) have intended to better understand the paleoecology of these
animals through isotopic analyses.
In the last two decades there has been an increase in studies on the
paleodiet of the South American megafauna based on isotopic (δ13C)
data. The published studies characterized the feeding patterns and the
paleoenvironment in which some megafauna taxa lived (França et al.,
2015).
In this sense, the present paper has the following objectives: (i) to
present the taxonomic identification of the material discussed here; (ii)
to assess the diets and niche width of the studied taxa through carbon
isotope (δ13C) analysis; and, (iii) to reconstruct the paleoenvironment
in which these taxa lived.

∗
Corresponding author. Programa de Pós-graduação em Genética, Biodiversidade e Conservação, Universidade Estadual do Sudoeste da Bahia – Campus Jequié,
Jequié, BA, Brazil.
∗∗
Corresponding author.
E-mail address: matdantas@yahoo.com.br (M.A.T. Dantas).

https://doi.org/10.1016/j.jsames.2019.102362
Received 10 January 2019; Received in revised form 25 August 2019; Accepted 11 September 2019
Available online 12 September 2019
0895-9811/ © 2019 Elsevier Ltd. All rights reserved.
J.d.A.d. Silva, et al.
Fig. 1. Location map of Anagé Municipality, Bahia, and the delimitation for the Brazilian Intertropical Region (modified from Dantas et al., 2013).
2. Material and methods
2.1. Study material
The studied material was recovered from a tank located in the Lagoa
de Pedra site, Anagé municipality, Bahia State (Fig. 1; coordinates UTM
24L 0271258 E; 8382652 S, Datum WGS84).
All material is currently housed at the scientific collections of the
Laboratory of Geosciences (Jequié campus) and of the Laboratory of
Geology (Vitória da Conquista campus), Universidade Estadual do
Sudoeste da Bahia – UESB, Bahia State, Brazil. The fossils were analyzed
in these laboratories and, when necessary, the specimens were me-
chanically prepared with the aid of proper instruments. The fragmented
specimens were glued with multipurpose instant adhesive and water-
proofed with shellac.
2.2. Taxonomy
The specimens were anatomically identified following, whenever
possible, Paula-Couto (1979). The taxonomic identification was made
in accordance with the groups: Order Tardigrada (e.g. Cartelle, 1992),
Order Cingulata (e.g. Porpino and Bergqvist, 2002), Order Notoungu-
lata (e.g. Roth, 1898), Order Cetartiodactyla (e.g. Scherer, 2013), and
Order Proboscidea (e.g. Mothé et al., 2012).
2.3. Isotopic paleoecology
Samples of adult individuals of Eremotherium laurilardi (Lund, 1842)
(one dentin sample), Holmesina paulacoutoi (Guerra & Bohórquez, 1984)
(one osteoderm fragment), Panochthus sp. (one osteoderm fragment),
and Notiomastodon platensis (Ameghino, 1888) (one enamel sample)
were used to obtain stable carbon and oxygen isotope values (δ 13C,
δ18O) from bioapatite structural carbonate. It is known that bone is
more susceptible to diagenetic processes that affect the stable isotope
2
Journal of South American Earth Sciences 96 (2019) 102362
composition of fossils. Since bone was the only material of Holmesina
paulacoutoi and Panochthus sp. available to perform this analysis, the
interpretation of these results were made more carefully.
The stable isotope analyzes were performed at the Center for
Applied Isotope Studies of the University of Georgia (Georgia, USA). All
samples were cleaned with distilled water in ultrasonic bath and then
left to dry naturally. The samples were then crushed into smaller
fragments to be treated with diluted 1N acetic acid to remove surface
adsorbed and secondary carbonates. This solution was periodically re-
newed to ensure that diagenetic carbonates were removed from the
interior of the sample and that fresh acid was allowed to reach all
surfaces uniformly.
The chemically cleaned samples were then reacted under vacuum
with 100% phosphoric acid to dissolve the mineral fraction and release
carbon dioxide from the hydroxyapatite. The resulting carbon dioxide
was cryogenically purified with other reaction products and catalyti-
cally converted to graphite (Cherkinsky, 2009).
Graphite 14C/13C ratios were measured using a CAIS 0.5 MeV ac-
celerator mass spectrometer. The sample ratios were compared to the
ratio measured from the standard Oxalic Acid I (NBS SRM 4990). The
13
C/12C ratios were measured separately with respect to PDB standard
values using a stable isotope ratio mass spectrometer.
The isotopic composition of hydroxyapatite can be preserved with
minimal or no significant diagenetic alteration (e.g. Bocherens et al.,
1996; Sánchez et al., 2004). All results are reported using the delta
notation, δ = [(Rsample/Rstandard − 1)*1000] (Coplen, 1994). The re-
ference standard for carbon isotope values (R = 13C/12C) is V-PDB, and
for oxygen isotope values (R = 18O/16O) is V-SMOW.
The interpretation of carbon isotope values in meso-mega-
herbivorous mammals is based on the known average values of C3
(μδ13C = −27 ± 3‰), C4 (μδ13C = −13 ± 2‰), and CAM plants
(δ13C values intermediate between C3 and C4 plants).
We estimated carbon enrichment (Ɛ*diet-bioapatite) for Holmesina
paulacoutoi (Ɛ = 13‰), Eremotherium laurillardi and Panochthus sp.
J.d.A.d. Silva, et al.
(Ɛ = 14‰), and Notiomastodon platensis (Ɛ = 15‰) following Tejada-
Lara et al. (2018). By considering these enrichment values, δ13C values
lower than −12‰ to −14‰ are typical of animals with diets that
consist exclusively of C3 plants, while δ13C values higher than −1‰ to
+1‰ are consistent with diets based mainly on C4 plants.
The relative proportion of different plant types ingested (C3 and C4)
by the analyzed specimens was determined by the following mixed
mathematical model (1; Phillips, 2012):
δ 13Cmix = δ 13C1f1 + δ 13C2 f2 1 = f1 + f2 (1)
In order to estimate the ecological measurements, we calculated the
isotope niche breadth (B) using the equation described by Levins
(1968), where pi is the relative proportion of individuals in isotope bin I
(2). The result was then standardized (BA) from 0 to 1 following
equation (3), where n is the total number of available isotope bins.
Values lower or equal to 0.5 are compatible with a specialist diet
strategy while values above 0.5 are compatible with a generalist diet
strategy.
1 be explained by the transportation of the carcasses and their long ex-
B=
Σpi2 (2)
BA =
B−1
N−1 (3)
In addition, the average niche overlap (O) was also calculated fol-
lowing Pianka (1973) index (4), where pi is relative proportion of in-
dividuals in bin i. Results between 0 and 0.3 indicate a low niche
overlap, between 0.3 and 0.7 there is a moderate overlap, and above
0.7 there is a high niche overlap.
Σpij . pik
Ojk =
Σpij2 . Σpik2 (4)
As the isotopic data do not reflect seasonal diet shifts, we followed
Dantas et al. (2017) to interpret the overlapping values as the effect of
competition for food resources during dry seasons. We also followed
this study to interpret that taxa presenting higher body mass values
were better competitors for food resources and, thus, when niche
overlap was moderate or high between taxa, the ones with higher body
masses were more likely to prevail in that niche.
Although there are no datings for the studied locality, the paleoe-
cological data from other tanks in northeastern Brazil indicate en-
vironmental stability and the available datings suggest a small time
variation within the fossils in those tanks (~32 ka, Poço Redondo,
Sergipe; França et al., 2014). Hence, based on this information, we
estimated the niche overlap of the studied taxa and proposed a pa-
leoenvironmental reconstruction (through carbon isotopic data) for the
Lagoa de Pedra site.
3. Results and discussion
3.1. Taxonomic composition, NISP, MNI and ontogenetic stage
The Lagoa de Pedra tank was excavated before the collection of the
fossil material, making it impossible to know which specimens were
buried in the same stratum or period. The Anagé fossil assemblage is
multitaxon and monodominant. Eremotherium laurillardi is the dominant
species and has the highest Number of Identifiable Skeletal Parts - NISP
recorded (91) here, followed by Panochthus sp. (NISP = 20), P. major
and N. platensis (NISP = 2), and H. paulacoutoi, indeterminate
Toxodontinae and Equidae (NISP = 1) (Table 1). The Minimum
Number of Individuals - MNI of E. laurillardi was three, represented by
three left radius (UESB 318PV06, UESB 318PV19, and UESB 318PV61)
and three right ulnas (UESB 318PV04, UESB 318PV07, and UESB
318PV20). All other taxa have MNI equals to one (Table 1).
The prevalence of E. laurillardi compared to other taxa may be the
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Journal of South American Earth Sciences 96 (2019) 102362
Table 1
Number of Identifiable Skeletal Parts (NISP), Minimum Number of Individuals
(MNI) and ontogenetic stage for taxa found in “Lagoa da Pedra”, Anagé, Bahia.
Taxa NISP MNI Ontogeny
Eremotherium laurillardi 99 3 adult and subadult
Panochthus sp. 20 1 adult
Holmesina paulacoutoi 1 1 –
Palaeolama major 2 1 adult
Notiomastodon platensis 3 1 adult
indeterminated Toxodontinae 1 1 –
result of the high resistance of its skeletal elements (Araújo-Júnior
et al., 2013) and the likely abundance of extant individuals during the
Pleistocene.
All materials were found disarticulated and most are fragmented or
fractured. A total of 117 specimens are partially complete, which al-
lowed their anatomical identification. Other 80 specimens are uni-
dentified fragments. The fragmentary condition of the material might
posure to environmental conditions, which eventually lead to the dis-
articulation of the skeletal elements (Araújo-Júnior et al., 2013) before
the final burial.
3.1.1. Eremotherium laurillardi (Lund, 1842)
Most of the specimens belongs to this taxon. The identified material
consists of two partial skulls (UESB 318PV29; UESB 318PV30), two
fragments of maxilla (UESB 318PV32; UESB 318PV38), isolated teeth
(e.g.: UESB s/n; UESB 318PV98), and other several postcranial elements
(Fig. 2A–E).
The presence of bones with epiphyses fused to the diaphyses in-
dicate that this is an assemblage composed of both adult and subadult
individuals (fragments of bone epiphyses are not anatomically identi-
fiable; UESB 318PV71, UESB 318PV72, and UESB 318PV74 has circular
depressions caused by growth cartilage).
The material was assigned to Eremotherium laurillardi due to the
following characters: presence of four prismatic molariforms in the
mandible (Fig. 2D–E), maximum height of the mandibular body lower
than the lengths of the m1-m4 (Fig. 2D–E), mandibular symphysis
ending close to the m1 (Fig. 2D–E), and tibia-fibula fused only proxi-
mally (not figured). In addition, the locality where this material was
found falls in the area of occurrence of that taxon, as recognized by
Cartelle and De Iullis (1995). Cartelle and De Iullis (1995, 2006) also
reinforce that E. laurillardi is a single intertropical species with Pan-
American distribution, from southern Brazil to northeastern United
States.
3.1.2. Panochthus sp.
The materials assigned to this taxon consist of the distal portion of
one left humerus (UESB 318PV174) and 19 isolated osteoderms (UESB
318PV175 – UESB 318PV194; Fig. 3A–D). The humerus belonged to an
adult individual, since the epiphysis and the diaphysis are completely
fused.
The group Panochthini is composed by glyptodonts endemic to
South America. In Brazil, this taxon occurs in the southern and north-
eastern regions (Ferreira et al., 2015). There are four Panochthus species
described to northeast Brazil (P. greslebini, P. jaguaribensis, P. oliveira-
roxoi, and P. rochai). According to Porpino et al. (2014), P. greslebini and
P. jaguaribensis are valid species within Panochthus and their diagnoses
are based on the characters of the caudal tube. The authors also argue
that P. rochai is a synonym of P. greslebini and that P. oliveira-roxoi is a
nomen dubium.
The osteoderms studied here exhbit homogeneous ornamental fig-
ures randomly distributed and no central figure, similar to those of
Panochthus greslebini described by Porpino and Bergqvist (2002).
However, isolated osteoderms cannot be identified at the species level
J.d.A.d. Silva, et al.
Fig. 2. Eremotherium laurillardi. Skulls fragments (A) 318PV29 and (B) 318PV30 in dorsal view; Dentary fragment (C) 318PV32 in occlusal view; Right dentary UESB
s/n in lateral (D) and occlusal (E) views.
(Porpino et al., 2014) and, thus, the material was regarded as Pa-
nochthus sp.
3.1.3. Holmesina paulacoutoi (Cartelle & Bohorquez, 1984)
The presence of this taxon in the assemblage is based on an osteo-
derm fragment of the moveable bands (UESB 318PV198; Fig. 3E).
According to Cartelle (1992), H. paulacoutoi is an autochthonous
species in the Brazilian Intertropical Region (BIR). Two genera of
pampatheriids are recognized in the Pleistocene deposits of South
America: Pampatherium and Holmesina. Both genera are distinguishable
by the morphology of their osteoderms (Rincon and White, 2007). The
osteoderm described here has a depressed marginal area and lacks a
central figure, which is morphologically similar to those already de-
scribed for Holmesina paulacoutoi.
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Journal of South American Earth Sciences 96 (2019) 102362
3.1.4. Toxodontinae Trouessart, 1898
The specimen assigned to this taxon is a complete and well pre-
served cuboid (UESB 318PV199; Fig. 3F–J).
Nasif et al. (2000) proposed a classification for Toxodontidae based
on cladistic analysis and using two subfamilies, Nesodontinae and
Toxodontinae. Guérin and Faure (2013) acknowledge two genera of
Toxodontinae in the Late Pleistocene of Intertropical Brazil: Toxodon
and Piauhytherium.
The cuboid has characters similar to those described for Toxodon
burmeisteri (= Toxodon platensis) by Roth (1898). Yet, because this
anatomical component is not considered diagnostic for this species or
genus, we assigned it as an indeterminate Toxodontinae.
3.1.5. Palaeolama major (Liais, 1872)
The material assigned to this species is a right humerus (UESB
J.d.A.d. Silva, et al. Journal of South American Earth Sciences 96 (2019) 102362

Fig. 3. Panochthus sp., left humerus UESB 318PV174 in ventral (A), (B) dorsal and distal (C) view, and osteoderms UESB 318PV194 (D); Holmesina paulacoutoi
fragment of mobile band osteoderm UESB 318PV198 (E); indetermined Toxodontinae, cuboid UESB 318PV198, proximal (A); lateral (B–D); dorsal (C) and distal (E)
views.

318PV172) and a right femur (UESB 318PV173; Fig. 4A–D) that be-
longed to adult individuals as the diaphyses and epiphyses are com-
pletely fused.
The humerus exhibits in its proximal diaphisys a wide and small
tuberosity separated by a condyle and a well developed delto-pectoral
crest that ends nearly halfway through the diaphysis. In anterior view,
the distal epiphysis presents a shallow pseudocoronoid fossa. In lateral
view, there are condylar crests and the olecranon forms a deep fossa –
shaped as an inverted “V” – that articulates with the ulna.
The femur is incomplete, lacking the lateral and medial condyles of
the distal trochlea. In anterior view, it is possible to observe the prox-
imal articulation (head of the femur).
Although we lack any specific diagnostic material, the morphology
and size of the humerus and femur (UESB 318PV172 and UESB
318PV173; Table 2) indicate that these specimens belonged to Palaeo-
lama major.
3.1.6. Notiomastodon platensis (Ameghino, 1888)
The material assigned to this taxon consists of two complete right
metacarpals II (UESB 318PV195) and III (UESB 318PV196) (Fig. 4E–H)
and 1 M fragment (UESB 318PV197). The metacarpal II is longer than
the metacarpal III, but both are robust (Table 2). The molar tooth
fragment presents a thick whitish enamel layer.
The family Gomphotheriidae is a Proboscidea group that arrived in

5
J.d.A.d. Silva, et al. Journal of South American Earth Sciences 96 (2019) 102362

Fig. 4. Palaeolama major, right humerus UESB 318PV172 in ventral (A) and dorsal (B) views; right femur UESB 318PV173 in ventral (A) and dorsal (B) views.
Notiomastodon platensis, right metacarpal II (UESB 318PV195) in dorsal (E) and ventral (F) views; right metacarpal III (UESB 318PV196) in dorsal (G) and ventral (H)
views.

Table 2
Measurements (in mm) of poscranial bones for Palaeolama major and Notiomastodon platensis from “Lagoa de Pedra”, Anagé, Bahia. Labels. (1)
Cartelle (1992);
(2)
Molena (2012); ∗incomplete measurement.
Taxa specimen (n) mean ± sd

length Breadth of

proximal end distal end

P. major Humerus (UESB 318PV172) (1) 353.75 (1) 97.80 (1) 75.30
Humerus(1) (8) 327.75 ± 21.40 (8) 105.13 ± 5.00 (8) 76.75 ± 5.26
Femur (UESB 318PV172) (1) 422.60* (1) 106.20 –
Femur (1) (8) 422.88 ± 10.20 (8) 103.38 ± 4.31 (8) 89.63 ± 4.63
N. platensis Mc II (UESB 318PV195) (1) 144.00 (1) 67.30 (1) 75.10
Mc II(2) (5) 123.28 ± 13.22 (5) 59.96 ± 8.94 (5) 71.00 ± 9.75
Mc III (UESB 318PV196) (1) 124.75 (1) 79.87 (1) 77.16
Mc III(2) (7) 148.31 ± 21.69 (7) 74.49 ± 13.67 (7) 75.56 ± 9.47

South America during the Great American Biotic Interchange (GABI) the non-diagnostic molar fragment. Therefore, UESB 318PV195, UESB
(Mothé et al., 2012). The diagnostic characters that define the genus 318PV196 and UESB 318PV197 could possibly belong to Notiomastodon
and species of Gomphotheriidae are mainly based on the morphology of platensis, but this inference needs to be considered carefully.
their incisive and molar teeth. The only dental material in our study is

6
J.d.A.d. Silva, et al. Journal of South American Earth Sciences 96 (2019) 102362

Fig. 5. (A) Principal Component Analyses (PCA) using δ13C and δ18O values for meso-megaherbivores from Anagé, Bahia; (B) Dendogram with clustering of taxa
according to their ecological guilds (Bootstrap test, N = 10.000).

3.2. Isotopic paleoecology
Based on the observations of Dantas et al. (2017) for extant meso-
herbivores and megaherbivores from Africa, and in the cluster and PCA
analyses performed here with the carbon and oxygen isotope values
(Fig. 5), we suggest that the taxa from Anagé belonged to two guilds:
grazers, that consumed more than 80% of C4 plants; and mixed-feeders,
with some taxa that had a higher consumption of C4 grasses and other
taxa that consumed more C3 plants.
Panochthus sp. (δ13C = 0.51‰) was assigned to the grazer guild and
considered as a specialist mesoherbivore (BA = 0.07; body
mass = 785 Kg) with a diet consisting of 96% of C4 plants. The group of
mixed-feeders was divided in two subgroups based mainly on the
oxygen isotope values. Holmesina paulacoutoi (δ13C = −6.86‰;
piC4 = 51%; BA = 1.00; bm = 120 Kg) and Eremotherium laurillardi
(δ13C = −9.20‰; BA = 0.47; piC4 = 20%; bm = 2212 Kg) had δ18O
values near 30‰, while Notiomastodon platensis, with had a major
consumption of C4 grasses (pi = 52%; δ13C = −4.72‰; BA = 1.00;
bm = 4940 kg) showed δ18O values higher than 32‰ (Table 3).
The carbon isotope data of Panochthus sp. and H. paulacoutoi from
other localities are not available for comparison. However, the data of
E. laurillardi and N. platensis are similar to those found in other fossi-
liferous localities from Bahia, such as Quijingue (E. laurillardi,
δ13C = −5.52‰; BA = 0.91) and Ourolândia (N. platensis,
μδ13C = −6.60 ± 1.77‰; μBA = 0.72 ± 0.35) (Sánchez et al., 2004;
Dantas et al., 2017). These results support the interpretation of Dantas
et al. (2013) that in the lower latitudes of the BIR these taxa used to
consume more C3 plants.
Overall, the niche overlap between taxa was high, varying from
0.75 to 1.00, due to the high consumption of C4 grasses. The only ex-
ception was E. laurillardi and Panochthus sp., which presented a mod-
erate niche overlap (0.28; Table 3).
In Africa, megaherbivores like Loxodon africana (bm = 5000 Kg)
that have a mixed-feeding diet are key species in the community or-
ganization. Their generalist diets allow them to feed both on C3 and C4
plants. Thus, these animals occupy a broad niche breadth
(BA = 0.71 ± 0.29; Dantas et al., 2017), which suppresses the popu-
lation of taxa that are exclusively grazers or browsers.

Table 3
Body mass (kg), Carbon (δ13C) and oxygen (δ18O) isotopic values, proportional contributions (pi) of food sources (C3 or C4 plants), standardized isotopic niche
breadth (BA) and niche overlap (O) for Late Pleistocene megamammals from Anagé/BA. (1) our data.
Taxa bm (kg) δ13C (‰) δ18O (‰) pi BA O

C3 C4 El Hp P Np

Anagé/BA El 2,212(1)
−9.20 30.54 0.80 0.20 0.47 – 0.85 0.28 0.84
Hp 120(1) −6.86 30.05 0.49 0.51 1.00 0.85 – 0.75 1.00
P 785(1) 0.51 29.11 0.04 0.96 0.07 0.28 0.75 – 0.76
Np 4,940(1) −4.72 32.66 0.48 0.52 1.00 0.84 1.00 0.76 –

Legends. El - Eremotherium laurillardi, Hp - Holmesina paulacoutoi, P - Panochthus sp., Np - Notiomastodon platen.

7
J.d.A.d. Silva, et al.
In the Pleistocene of Anagé, the key species role was possibly oc-
cupied by N. platensis, as it fed equally on C3 and C4 plants (Table 3),
had a broad niche breadth (BA = 1.00), and their estimated body
masses (bm = 4940 kg) were high relative to other taxa from the same
locality.
By assessing the δ13C values of the meso-megamammals from
Anagé, we suggest that the environment in which these animals lived
was composed of open forests areas, as three taxa showed an estimated
consumption of more than 51% of C4 plants (Table 3).
4. Final remarks
The taxonomic composition of the specimens analyzed here was
similar to those found in other tanks in Bahia. The Lagoa de Pedra tank
in Anagé comprises one of the most diverse fossil assemblages in the
Bahia State, with a record of seven taxa.
Stable isotopes applied in paleoecological studies help us to better
understand the diet and the paleoenvironment in which ancient taxa
lived. In the present study, the stable isotope data show the presence of
two guilds in Anagé: grazers (Panochthus sp.) and mixed-feeders (H.
paulacoutoi, N. platensis and E. laurillardi). These animals lived in an
open environment where N. platensis could have had a major role in the
organization of this mammal assemblage.
Acknowledgments
The authors would like to thank the FAPESB for the scolarship
granted to J.A. Silva (Process 11554/2015) as well as the CNPq for
granting the research fellowship and the financial support for the stable
isotope analyses of the projects approved for M.A.T. Dantas (PQ/CNPq
308122/2016-0; Universal process 404684/2016-5). We would also
like to thank Thais Rabito Pansani and Camila Bernardes for translating
and revising the english of this manuscript; and the referees for their
valuable critics and suggestions that allowed the improvement of our
paper.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.jsames.2019.102362.
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