J Mammal Evol

DOI 10.1007/s10914-016-9328-y

ORIGINAL PAPER

Baraguatherium takumara, Gen. et Sp. Nov., the Earliest

Mylodontoid Sloth (Early Miocene)
from Northern South America
Ascanio D. Rincón 1 & Andrés Solórzano 1 & H. Gregory McDonald 2 &
Mónica Núñez Flores 1

# Springer Science+Business Media New York 2016

Abstract We report a new genus and species of sloth, based South America. The presence of a vasodentine layer in the
on a partial mandible and associated femur, from the early core of the tooth is quite similar to Octodontotherium,
Miocene of Venezuela. Baraguatherium takumara, gen. et Paroctodontotherium, and Orophodon and allows assignment
s p . n o v. , r e p r e s e n t s t h e e a r l i e s t m e m b e r o f t h e of this new taxon to the Mylodontoidea.
Mylodontoidea recognized from northern South America.
Phylogenetically and morphologically, Baraguatherium pos- Keywords Baraguatherium takumara . Castillo Formation .
sesses some plesiomorphic characters: a vasodentine layer in Early miocene . Venezuela . Sloth diversity
the core of the tooth similar to Octodontotherium,
Paroctodontotherium, and Orophodon; molariforms parallel
to the long axis of the toothrow; teeth with a very thin layer Introduction
of cementum; mf1-mf3 series of similar size and bilobate; mf3
conspicuously piriform; and occlusal surface of tooth beveled, The isolation of South America as an island continent, through
which places it at the base of the Mylodontidae clade. most of the Cenozoic, resulted in the evolution of a number of
Baraguatherium was found in continental deposits that also distinctive endemic linages of mammals (Woodburne 2010;
preserve abundant wood and leaves associated with a near Goin et al. 2012). Among these, one of the more distinctive
shore marine complex, indicating that Baraguatherium lived and diverse are the Xenarthra, which today includes anteaters,
in a coastal tropical forest in the early Miocene in northern armadillos, and sloths, and two extinct clades, glyptodonts
and pampatheres (McDonald 2003). Within the Xenarthra,
sloths are an extremely diverse lineage in terms of the number
* Ascanio D. Rincón
of genera (more than 90 named genera; McKenna and Bell
paleosur1974@gmail.com 1997; Wilson and Reeder 2005), which along with many sub-
sequently described genera, display a wide range of body sizes
Andrés Solórzano and diversity of locomotor and feeding adaptations that is
solorzanoandres@gmail.com reflected in the variety of habitats in which they lived
H. Gregory McDonald (Muizon and McDonald 1995; White 1997; McDonald
hmcdonald@blm.gov 2005; McDonald and De Iuliis 2008; Shockey and Anaya
Mónica Núñez Flores 2010; Toledo et al. 2014).
nuez.monica@gmail.com The earliest putative sloth, Pseudoglyptodon, comes from
the late Eocene of Cerro Blanco, Argentina (McKenna et al.
1
2006; Gaudin and Croft 2015), and specimens referred to this
Laboratorio de Paleontología–Centro de Ecología, Instituto
Venezolano de Investigaciones Científicas (IVIC), Km 11 de la
taxon are also known from the early Oligocene of Chile (Wyss
Carretera Panamericana, Edo. Miranda. Aptdo. 21.827, et al. 1990) and Argentina (Dozo et al. 2014), and the late
Caracas 1020-A, Venezuela Oligocene of Bolivia (Engelmann 1987; Pujos and De Iuliis
2
Bureau of Land Management, Utah State Office, 440 West 200 2007). Sloths became more diversified during the late
South, Salt Lake City, UT 84101, USA Oligocene (Deseadan SALMA), based mainly on records
 J Mammal Evol

from Argentina and Bolivia, and by that time are represented yet known. To date, the earliest records of sloths in northern
by several distinct lineages (e.g., Pujos and De Iuliis 2007; South America are from the early Miocene portion of the
McDonald and De Iuliis 2008; Shockey and Anaya 2010). Castillo Formation, Venezuela, and are based only on frag-
The greatest diversity of sloths is documented for the early mentary material (‘Tardigrada indeterminate’ sensu Sánchez-
Miocene (Santacrucian in South America) and the Pleistocene Villagra et al. 2004), or not-yet appropriately described spec-
(Ensenadan and Lujanian in South America, Irvingtonian, and imens (‘Orophodontidae indeterminate’ sensu Rincón et al.
Rancholabrean in North America) (McKenna and Bell 1997). 2014). The only other sloth, Pseudoprepotherium
Whether the greater diversity during these intervals is merely venezuelanum, is from the ‘early’ Miocene of Río Yuca,
an artifact of the number of known sites or actually reflects Venezuela, and is based on a single femur (Collins 1934).
periods of evolutionary diversification is not known at this But it must be emphasized that the age of this taxon needs to
time (Rincón et al. 2015). be confirmed (Rincón et al. 2014).
In contrast to the historically large number of extinct sloth The purpose of this paper is to provide a detailed morpho-
taxa described from the southern part of South America (pri- logical description of the specimen (recovered from the early
marily Argentina), the fossil record of sloths in the northern Miocene bed of Castillo Formation) previously recognized as
portion of the South America has not been studied in similar ‘Orophodontidae indeterminate’ by Rincón et al. (2014),
detail, and is not as well known. Recent research, however, which represents a new taxon distinct from other members
has demonstrated that during the Neogene the sloths in the of the Mylodontoidea. In order to provide a preliminary un-
northern part of the continent were much more diverse taxo- derstanding of its broader relationships to other sloths, we also
nomically and ecomorphologically than previously thought. provide a phylogenetic hypothesis for the new taxon based on
In northern South America, sloth remains are best known from cranial and postcranial features.
middle and late Miocene localities. The three best documented
faunal assemblages, with the greatest taxonomic diversity of
sloths includes: 1) La Venta (middle Miocene, Laventan Geological Setting
SALMA, 11.8–13.5 Ma, Colombia), with eight or nine spe-
cies from at least four families, including Megatheriidae, The Castillo Formation was deposited in the Falcón Basin in
Nothrotheriidae, Megalonychidae, and Mylodontidae northwestern Venezuela, which covers the state of Falcón and
(Hirschfeld 1985; McDonald 1997); 2) Solimões Formation parts of the states of Zulia, Lara, and Yaracuy (Rincón et al.
(late Miocene, Huayquerian SALMA, (9.0–6.8 Ma, Brazil) 2014). Early geological studies of the Castillo Formation
with eight known taxa from five families, Orophodontidae, (Wheeler 1960) suggested that it was deposited in mainly
Megalonychidae, Nothrotheriidae, Mylodontidae, and shallow shore and marine environments that included a shal-
Megatheriidae (Cozzuol 2006; Negri et al. 2010); and low freshwater to brackish facies intertongued with a local
3) the Urumaco sequence (middle Miocene to early continental facies. In general, the age of the Castillo
Pliocene, Huayquerian SALMA, ~12–6 Ma, Venezuela) Formation ranges from late Oligocene in the north to early
with ten taxa represented by two families, Mylodontidae Miocene in the south of the Falcón Basin (Wheeler 1960;
and Megatheriidae (see Linares 2004; Carlini et al. Johnson et al. 2009; Rincón et al. 2014).
2006; Rincón et al. 2015 for details). Since the year 2000, ongoing field work in the Castillo
It has been proposed that some of these sloth assemblages Formation at the Cerro La Cruz locality has resulted in the
still included members of very basal sloth groups and/or the recognition of a diverse vertebrate assemblage (see Rincón
earliest representatives of lineages or clades known only from et al. 2014 and references cited therein) including new taxa
older deposits of the significantly better sampled areas of such as the holotype of the oldest Neotropic gavialoid,
southern South America (Hirschfeld 1985; Carlini et al. Siquisiquesuchus venezuelensis (Brochu and Rincón 2004)
1997, 2006; Rincón et al. 2015), suggesting that these lineages found near Siquisique, Lara State.
survived much later in the more tropical northern portions of The geology, chronology, and paleontology of the locality
South America. A diversity of factors, such as the poorly with the greatest diversity of vertebrates from the Castillo
known phylogenetic affinities of these earlier sloth lineages Formation, Cerro La Cruz, were recently updated by Rincón
and the scarcity of early Neogene fossil localities in northern et al. (2014). The lithology of the Cerro La Cruz sequence
South America have limited our understanding about the di- consists of alternating packages of siliciclastic and carbonate
versity, origin, and evolution of sloths in the Neotropics. sediments, deposited principally in a near shore marine envi-
Whether they have always been present in the Neotropics ronments (subtidal shallow marine, intertidal near man-
and merely continued to survive in this region after groves), with episodes of continental environments of appar-
disappearing from the southern part of the continent, or shifted ently short duration (see Rincón et al. 2014 for details). In
their range northward into the more tropical part of the conti- addition, four isotopic ages obtained from analyses of stron-
nent in response to environmental change in the south is not tium ( 87 Sr/ 86 Sr) ratios suggest that the Cerro La Cruz
J Mammal Evol

sequences were deposited during the early Miocene (17.21– characters were coded as such, as implemented in TNT
19.27 Ma; Burdigalian). They overlap with the older part of (Goloboff et al. 2008). We also provide a second phylogenetic
the Santacrucian SALMA. Cerro La Cruz is located near La analysis based on the postcranial elements (see coding in
Mesa village, north of Carora town, Lara State, near the Appendix 2) of the new Castillo sloth to independently deter-
southern-most exposures of the Sierra de La Baragua, north- mine the affinities of the new taxon, coding its postcranial
western Venezuela (Fig. 1). elements following the data set of Rincón et al. (2015),
consisting of 25 characters of the femur and tibia, across 21
taxa, and using Megalonyx as the outgroup. This permits us to
Material and Methods evaluate the affinities of the new taxa with other Venezuelan
and northern South American sloths that are known mostly
The specimens described here are housed at the Instituto from the femur and tibia.
Venezolano de Investigaciones Científicas (IVIC),
Paleontological collection, in Caracas, Venezuela. All mea- Search Methods Both data sets were analyzed using the TNT
surements are in millimeters and were taken with a digital 1.1 software (Goloboff et al. 2008). All characters were treated
caliper. The comparison of the newly described taxon princi- as non-additive (unordered); gaps were treated as missing.
pally is with the other mylodontoid sloths from northern South The characters were analyzed using equal weights methodol-
America and with the closely related Mylodontoidea. ogy. The heuristic parsimony analysis of 1000 replicates was
performed using the ‘traditional search option.’ The swapping
algorithm used was tree bisection reconnection (TBR), with
Molariform Notation We follow the notation proposed by
ten trees saved per replication, collapsing the trees after each
Carlini and Scillato-Yané (2004), and followed by others au-
search.
thors (McKenna et al. 2006; Shockey and Anaya 2010), in
designating the four lower teeth as cf1 and mf1–3.
Following Gaudin (2004) and Shockey and Anaya (2010), Body Mass To calculate the body mass of the new taxon, we
we avoid the use of the “orophodont” notation, because the used the predictive regression equation based on measure-
dentition of the new taxon described here is unambiguously ments of the femur derived from extant mammals developed
homologous to those of other mylodontids. by Scott (1990).

logmass ¼ 3:4855*logfl–2:9112
Data Sets In order to estimate the phylogenetic affinities of
the new sloth described here, we use the data set of Gaudin where, fl is the femur length.
(2004) consisting of 286 osteological characteristics of the
skull, lower jaw, dentition, and hyoid arch, and the 34 sloth Institutional Abbreviations IVIC-P, Colección de
genera including the new taxon described here (see Appendix Paleontología, Instituto Venezolano de Investigaciones
1 for details); two outgroup taxa were utilized (Glyptodon and Científicas, Caracas, Venezuela. MACN, Museo Argentino
Myrmecophaga). In contrasts with Gaudin (2004), all multi- de Ciencias Naturales Bernardino Rivadavia, Buenos Aires,
state characters were treated as unordered. Polymorphic Argentina. MBLUZ–P, Museo de Biología de la Universidad

Fig. 1 Location and geological setting of Cerro La Cruz, Castillo Modified from Rincón et al. (2014). Abbreviations: Tpem = Matatere
Formation, Lara State, Venezuela. Black star shows the location of the Formation (Eocene), Tolmc = Castillo Formation (early Miocene), Qal
analyzed section. a, Geological map of Cerro La Cruz locality; b, = Alluvial (Quaternary).
Regional; c, Continental map of the location of the described section.
 J Mammal Evol

del Zulia, Sección de Paleontología, Maracaibo, Venezuela. having a piriform last lower molariform (mf3). Differs from
MCC, Museo de Ciencias de Caracas, Venezuela. MNHN, Paroctodontotherium, Octodontotherium, and Orophodon in
Muséum national d’Histoire naturelle, Paris, France. UATF, having molariforms with a thin layer of cementum. Differs
Museo de la Universidad Autonoma Tomas Frias, Potosí, from Bolivartherium, Orophodon, and Pseudoprepotherium
Bolivia. UCMP, The University of California Museum of confusum by the presence of three bilobate lower molariforms.
Paleontology Berkeley, California. YPM-PU, Princeton Differs from Brievabradys by its larger size, the presence of
University Collection in the Peabody Museum of Natural three bilobate lower molariforms, and the parallel lower
History, Yale University, New Haven, Connecticut. toothrow. Differs from Octodontotherium by the absence of
a marked asymmetric step between the anterior and posterior
Others Abbreviations SALMA, South American Land lobes in the last lower molariform. Differs from
Mammals Age; Ma, millions of years; Mf and mf, upper and Octodontobradys by molariforms more quadrangular (in oc-
lower molariform tooth; Cf, upper caniniform. clusal view) and mf3 piriform. Differs from Pseudoglyptodon
spp. by the presence of bilobed (not trilobed) lower
molariforms.
Systematic Paleontology
Derivation of Name Takumara, from Takumará an Ayamán
Xenarthra Cope, 1889 word (a now extinct aboriginal language from north central
Folivora Delsuc, Catzeflis, Stanhope, Douzery 2001 Lara and Falcón State, Venezuela), which means sloth
Mylodontoidea Gill, 1872 (Oramas 1916).
Mylodontidae Gill, 1872
Holotype IVIC-P-1828; right mandible with mf1-mf3 series,
Baraguatherium, gen. nov. but lacking the first tooth, the coronoid and angular processes,
and the anterior part of the mandible; and associated second
Diagnosis. As for the type and only species. upper molariform (Figs. 2a–c, 3a–b, 4a).
Derivation of name. Baragua from Sierra de La Baragua,
the northern cordillera of Lara State, Venezuela, and therium Paratype IVIC-P-2917; femur, lacking the femoral head and
(Greek for beast). greater trochanter (Fig. 5).
Type Species. Baraguatherium takumara, new genus and
species. Referred Material MBLUZ-P-5043 and MBLUZ-P-5044;
Occurrence. Early Miocene (Burdigalian), ca. earliest both tooth fragments (Sánchez-Villagra et al. 2004)
Santacrucian SALMA, Castillo Formation, northwestern (Fig. 2d–e).
Venezuela, northern South America.

Baraguatherium takumara, sp. nov. Occurrence

Tardigrada gen. et sp. indet; Sánchez-Villagra et al. (2004): Cerro La Cruz, 20 km north of Carora Town, Lara State, north-
107–109, figs. 1c–h. western Venezuela (Fig. 1). All the material described came from
Orophodontidae gen. et sp. indet; Rincón et al. (2014): 516, the bone-bed locality informally called “valle de los vertebrados”
fig. 5d. (vertebrate valley), where most of the continental mammals of
Diagnosis. Baraguatherium differs from other sloths in the Cerro La Cruz have been recovered (see Rincón et al. 2014 for
following character combination: toothrows parallel in occlu- details). This bone-bed comprises the Unit C of Rincón et al.
sal view; long axis of molariforms parallel to the long axis of (2014) with an age constraint between 17.21–18.27 Ma (early
the toothrow; mandibular symphysis extends to the middle of Miocene; Burdigalian; ca. Santacrucian SALMA).
the second lower tooth (mf1); diastema absent; tooth with a
very thin layer of cementum; mf1-mf3 series of similar size
and bilobate, with mf3 conspicuously piriform (anterior lobe Description and Comparisons
much wider than the posterior lobe); occlusal surface of tooth
beveled. Femur diaphysis straight; third trochanter positioned at Mandible The incomplete mandible of Baraguatherium lacks
the middle of the diaphysis and projects posteriorly; the first lower tooth (cf1), the coronoid and angular processess,
ectepicondyle and entepicondyle robust and project prominently. and the anterior part of the mandible. The horizontal ramus of the
Baraguatherium differs from members of Megatheroidea mandible is relatively broad and massive as in most
and several Miocene mylodontoids (except Brievabradys, plesiomorphic sloths (e.g., Orophodon, Octodontotherium, and
Orophodon, Octodontotherium, and Octodontobradys) in Pseudoglyptodon; Pujos and De Iuliis 2007). Baraguatherium
J Mammal Evol

Fig. 2 Holotype of Baraguatherium takumara, gen. et sp. nov. Mandible (IVIC-P-1828) in lateral a, lingual b, and c occlusal views. d second upper
molariform in occlusal view (IVIC-P-1828); e possible upper caniniform (MBLUZ-P-5044)

as in many Mylodontidae has a horizontal toothrow in lateral Catonyx, and unlike the rest of Mylodontidae where the toothrow
view (Fig. 3), while in occlusal view (Fig. 4), the lower toothrows is anteriorly divergent. Baraguatherium is similar to other “bas-
appear to be parallel, as in Octomylodon, Scelidotherium, and al” mylodontid sloths (e.g., Paroctodontotherium, Brievabradys,

Fig. 3 Comparison among late Oligocene (Deseadan SALMA) to haploides mandible in lateral view (MNHN-des-277); d, Brievabradys
middle Miocene (Laventan SALMA) mylodontid sloths. a–b. laventensis (holotype) mandible in lateral view (LV-4-12); e,
Baraguatherium takumara, gen. et sp. nov. (holotype) mandible (IVIC- Octodontotherium grande mandible in lateral view (MNHN-des-246);
P-1828); a, in lateral view, and b, in lingual view; c, Orophodon f, Octodontotherium grande mandible in lateral view (MACN-12,777)
 J Mammal Evol

Fig. 4 Occlusal view of some late Oligocene (Deseadan SALMA) to Octodontotherium grande mandible (MACN-12,777); d, Orophodon
middle Miocene (Laventan SALMA) mylodontid sloths. a, haploides (holotype) mandible (MNHN-des-277); e, Brievabradys
Baraguatherium takumara (holotype) mandible (IVIC-P-1828); b, laventensis (holotype) mandible (LV-4-12)
Paroctodontotherium calleorum palate (holotype UATF-V-000,127); c,

Octomylodon, Octodontotherium, Pseudoprepotherium posteriorly, reaching the third lower molariform (the mf2). The
confusum) in that the long axis of the molariforms is parallel or mandibular symphysis appears to be distinctively narrow
orthogonal to the long axis of the toothrow (Fig. 4). In other (Table 1).
related sloths, as in Octodontobradys, the toothrows are slightly In Baraguatherium the inferior edge of the mandible is
convergent anteriorly. The mandibular symphysis of straight and horizontal; and the ascending ramus of the man-
Baraguatherium is parallel to the toothrow, and extends posterior dible (in lateral view) extends in front of the posterior
to the first lower molariform, reaching the middle portion of the molariform, similar to Octodontobradys, Brievabradys,
second lower tooth (the mf1; Fig. 2c), unlike Brievabradys, Octomylodon, Nematherium, Scelidotherium, Catonyx, and
Octomylodon, and Octodontotherium, where the symphysis ends Thinobadistes, while, in Octodontotherium,
anterior to first lower tooth (Gaudin 2004) and differs from Pseudoprepotherium, Pleurolestodon, and Mylodon, the as-
Octodontobradys in which the symphysis extends more cending ramus extends only partially in front of the posterior

Fig. 5 Baraguatherium takumara femur (paratype IVIC-P-2917); a, posterior view; b, anterior view; c, medial view; d, distal view
J Mammal Evol

Table 1 Measurements of Baraguatherium takumara (IVIC-P-1828) of Octodontotherium shows a higher mesial than distal facet,
from the Castillo Formation, Venezuela in mm. Tooth lengths are mesial-
which results in a step-like facet. A similar condition occurs,
distal dimensions and widths are transverse dimensions
although to a lesser degree, in Octomylodon, Orophodon,
Element Length (mm) Width (mm) Paroctodontotherium, and Brievabradys (Gaudin 2004; Pujos
and De Iuliis 2007).
Toothrow length (mf1-mf3) 68.0 -
The configuration of the lower dentition of Baraguatherium
Total length of mandible >123.0 -
resembles that of Octodontotherium grande and
Maximum mandible width - 35.6
Octodontobradys paraensis (Santos et al. 1993), in which there
Mandibular symphysis - 20
are least three bilobate lower molariforms (the posterior three
Dentitions
teeth). The enigmatic Octomylodon also exhibits three lower
mf1 20.4 15.8 bilobate molariforms (Scillato-Yané 1977), but the dental formu-
mf2 21.2 16.8 la of this genus only includes three lower molariforms, thus
mf3 21.0 16.7 differing from the four lower teeth present in Baraguatherium
Mf2 24.0 19.2 and all other mylodontids. While the molariforms are similar to
one another in that all are distinctly bilobate, there are slight
differences in size and shape between each consecutive tooth in
molariform (Gaudin 2004). Baraguatherium has the horizon- the series. The mf3 is bilobate and piriform in shape (the anterior
tal ramus blending uniformly into the ascending ramus, and lobe is significantly wider than the posterior lobe). This condition
lacks an ascending ramus with internal ridge running oblique- is similar to Octodontotherium, Octodontobradys, Brievabradys,
ly vertically from ventral edge near the base of the angle, Orophodon, Pseudoprepotherium confusum, Glossotherium,
toward the last tooth. Baraguatherium also lacks the medial Lestodon, and Mylodon (mf3 as bilobate molariform, with an
ridge running along the anterior edge of the coronoid process antero-posterior elongation). All mylodontids (except
and the medial fossa on the angular process. Urumacotherium) have a bilobate last lower molariform, which
is considered as possibly symplesiomorphic for mylodontids
Dentition The arrangement of the teeth of Baraguatherium is sloths (see Rinderknecht et al. 2010). In Baraguatherium the
very distinctive. Although the mandible is broken in front of mf2 is the longest molariform of the mandible (Fig. 2c), although
the first molariform, the posterior part of the caniniform (cf1) all lower molariforms are closely similar in size (Table 1). The
alveolus is present, and it is very close of the second molariform mf2 of Baraguatherium is bilobate in shape (in cross-section),
(Fig. 2c). Therefore, we infer that in Baraguatherium a diastema while this tooth is elongate and irregularly lobate in mylodontids
is absent between the caniniform and the first molariform alveoli. (e.g., Lestodon, Thinobadistes, Glossotherium, and
At least the lower teeth are molariform in shape, similar to those Pleurolestodon), or S-shaped in scelidotheres (e.g.,
of Octodontobradys. Nematherium, Scelidotherium, and Catonyx). The mf1 of
In transverse section, the upper and lower molariforms of Baraguatherium is also bilobate in shape (in cross-section), sim-
Baraguatherium shows three distinctive tooth tissues ilar to Octomylodon, Octodontotherium, and Octodontobradys.
(following Green and Kalthoff 2015). The core of the teeth is In other mylodontids the mf1 is anteroposteriorly ovate in shape
represented by a distinctive and relatively thin layer of (e.g., Pseudoprepotherium confusum, Brievabradys, Lestodon,
vasodentine, surrounded by a thick layer of orthodentine Thinobadistes, Mylodon), irregularly lobate (e.g., Nematherium,
(representing the thickest dentine layer), followed by a thin layer Glossotherium, Pleurolestodon, Paramylodon), or irregularly
of cementum around the outside of the molariform (Fig. 2c). lobate and elongate mesiolabially to distolingually, compressed
This pattern is similar to that observed in Brievabradys and perpendicular to the long axis (e.g., Scelidotherium and
Octodontobradys, while in the rest of the known sloths the ce- Catonyx). With respect to the upper molars of Baraguatherium,
mentum layer is generally thicker (e.g., Paroctodontotherium, the only recovered upper molariform (the Mf2) (Fig. 2d),
Pseudoprepotherium confusum; Gaudin 2004; Shockey and which was found in association with the mandible, is also
Anaya 2010). In Baraguatherium the outer portion of the bilobate in shape (in cross-section), as in Paroctodontotherium,
orthodentine is corrugated. This feature is apparently common Octomylodon, and Octodontotherium. In other taxa, the Mf2
in advanced or later mylodontids (e.g., Mylodon, Paramylodon), is either ovate anteroposteriorly (e.g., Brievabradys and
but has not been previously observed, until now, in the late Pseudoprepotherium confusum ), or lobate (e.g.,
Oligocene to early Miocene mylodontids, such as Scelidotherium, Catonyx, Nematherium, Lestodon, and
Octodontotherium and Nematherium (Green and Kalthoff 2015). Thinobadistes; Gaudin 2004). Finally, it should be noted that
The molariforms of Baraguatherium are slightly longer than the bilobation of both the lower and upper molariforms is sym-
wide, while in other related taxa, such as Octodontobradys metrical and the labial and lingual lobes are similar in depth.
puruensis, the molariform are much more elongate mesially- Though the available material of Baraguatherium is not
distally. The occlusal surface (best observed in the lateral view) directly comparable to the holotype of Paroctodontotherium
 J Mammal Evol

(which is only known from a partial skull), as previously men- the diaphysis is cylindrical to oval in cross-section). The third
tioned, the latter genus has a thick layer of cementum on the trochanter projects posteriorly, and is positioned at the middle
upper teeth (see Fig. 1a of Shockey and Anaya 2010), in of the diaphysis, as occurs in Pseudoprepotherium confusum
contrast to Baraguatherium where the cementum is signifi- and Bolivartherium urumaquensis. The ectepicondyle and
cantly thinner (Fig. 2c). Both genera differ in size as in the entepicondyle are robust and project prominently laterally
holotype of Paroctodontotherium the anteroposterior length and medially, respectively. The patellar and condylar surfaces
of the Mf1-Mf3 is around 55 mm, whereas in are connected. The patellar surface is shorter than wide. The
Baraguatherium, the similar measurement (the mf1-mf3 se- medial condyle is larger than the lateral (Rincón et al. 2015).
ries), is larger (68 mm length), although the interdentary space The above described features of the femur show that
between the upper teeth of Paroctodontotherium is larger than Baraguatherium differs from the other mylodontoids known
the same space in the lower dentition of Baraguatherium. The from Miocene of northern South America: Venezuela (e.g.,
upper Mf2 of Paroctodontotherium is 13.4 mm length (mm) Eionaletherium, Mirandabradys spp. and Bolivartherium,
and 11.9 mm width, while the same teeth in Baraguatherium Pseudoprepotherium venezuelanum), or Colombia
is 65 % larger (Table 1). In any case, is clear that (Pseudoprepotherium confusum).
Baraguatherium is larger than Paroctodontotherium.
It is possible to differentiate Baraguatherium from the oth-
e r k n o w n Ve n e z u e l a n M i o c e n e m y l o d o n t o i d s , Phylogenetic Affinities Following the procedures outlined in
Mirandabradys and Bolivartherium (Carlini et al. 2006). the Materials and Methods and using osteological features of the
Mirandabradys socorrensis (recovered from the middle skull, lower jaw, dentition, and hyoid from Gaudin (2004), we
Miocene of the Socorro Formation) has laterally expanded recovered 12 most parsimonious trees (MPT) from the TNT
upper mola riforms, a condition not observed in analysis. Each of those trees has a consistency index (CI) of
Baraguatherium. Bolivartherium (known from two species) 0.398 and a retention index (RI) of 0.755. A strict consensus tree
has a diastema between the two most mesial teeth (this condi- of these MPT is illustrated in Fig. 6a. The strict consensus is in
tion is absent in Baraguatherium); the molariform series is general agreement with those of Gaudin (1995, 2004), with
s u b p a r a l l e l t o a n t e r i o r l y d i v e rg e n t ( p a r a l l e l i n Bradypus as a sister group of all others sloths and two well-
Baraguatherium), and the mf1 and mf2 are subtrapezoidal in defined clades among sloths, the Mylodontoidea and the
shape (mf1 and mf2 bilobate in Baraguatherium), with the Megatherioidea (see Gaudin 2004 for details). Baraguatherium
mf3 typically bilobate (see Carlini et al. 2006 for details). is placed together with Octomylodon in an unresolved basal
Previously, two isolated sloth molariforms were recovered position among the Mylodontoidea, and as a sister group to all
from the same stratigraphic level as the holotype of remaining mylodontoids, including Nematherium (Fig. 6a).
Baraguatherium (MBLUZ-P-5044; MBLUZ-P-5043; However, this position must be considered quite tentative. The
Sánchez-Villagra et al. 2004), and their similar size, the pres- limited material thus far discovered precludes a definitive polar-
ence of the three distinctive tooth tissues, and the close geo- ization of some characters considered by Gaudin (2004) as di-
graphic and stratigraphic provenance of these specimens agnostic for the Mylodontidae, such as a dorsoventrally short,
(same bone-bed, and only around 50 m east) suggest that they anteroposteriorly broad mandibular coronoid process, and an
should also be referred to Baraguatherium. The importance of anteroposteriorly short, dorsoventrally deep angular process of
these specimens, especially for MBLUZ-P-5044, is that it the mandible.
probably represents a basal fragment of an upper caniniform, A second line of evidence for the phylogenetic affinities of
based on its ovate cross-section and the ovate shape of the core Baraguatherium is based on the femur (Fig. 6b). In this sep-
of the tooth, with a thin layer of vasodentine and the presence arate analysis we recovered six most parsimonious trees
of a distinctive thin outer layer of cementum (see Sánchez- (MPT) from the TNT analysis. These trees have a consistency
Villagra et al. 2004: fig. 1g–h). index (CI) of 0.373 and a retention index (RI) of 0.631. The
topology of the consensus tree resembles that described in
Femur The femur of Baraguatherium (Fig. 5) is distinctively Rincón et al. (2015)), where two main clades were recovered,
flat and wide. The distal epiphysis is completely ossified to the Mylodontoidea and Megatheroidea. Baraguatherium falls in-
diaphysis, which means that is an adult individual. Although to the Mylodontoidea, in an unresolved clade together with
the specimen (IVIC-P-2917) lacks the head of the femur and two other northern South America taxa, Pseudoprepotherium
the lesser trochanter, it shows some similarities to the other confusum (middle Miocene, Colombia) and Bolivartherium
Miocene mylodontoids from the Neotropics (Rincón et al. urumaquensis (late Miocene, Venezuela). These three taxa
2015). The shaft of the femur is straight, similar to share two features, the third trochanter projects posteriorly
Urumacotherium. The femur diaphysis is anteroposteriorly from the diaphysis of the femur relative to the lateral margin
flattened, as occurs in most mylodontoids (except in of the greater trochanter and third trochanter positioned at the
Bolivartherium urumaquensis and Urumacotherium, in which middle of the diaphysis.
J Mammal Evol

Fig. 6 Results of phylogenetic analyses. a, strict consensus cladogram of of for the femur and tibia. The dark pentagon represents the node of the
the 12 most parsimonious trees (CI = 0.398; RI = 0.755) based mainly on Mylodontoidea/Mylodontidae, while the white circle represents the
the data in Gaudin (2004) for osteological characteristics of the skull, Megatheroidea clade. The new taxon described in this paper is in bold
lower jaw, dentition and hyoid; b, strict consensus cladogram of six type. Note in both cladograms the location of the early Miocene new sloth
most parsimonious trees (CI = 0.373; RI = 0.631), based on the data set Baraguatherium takumara, gen. et sp. nov., among the Mylodontoidea

Discussion molariforms composed of a greater percentage of orthodentine

and lacking the internal channel of vasodentine. Although the
With a well constrained age between 17.21 and 18.27 Ma lower mf3 of Baraguatherium is bilobate, the presence of this
(Burdigalian, early Miocene; Rincón et al. 2014), B. takumara vasodentine layer in the core of the tooth precludes assigning it
constitutes the earliest unequivocal sloth so far known in north- to the Mylodontidae based on the criterion of Rinderknecht et al.
ern South America. The Santacrucian SALMA age is usually (2010). In contrast, this condition is considered diagnostic of the
considered to be between 16.3–17.5 Ma (Flynn and Swisher family Orophodontidae (Hoffstetter 1956). Hoffstetter (1982)
1995) or 16–17.8 Ma (Marshall et al. 1986). In contrast, Croft considered the Orophodontidae to be an early tardigrade side
et al. (2004, 2007) reported an age estimate of 17.0–19 Ma for branch that apparently disappeared during the Oligo-Miocene
the Santacrucian SALMA based on a locality from northern transition. Hoffstetter (1982) characterized the family as having
Chile. If we accept the proposal of Croft et al. (2004, 2007), this dasypodid teeth (small cap of cementum surrounding a thick cap
would extend the length of the Santacrucian SALMA to between of osteodentine and a reduced central nodule of vasodentine) and
16 and 19 Ma. The latter dates indicate that the Cerro La Cruz considered them convergent with the later tardigrades in the
outcrops and the bone-bed where the specimens of B. takumara post-cranial skeleton.
were recovered at least partially overlap the Santacrucian The affinities of the Orophodontidae to other sloths have
SALMA. not yet been determined, mostly due to the small number of
Specimens MBLUZ-P-5044 and MBLUZ-P-5043 (here re- incomplete specimens. Various authors have placed the
ferred to Baraguatherium) were previously considered to be Orophodontidae as basal mylodontoids (e.g., Engelmann
closely related to the Santacrucian megathere, Planops magnus, 1985; Villarroel 2000; Gaudin 2004; Shockey and Anaya
and the later northern Neotropical Mylodontidae, 2010), within the Megatherioidea (Pujos and De Iuliis
Urumacotherium garciai (late Miocene, Urumaco Formation, 2007), or even outside of the Phyllophaga sensu stricto
Venezuela), Brievabradys laventensis or Pseudoprepotherium (Hoffstetter 1954, 1969). But the morphological affinities of
confusum (both from the middle Miocene, La Venta, Baraguatherium to Paroctodontotherium, which was includ-
Colombia; Sánchez-Villagra et al. 2004). However, in light of ed into Mylodontidae (Shockey and Anaya 2010), allow us to
the more diagnostic material recovered from Cerro La Cruz, consider the uncertain Orophodontidae clade as basal
these relationships must be reassessed. Mylodontoidea. Usually five genera were included in the
Rinderknecht et al. (2010) characterized members of the Orophodontidae clade, Orophodon, Octodontotherium,
Mylodontidae as having a bilobate last upper and lower Octodontobradys, and Brievabradys (McKenna and Bell
 J Mammal Evol

1997). However, in the most recent revision of the affinities of washboard-like (corrugated) structure on the outer portion of
this clade, the status of Octodontobradys and Brievabradys is the orthodentine is phylogenetically understood as being a de-
considered uncertain (Pujos and de Iuliis 2007), restricting the rived feature for stratigraphically younger taxa within both the
‘orophodontids’ to those genera recovered from the Deseadan Scelidotheriidae and Mylodontidae (Green and Kalthoff 2015).
SALMA of central and southern South America, The phylogenetic hypotheses based on the osteological fea-
Octodontotherium and Orophodon. Baraguatherium is clearly tures of the skull, lower jaw, dentition, and hyoid (Fig. 6a;
distinguished from those Deseadan basal Mylodontoidea following Gaudin 2004) and the femur and tibia features
(Octodontotherium and Orophodon) by its bilobate molariform (Fig. 6b following Rincón et al. 2015) indicate that
teeth (mf1-mf3) with the occlusal surfaces almost flat, distal Baraguatherium seems to be a basal member of Mylodontidae
molariforms with symmetrical lobes, and the teeth arranged or at least of the Mylodontoidea. Based on the criteria of Pujos
along a rectilinear anteroposterior axis (see Pujos and De Iuliis and De Iuliis (2007), Baraguatherium would not be allocated to
2007 for details). the ‘Orophodontidae.’ However, the presence of
The dentition of Baraguatherium more closely resembles the Baraguatherium in the early Miocene of northern South
dentition of the later taxon Octodontobradys (late Miocene – America represents a distinct lineage from all of the other known
Pliocene, Solimões Formation, Brazil) in having the three lower sloths from the Neotropics.
molariform conspicuously bilobate. According to Santos et al. During the early Miocene there are only two known
(1993) and Guilherme et al. (2011), Octodontobradys belongs to mylodontoids in southern South America, the Santacrucian gen-
the subfamily Octodontobradyinae, while Pujos and De Iuliis era, Nematherium and Analcitherium. Mones (1986) listed up to
(2007) considered its dentition to be more reminiscent of the six species for Nematherium and one for Analcitherium. In con-
Scelidotheriinae, possibly representing a relict primitive trast, with the discovery of Baraguatherium plus the tentative
scelidothere that conserved a multi-lobate dentition. However, record of Pseudoprepotherium venezuelanun from the ‘early’
many characteristics of Baraguatherium (e.g., broader Miocene of Río Yuca Formation (Venezuela), there probably
molariforms, anteriorposteriorly shorter mandibular symphysis) exist at least two mylodontoids of this age in northern South
also differentiate it from Octodontobradys. Finally, the America.

Table 2 Estimated body mass of selected mylodontoid sloths calculated from total femur length (in cm), using the equation of Scott (1990). Note that
giant mylodontids first appear in the middle Miocene. (Table modified from Rincón et al. 2015)

Taxa Femur length (cm) Body mass (kg) Biochronological age

Lestodon armatus 73 3825 Pleistocene

Glossotherium myloides 49 953 Pleistocene
Glossotherium myloides 46 765 Pleistocene
Glossotherium wegneri 47 824 Pleistocene
Paramylodon harlani 54.6 1390 Pleistocene
Scelidotherium leptocephalum 45 708 Pleistocene
Thinobadistes segnis 44.3 671 Late Miocene-Pliocene-Pleistocene
Mirandabradys zebasi 57.1 1625 Early Pliocene
Mirandabradys zebasi 53.5 1295 Early Pliocene
Mirandabradys urumaquensis 59.4 1864 Late Miocene
Urumacotherium garciai 61.5 2104 Late Miocene
Eionaletherium tanycnemius 51 1096 Late Miocene
Bolivartherium urumaquensis 58 1716 Late Miocene
Simomylodon uccasamamensis 32.5 228 Late Miocene
Mirandabradys socorrensis 56 1518 Middle Miocene
Pseudoprepotherium confusum 47 824 Middle Miocene
Pseudoprepotherium confusum 48 887 Middle Miocene
Pseudoprepotherium confusum 48.4 913 Middle Miocene
Pseudoprepotherium venezuelanun 41.9 552 Early Miocene
Baraguatherium takumara 40.6 495 Early Miocene
Baraguatherium takumara 46 765 Early Miocene
Chubutherium ferelloi 38.3 404 Late Oligocene-Early Miocene
J Mammal Evol

Paleoecological Affinities Although broken, the femur of (Collins 1934), represents the unequivocally earliest
Baraguatherium preserves 60–70 % of the total femur length. mylodontoid sloth in this part of the subcontinent. Its discov-
The length of the Castillo specimen is 345 mm and probably ery also offers a new view of the early sloth origins,
represents 75–85 % of the total length of the femur, suggesting evolution, and adaptations in the Neotropics in contrast to
an estimated complete length around to 406–460 mm. While the traditional focus on the evolution of the group based on
at the larger end of the size range for sloths (see electronic records from the southern part of the continent, which took
supplementary material for details) with an estimated body place in more temperate environments. Our phylogenetic
mass over the 500 kg (ranging probably between 495 and hypothesis based independently on a partial mandible and a
765 kg), Baraguatherium is in the same size range as the other partial femur is in agreement with the Gaudin (2004) hypoth-
northern South America mylodontid, Pseudoprepotherium esis that there are two well-defined Folivora clades:
venezuelanum, and is much smaller than the later mylodontids Mylodontoidea and Megatherioidea, and Baraguatherium
from the Urumaco sequence (middle Miocene to early Pliocene; phylogenetically and morphologically belongs to a
Venezuela), Mirandabradys spp., Urumacotherium, and unique lineage within the basal Mylodontoidea present
Bolivartherium (see Table 2). In addition, Baraguatherium ap- in northern South America. That Baraguatherium
pears to be significantly larger compared to the other early inhabited lowland coastal tropical forest indicates a hab-
Miocene (Santacrucian) mylodontids, Analcitherium and itat preference quite different from the traditional view
Nematherium, which have body masses in the order of 100 kg that mylodontoids are only closely associated with open
(Bargo et al. 2012; Toledo et al. 2014). In general, Santacrucian grassland habitat and that during the early taxonomic
sloths range in body mass from 40 to 100 kg (Bargo et al. 2012), diversification of the clade there was ecological diversi-
and are significantly smaller than the larger late Miocene ty of the group as well.
or Pleistocene sloths. Finally, some of the Desedean
Mylodontoidea, such as Octodontotherium and
Paroctodontotherium, with a body mass of 270 and 47 kg, re-
spectively (Shockey and Anaya 2010), are also smaller than Acknowledgments We wish to thank the Instituto del Patrimonio
Baraguatherium. Cultural (IPC), Venezuela, for fossil collection permissions to this project.
Baraguatherium was most likely to have been a robust, We thank Sr. P. Gómez†, Ing. M.E. Mendoza, Lic. M. López, and Dr. F.
Urbani for their help and camaraderie during the field trips. The major
medium-sized mylodontoid as its femur exhibits strong simi-
funding support of this work was provided by Centro de Ecología,
larities to the later mylodontoids from the Urumaco sequence. Instituto Venezolano de Investigaciones Científicas (IVIC) grant 1096
In this sense, the locomotor system of Baraguatherium like to A.D.R. Also, we thank to B. Shockey and an anonymous reviewer
these other sloths is indicative of terrestrial quadrupedal for all the improvements to that manuscript.
habits. Baraguatherium was found in one of the two continen-
tal episodes within the mainly marine environments of the
Castillo Formation. This suggests that Baraguatherium, like
Appendices
Eionaletherium tanycnemius from the late Miocene of
Urumaco (Rincón et al. 2015), inhabited a low line coastal
Appendix 1
habitat. Also present in the level where Baraguatherium was
found, there is abundant wood and leaves associated with
Character coding for the new early Miocene sloth,
mangrove habitat along with marine invertebrates, intermixed
Baraguatherium takumara following the character descrip-
with fresh water vertebrates such as the turtle Chelus and the
tions of Gaudin (2004).
river fishes Colossoma and Mylossoma (Rincón et al. 2014).
Baraguatherium 12001010202??0100?0- - - - 0- -
These associated taxa provide additional evidence that allows
???1?566?32100??00????????????????1?1???0??????100???
to us to interpret the preferred habitat of Baraguatherium as a
??????????????????????????????????????????????????????????
coastal tropical forest transversed by small rivers (Rincón
??????????????????????????????????????????????????????????
et al. 2014).
??????????????????????????????????????????????????????????
?????????????????????????????????
Conclusions
Appendix 2
The discovery of B. takumara in the early Miocene of
Venezuela increases the diversity of taxa recognized in the Character coding for the new early Miocene sloth,
northern part of South America and, with the possible excep- Baraguatherium takumara following the character descrip-
t i o n o f t h e p o o r l y k n o w n P s e u d o p re p o t h e r i u m tions of Rincón et al. (2015).
venezuelanum whose age assignment needs to be confirmed Baraguatherium 0- - 10- - 00- - - - 10-1- - - - - - - -
 J Mammal Evol

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