Late Pleistocene Pachyarmatherium of Venezuela
7
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
DOI: 10.5281/zenodo.7951590 / Anartia, 35 (diciembre 2022): 7-13
New record of Pachyarmatherium (Cingulata:
Pachyarmatheriidae) from the Late Pleistocene in Venezuela
Nuevo registro de Pachyarmatherium (Cingulata: Pachyarmatheriidae)
para el Pleistoceno tardío de Venezuela
Andrés E. Reyes-Céspedes
1
, Alfredo A. Carlini
2
& Jorge D. Carrillo-Briceño
3
1
Programa de Magister en Paleontología, Universidad Austral de Chile, Valdivia, Chile.
2
Laboratorio de Morfología Evolutiva y Desarrollo (Morphos), Facultad de Ciencias Naturales y Museo, UNLP, CONICET
(Consejo Nacional de Inestigaciones Cientícas y Técnicas), Buenos Aires, Argentina.
3
Universität Zürich, Paläontologisches Institut und Museum, Zürich, Switzerland.
Correspondence: jorge.carrillo@pim.uzh.ch
(Received: 08-09-2022 / Accepted: 15-12-2022 / On line: 19-05-2023)
ABSTRACT
e Falcón state, in northwestern Venezuela, preserves one of the oldest localities documenting human presence on the
Americas in association with megafauna remains. One of these localities is the Late Pleistocene Taima-Taima site, which
is located in the vicinity of the Caribbean coast. Excavations since the 1960s in the Taima-Taima site, and surroundings,
have oered new insights into the faunal assemblages that inhabited this arid coastal savannah region during the Late Pleis-
tocene. Isolated osteoderms collected in recent prospections allow us to report here the presence of the extinct cingulate
Pachyarmatherium cf. brasiliense and Pachyarmatherium sp. from the Taima-Taima site, and the new locality Cucuruchú
(Las Dunas). e record of these taxa in the Late Pleistocene of the Falcón state increases the known paleodiversity of
Cingulata for the region and expands the geographical distribution of the genus, which is poorly known in South America.
Keywords: armadillos, Cenozoic, fossils, northern South America, Taima-Taima, Xenarthra.
RESUMEN
El estado Falcón, en el noroeste de Venezuela, conserva una de las localidades más antiguas que documentan la presencia
humana en las Américas en asociación con restos de megafauna. Una de estas localidades es el sitio Taima-Taima del Pleisto-
ceno Tardío, que se encuentra en las inmediaciones de la costa caribeña. Las excavaciones llevadas a cabo desde la década de
1960 en este sitio y sus alrededores han ofrecido nuevos datos sobre los conjuntos faunísticos que habitaban esta región de
sabana costera durante el Pleistoceno Tardío. Osteodermos aislados recolectados en prospecciones recientes nos permiten
reportar aquí la presencia del extinto cingulado Pachyarmatherium cf. brasiliense y Pachyarmatherium sp. en el sitio Taima-
Taima, y la nueva localidad de Cucuruchú (Las Dunas). El registro de estos taxones en el Pleistoceno Tardío del estado
Falcón incrementa la paleodiversidad conocida de Cingulata para la región y amplía la distribución geográca del género,
cuyos reportes son escasos en Sudamérica.
Palabras clave: armadillos, Cenozoico, fósiles, norte de América del Sur, Taima-Taima, Xenarthra.
INTRODUCTION
Pachyarmatherium was originally erected by Down-
ing & White (1995) when describing the type species
Pachyarmatherium leiseyi on the basis of numerous osteo-
derms, cranial and postcranial remains from the Pliocene–
Pleistocene in the USA. In South America, the fossil re-
cord of Pachyarmatherium is restricted mainly to isolated
A. E. Reyes-Céspedes, A. A. Carlini & J. D. Carrillo-Briceño
8
osteoderms and postcranial remains reported from Brazil,
Peru and Venezuela (Rincón & White 2007, Porpino et
al. 2009, Martínez & Rincón 2010). Rincón & White
(2007, gs. 2–3) described the species Pachyarmatherium
tenebris, based on few isolated osteoderms recovered from
Late Pleistocene cave sediments in the eastern Falcón state,
western Venezuela (Rincón & White 2007, g. 1). Porpi-
no et al. (2009), based on numerous osteoderms and post-
cranial remains, described the species Pachyarmatherium
brasiliense, for the late Pleistocene–Holocene in the Rio
Grande do Norte region, northeastern Brazil. Isolated os-
teoderms assigned to P. tenebris from the late Pleistocene
of Peru by Martínez & Rincón (2010) have been interpret-
ed as morphologically indistinguishable from those of P.
brasiliense, leading these authors to consider both species
as synonymous.
Late Pleistocene deposits in the Falcón state preserve
one of the best-recorded paleodiversity records of Cin-
gulata from that time in Venezuela. Such nds are known
from archaeological and paleontological excavations, in-
cluding some specimens collected on the surface in karst
systems (see Carrillo-Briceño 2015). Apart from Pachyar-
matherium, the fossil record of the Falcón state also in-
cludes glyptodonts with the genera Glyptodon (e.g., Royo
y Gómez 1960, Rincón & White 2007, Chávez-Aponte
et al. 2008a) and Glyptotherium (Carlini et al. 2008, 2022,
Carlini & Zurita 2010), the pampatheres Holmesina and
Pampatherium (Rincón 2004, Aguilera 2006, Chávez-
Aponte et al. 2008a, Carrillo-Briceño 2015), and the dasy-
podid Propraopus (Royo y Gómez 1960, Rincón & White
2007). In this contribution, we describe new fossil remains
assignable to Pachyarmatherium cf. brasiliense, which were
collected in the archaeological/paleontological site of
Taima-Taima and its surroundings in the Venezuelan state
of Falcón. e Taima-Taima site is located in the vicinity
of the Caribbean coast (Fig. 1), a region internationally
known to contain some of the oldest localities document-
ing human presence on the American continent (Bryan et
al. 1978, Carlini et al. 2008, 2022, and references therein).
Possible evidence of hunting on glyptodonts was reported
recently for the Taima-Taima and the Muaco (also in the
Falcón state) sites by Carlini et al. (2022). e presence
of Pachyarmatherium cf. brasiliense in the Late Pleistocene
of the Falcón state increases the known paleobiodiversity
of Cingulata for the region and expands the geographic
distribution of the species.
MATERIAL AND METHODS
Four isolated osteoderms, one (MTT-V-320) collected
from the surface washed sediments adjacent to the old ex-
cavations in the Taima-Taima site (11° 29’ 57’’ N / 69° 31’
18’’ W), and the other three (MTT-V-212, -358, -479)
from the surroundings of the Taima-Taima park, a local-
ity referred here as Cucuruchú (Las Dunas) site (11° 30’
10’’ N / 69° 30’ 17’’ W). e later site is located on the
coastal area, near the mouth of the Cucuruchú Creek, ap-
proximately two kilometers east of the Taima-Taima site
(Fig. 1). To avoid the total loss of the specimens, due to the
erosion of the outcrops, these were collected by one of the
authors (AERC) during a paleontological rescue that took
place in 2015 as part of a workshop on protection of pale-
ontological heritage (Zavala & Reyes 2017). Nowadays on
the Taima-Taima excavation site there is an in situ museum
and a park (Fig. 1A–B) that was opened in the year 2000
(see Aguilera 2006, Carrillo-Briceño 2015). e studied
remains were compared with published bibliography, and
measurements were taken using a digital caliper. Com-
parative measurements presented in Table 1 were taken
from Downing & White (1995; for P. leiseyi), Rincón &
White (2007; for P. tenebris), and Porpino et al. (2009; for
P. brasiliense). e specimens are housed in the paleonto-
logical collection of the local Museum of Taratara with the
acronym MTT-V- (Museo de Taratara-Vertebrados).
GEOLOGICAL CONTEXT
e study area is located within the Taima-Taima
Park polygonal area (Fig. 1A), which covers an exten-
sion of approximately 14.8 km². e Taima-Taima site is
approximately 18 km NE of the city of Coro, and 3 km
NW of the Taratara town. e area is characterized by a
continuous exposure of folded strata that constitute La
Vela anticline (see Benites-Palomino et al. 2021, and refer-
ences therein). ere are no formally dened Pleistocene
sedimentary units in this area; most Pleistocene deposits
occur on rounded cobble and pebble layers, eroded and
deposited from the underlying Miocene limestone lay-
ers (Cruxent 1970, Bryan & Gruhn 1979, Carlini et al.
2022). Water springs are also common in the area (Bryan
et al. 1978; Aguilera 2006). Abundant remains of Pleis-
tocene fauna have been reported for the site of Taima-
Taima from the successive excavations carried out since
the 1960s, and include turtles (Testudinidae), bats (Phyl-
lostomidae), ground sloths (Megatheriidae, Mylodon-
tidae), glyptodonts (Glyptodontidae), native ungulates
(Macraucheniidae, Toxodontidae), artiodactyls (Cam-
elidae, Tayassuidae, Cervidae), perissodactyls (Equidae),
carnivore (Felidae, Canidae, Ursidae), and proboscideans
(Gomphotheriidae) (Casamiquela 1979; Bocquentin-
Villanueva 1982a; Chávez-Aponte et al. 2008b; Carrillo-
Briceño 2015, Carlini et al. 2022; and references therein).
Late Pleistocene Pachyarmatherium of Venezuela
9
Figure 1. A. Geographical location of the fossiliferous localities and the Taima-Taima Park. B. Taima -Taima in situ open museum.
C.Cucuruchú (Las Dunas) site.
Table 1. Range of osteoderm measurements of the dierent species of Pachyarmatherium from the literature and measu-
rements of the four specimens discovered at Taima-Taima and the Cucuruchú (Las Dunas) sites. Expressed in millimeters.
Species Length Width icknes Reference
P. leiseyi - 8.1 – 22.9 5.5 – 13.3 Downing & White 1995
P. tenebris 17.6–32.1 14.8–28.5 6.6–9.8 Rincón & White 2007
P. brasiliense 10.1–26.7 14.5–28 7–18.4 Porpino et al. 2009
MTT-V-320 18.96 17,91 10.36 is paper
MTT-V-212 19.76 16.95 14.72 is paper
MTT-V-358 19.80 17.46 14.29 is paper
MTT-V-479 15.03 13.48 10.26 is paper
A. E. Reyes-Céspedes, A. A. Carlini & J. D. Carrillo-Briceño
10
Excavations at the Taima-Taima site during the 1960s and
1970s reported lithic artifacts associated with megafaunal
remains (e.g., Bryan et al. 1978). Possible evidence of hunt-
ing on glyptodonts was also reported recently for the Tai-
ma-Taima site by Carlini et al. (2022). e Taima-Taima
assemblage was dated, based on several C14 analyses, to be
between ~ 17,300calybp [calibrated years before the pres-
ent using IntCal20 (Ramsey 2009)] (14,200 ± 300ybp)
and ~ 15,780calybp (12,980 ± 85ybp) (Bryan & Gruhn
1979; Bryan etal. 1978). e osteoderm MTT-V-320 was
collected from washed surface sediments accumulated be-
side the area of the old excavations, which makes the accu-
rate identication of the fossiliferous level from which this
specimen comes, highly dicult. However, we do not rule
out that the specimen comes from one of the fossiliferous
layers reported for the Taima-Taima site (see Carlini et al.
2022, g. 3).
Regarding the Cucuruchú site (Las Dunas), the locality
was discovered on the right margin of the mouth of the
Cucuruchú Creek, and the Pleistocene deposits are over-
laying Miocene rocks (Bocquentin-Villanueva 1982b).
e fossiliferous locality is characterized by a conglomer-
ate of alluvial origin with a layer of sand or paleodunes on
top (Fig. 1C); both layers yield fossil vertebrates. e age
for the Cucuruchú (Las Dunas) site is estimated based on
its faunal composition (under study), which corresponds
to a typical Late Pleistocene (Lujanian) mammal assem-
blage.
SYSTEMATIC PALEONTOLOGY
XENARTHRA Cope, 1889
CINGULATA Illiger, 1811
†PACHYARMATHERIIDAE Fernicola et al., 2018
†Pachyarmatherium Downing & White, 1995
†Pachyarmatherium brasiliense Porpino et al., 2009
†Pachyarmatherium cf. brasiliense.
(Fig. 2A1–A2)
Referred specimens. MTT-V-320 is a xed osteoderm
from a section of the carapace not corresponding to a mov-
able band.
Locality. Taima-Taima site (Fig. 1).
Description. e osteoderm is hexagonal, with a simi-
lar size to P. brasiliense, larger and thicker than P. ten-
ebris (Fig. 2A1–A2, Table 1). e dorsal surface is well
ornamented and almost at as in P. brasiliense (Porpino
et al. 2009, g. 2; Oliveira et al. 2013, g. 2), and dier-
ent from P. leiseyi and P. tenebris where the dorsal surface
is convex (Downing & White 1995, gs. 1–2; Rincón
& White 2007, gs.2–3). MTT-V-320 has four periph-
eral antero-lateral gures slightly higher than the central
one; the two proximal are larger than the lateral gures,
which reduce in size until disappearing on the margin as
a sharp tip pointing backwards; the central gure is larger,
rounded to subrounded, similar to P. brasiliense (Porpino
et al. 2009, g. 2; Oliveira et al. 2013, g. 2), and dierent
from P. leiseyi and P. tenebris (Downing & White 1995,
gs. 1–2; Rincón & White 2007, gs. 2–3) whose shape
is mostly polygonal, covering a little more than half of the
total exposed surface and posteriorly displaced to reach
the posterior border. is gure is surrounded by a main
sulcus that is shallow and wide, a feature that dierenti-
ates it from P. leiseyi and P. tenebris (Downing & White
1995, gs. 1–2; Rincón & White 2007, gs. 2–3). e
radial sulci are short, deep and as wide as the one that lim-
its the central gure; it has a foramen at the interception
between the main and radial sulci in the anterior region,
a common feature in osteoderms of Pachyarmatheriidae
and Glyptatelinae. Numerous small but well-marked fo-
ramina are located on the main sulcus and on the surface
of the central gure (Fig. 2). e later shows six small
and well-marked foramina, aligned longitudinally in two
parallel rows of three each, a feature not reported in any
of the species of Pachyarmatherium described so far. e
internal surface is at, as in P. brasiliense (Oliveira et al.
2013, g.2) and unlike P. leiseyi and P. tenebris where it
is somewhat concave, and has four large and well-marked
vascular foramina.
†Pachyarmatherium sp.
(Fig. 2B1–D2)
Referred specimens. ree isolated xed osteoderms
from the part of the carapace not corresponding to a mov-
able band. (MTT-V-212, -358, -479).
Locality. Cucuruchú (Las Dunas) site (Fig. 1).
Description. Osteoderms are hexagonal to heptagonal
(e.g., MTT-V-212), very thick in the anterior portion and
reducing its thickness to the distal portion, but thicker
than the described osteoderms in P. leiseyi, P. tenebris and
P. brasiliense in the later part (Downing & White 1995,
Rincón & White 2007; Porpino et al. 2009); slightly lon-
ger than wide, including a rough and convex external sur-
face with the central gure forming the highest relief as
in P. tenebris (Rincón & White 2007, gs. 2–3) and less
pronounced than in P. leiseyi (Downing & White 1995,
gs.1–2). ere are four to six peripheral gures, which
are larger on the anterior margin, reducing signicantly
in size distally; the central gure is rounded or polygonal
as in P. leiseyi and P. tenebris (Downing & White 1995,
gs.1–2; Rincón & White 2007, gs. 2–3), dened by
Late Pleistocene Pachyarmatherium of Venezuela
11
a deep and well-marked main sulcus. e central gure
is large, encompassing most of the external surface of the
osteoderm and posteriorly displaced. e radial sulci are
deep, short and well-dened (except in MTT-V-212). e
piliferous foramina are at the intersection between the
main/central sulcus and the peripheral sulci (Fig. 2B1,
C1, D1), being large and well-marked. Specimen MTT-
V-358 has a large foramen on the central gure and a series
of medium to small-sized foramina scattered throughout
this area, giving it a rough and stippled appearance. e
internal surface is at and somewhat concave, as in P. lei-
seyi (Downing & White 1995, g. 1), and exhibits three to
ve vascular foramina.
DISCUSSION
Despite the abundant cingulates remains recovered
from the Taima-Taima and Cucuruchú sites since the rst
excavations in the locality in the early 1960s, these have
been identied exclusively as glyptodontids (Casamiquela
1979, Ochsenius 1980, Bocquentin-Villanueva 1982a, b,
Aguilera 2006). Primarily on the basis of dorsal carapace
osteoderms, these materials have been reported as be-
longing to the genus Glyptodon (Bocquentin-Villanueva
1982a, Aguilera 2006). Nevertheless, detailed studies of
these specimens, including skulls, postcranial skeletons,
dorsal carapaces and caudal rings, from the Taima-Taima
and Cucuruchú sites, as well as other Late Pleistocene sites
in the surroundings (e.g., Muaco and Quebrada Ocando),
indicates that all of them are in fact assignable to Glypto-
therium and morphologically similar to G. cylindricum
(Carlini et al. 2008, 2022). e pampatheres Holmesina
and Pampatherium, as well as the dasypodid Propraopus
have been reported from other fossiliferous localities in
the Falcon state (Rincón 2004, Aguilera 2006, Chávez-
Aponte et al. 2008a, Carrillo-Briceño 2015). e presence
of Pachyarmatherium in the Taima-Taima and Cucuruchú
(Las Dunas) increases the known paleobiodiversity of cin-
gulates for these sites and the geographic distribution of
the genus in the region, whose previous record included
only the presence of P. tenebris in eastern Falcón state
(Rincón & White 2007).
e morphological characteristics of the Taima-Taima
specimen MTT-V-320 i.e., at external surface, rounded
central gure, shallow and wide sulci and at internal sur-
face (Fig. 2A1–A2) are similar to those of P. brasiliense.
However, due to the scarcity of material its specic iden-
tication is tentative until new material is found and the
description can be rened. According to the morphologi-
cal characteristics of the specimens from the Cucuruchú
(Las Dunas) site i.e., thick osteoderms, longer than wide,
rough and convex external surface, rounded to polygonal
central gure, deep and wide sulci, at to concave internal
surface (Fig. 2B1–D2). ese dier from the Taima-Taima
osteoderms, as well as from P. tenebris and P. leiseyi. How-
ever, considering that these osteoderms could belong to a
non-homologous area of the carapace, only new remains
will allow us to have a more precise taxonomic assignation.
e phylogenetic placement of Pachyarmatherium has
been somewhat problematic, occupying dierent taxo-
nomic positions, from Dasypodoidea (Downing & White
1995, Rincón & White 2007, Oliveira et al. 2013), Glypto-
Figure 2. A1-D2. Pachyarmatherium osteoderms. A1-A2. Pachyarmatherium cf. brasiliense (MTT-V-320) from the Taima-Taima site.
B1-D2. Pachyarmatherium sp. (B1-B2: MTT-V-358, C1-C2: MTT-V-212, D1-D2: MTT-V-479) from the Cucuruchú (Las Dunas)
site. Views: dorsal (A1, B1, C1, D1) and lateral (A2, B2, C2, D2). Comparison with images in Downing & White (1995, gs. 1–2),
Rincón & White (2007, gs. 2–3), Porpino et al. (2009, g. 2), and Oliveira et al. (2013, g. 2).
A. E. Reyes-Céspedes, A. A. Carlini & J. D. Carrillo-Briceño
12
dontoidea, Glyptatelinae (McKenna & Bell 1997, Vizcaíno
et al. 2003), Cingulata incertae sedis as a sister group of the
clade formed by pampatheres and glyptodonts (Porpino
et al. 2009), to Dasypodinae Dasypodini (Oliveira et al.
2013). More recently, Fernicola et al. (2018) described
Neoglyptatelus uruguayensis, based on very complete post-
cranial materials from the late Miocene of Uruguay, allow-
ing these authors to propose the family Pachyarmatheriid-
ae. is would include the species Neoglyptatelus originalis
and Neoglyptatelus sincelejanus from the Middle and Late
Miocene of Colombia (Carlini et al. 1997, Villarroel &
Clavijo 2005), and the genus Pachyarmatherium from the
Plio–Pleistocene of North, Central and South America
(Downing & White 1995, Porpino et al. 2009). ey are
characterized mainly by a carapace divided into two parts
(scapular and pelvic shields) without intermediate dorsal
mobile bands, and the central gure of the osteoderms
displaced posteriorly. e absence of complete or well-pre-
served cranial and dental remains has prevented the study
to infer on ecological preferences of this family. Downing
& White (1995) have suggested myrmecophagous feeding
habits for P. leiseyi, similar to those of Dasypus noemcinc-
tus, based on some mandibular characters and the presence
of claws, possibly employed for digging. e paleoenviron-
mental characteristics inferred for the Leiseyi Shell Pits
locality place P. leiseyi in a mixed environment such as a
coastal mangrove bay in an estuary of a major river, with
swampy areas (Rich & Newson 1995, Downing & White
1995). Likewise, Rincón & White (2007) propose that P.
tenebris inhabited a mixed environment with a predomi-
nance of open savannahs with wooded patches. Similar
paleoenvironments characterized by dry coastal savannas
with the presence of wooded patches have been suggest-
ed for the Taima-Taima area during the Late Pleistocene
(Ochsenius & Gruhn 1979, Ochsenius 1980).
CONCLUSION
In the present study, we report the presence of Pachyar-
matherium cf. P. brasiliense and Pachyarmatherium sp.
from the Taima-Taima and the Cucuruchú (Las Dunas)
sites within the area of the Taima-Taima Park, expanding
the geographical distribution of the genus in Venezuela
during the Late Pleistocene. Although both Taima-Taima
and the Cucuruchú (Las Dunas) sites are located closely
and could be also chronologically contemporaries, the
morphological characteristics of the osteoderms from
both sites allow us to dierentiate them. Nevertheless, for
now the scarcity of the material does not allow identifying
them precisely at the species level. eir presence increases
the diversity of fossil mammals in these fossiliferous locali-
ties and demonstrates the great paleontological potential
for future studies in this region.
ACKNOWLEDGEMENTS
We thank the community of Taratara and the Centro
de Investigaciones Arqueológicas, Antropológicas y Pale-
ontológicas (CIAAP) de la Universidad Nacional Experi-
mental Francisco de Miranda (UNEFM) for the valuable
collaboration. Many thanks to Dinorah Cruz, Isabel De
Jesús, and the Instituto del Patrimonio Cultural de Vene-
zuela (IPC) for their support and kindly providing collect-
ing permits. We greatly appreciate comments and sugges-
tions from the editor and three anonymous reviewers. We
are thankful to Kevin Le Verger for his valuable comments
and suggestions on the manuscript.
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