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Paleobiogeography, biostratigraphy and systematics of the Hoplophorini
(Xenarthra, Glyptodontoidea, Hoplophorinae) from the Ensenadan Stage
(early Pleistocene to early-middle Pleistocene)
Alfredo E. Zurita
a
,
*
, Alfredo A. Carlini
b
,
c
, Gustavo J. Scillato-Yane
´
b
a
Centro de Ecologı
´a Aplicada del Litoral (CECOAL-CONICET), y Universidad Nacional del Nordeste, Ruta 5, km. 2,5 (3400), Corrientes, Argentina
b
Departamento Cientı
´fico Paleontologı
´a de Vertebrados, Museo de La Plata, Facultad de Ciencias Naturales y Museo, Paseo del Bosque s/n
, 1900 La Plata, Argentina
c
Pala
¨ontologisches Institut und Museum, Universita
¨tZu
¨rich, Karl Schmid-Straße 4, CH-8006 Zu
¨rich, Switzerland
article info
Article history:
Available online 10 July 2009
abstract
Neosclerocalyptus Paula Couto (¼Hoplophorus ¼Sclerocalyptus) is a Pleistocene genus of Glyptodontidae
Hoplophorini (¼Sclerocalyptini) that includes several (ca. twelve) species, many of which have been
recognized by typological/morphological taxonomic criteria. Four species have been described for the
Ensenadan Stage (early Pleistocene to early-middle Pleistocene) of the Pampean region, Argentina.
However, this study shows only two of them to be valid: Neosclerocalyptus pseudornatus and
Neosclerocalyptus ornatus. An evident synapomorphy of Neosclerocalyptus is the notable pneumatization
and lateral expansion of the fronto-nasal sinuses, which becomes evident in N. pseudornatus (ca. 1.07–
0.98 Ma) and even more so in N. ornatus (ca. 0.98–0.40 Ma). This character, interpreted here as a probable
response to the cold and arid/semiarid Pleistocene climate, is maximally manifested in the taxa from the
middle Pleistocene (Bonaerian Stage) and late Pleistocene (Lujanian Stage). Neosclerocalyptus is very
common in the Pampean region and north-central Argentina, but very scarce or absent in the Argenti-
nian Mesopotamia, Uruguay and southern Brazil, areas that were subject to relatively more humid and
warmer climates during most of the Pleistocene. From a biogeographical perspective, both Ensenadan
species are restricted to the current Pampean region. N. pseudornatus is recorded in the ‘‘Toscas’’ (caliche
duricrusts) of Rı
´o de La Plata (Buenos Aires City and Olivos) and Mar del Plata (Buenos Aires province),
while N. ornatus is recorded in Mar del Plata and San Pedro (Buenos Aires province), and Granadero
Baigorria (Santa Fe province).
Ó2009 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
The Glyptodontidae were one of the most conspicuous groups of
Xenarthra in South America during most of the Cenozoic. From
a taxonomic perspective, this clade experienced significant diver-
sification (ca. 65 genera; see McKenna and Bell, 1997), as well as
extensive temporal distribution (late Eocene–early Holocene;
Hoffstetter, 1958; Paula Couto, 1979; Politis and Gutierrez, 1998;
Carlini and Scillato-Yane
´, 1999; Cione et al., 2003; Zurita et al., 2005).
Even though Glyptodontidae taxonomy is complex, some
consensus exists at present for the recognition of five subfamilies:
a) Glyptatelinae (late Eocene – late Miocene – late Pleistocene?
Scillato-Yane
´, 1977, 1986; Downing and White, 1995; Carlini et al.,
1997, 2004, 2008a; Vizcaı
´no et al., 2003; Bostelmann et al., 2008);
b) Propalaehoplophorinae (late Oligocene–middle Miocene; Scott,
1903–1904; Scillato-Yane
´, 1977, 1986; Bondesio et al., 1980); c)
Glyptodontinae (middle Miocene–early Holocene; Cabrera, 1944;
Castellanos, 1953; Carlini and Scillato-Yane
´, 1999; Carlini et al.,
2008b); d) Doedicurinae (late Miocene–early Holocene; Hoff-
stetter, 1958; Paula Couto, 1979); e) Hoplophorinae
(¼Sclerocalyptinae) (middle Miocene–early Holocene; Scillato-
Yane
´and Carlini, 1998a; Scillato-Yane
´et al., 1995; Carlini and
Scillato-Yane
´, 1999; Zurita et al., 2005; Zurita, 2007).
In this context, the Hoplophorinae are probably the Glypto-
dontidae subfamily with greatest taxonomic complexity and
Abbreviations: GABI, Great American Biotic Interchange; M–m, upper and lower
molariforms, respectively; CC, Museo Universitario ‘‘Florentino y Carlos Ameghino’’,
Universidad Nacional de Rosario (ex Instituto de Fisiografı
´a y Geologı
´a ‘‘Alfredo
Castellanos’’), Rosario; MACN, Seccio
´n Paleontologı
´a de Vertebrados, Museo
Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’; MLP, Divisio
´n Paleon-
tologı
´a Vertebrados, Facultad de Ciencias Naturales y Museo, Universidad Nacional
de La Plata; MSP, Museo Paleontolo
´gico Municipal ‘‘Fray Manuel de Torres’’, San
Pedro; MMP, Museo Municipal de Ciencias Naturales del Mar del Plata ‘‘Lorenzo
Scaglia; RCS, ‘‘Royal College of Surgeons’’, London, UK.
*Corresponding author.
E-mail address: azurita@cecoal.com.ar (A.E. Zurita).
Contents lists available at ScienceDirect
Quaternary International
journal homepage: www.elsevier.com/locate/quaint
1040-6182/$ – see front matter Ó2009 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2009.06.029
Quaternary International 210 (2009) 82–92
Author's personal copy
morphological heterogeneity. This is due to the remarkable number
of taxa included within its seven recognized tribes (Hoplophorini,
Neuryurini, Palaeohoplophorini, Plohophorini, Panochthini, Loma-
phorini and Neothoracophorini) and the poor morphological
characterization of most Hoplophorinae species (Zurita, 2007).
Indeed, these have been mostly recognized on the basis of typo-
logical criteria characteristic of the 19th and early 20th centuries. In
the words of Hoffstetter (1958: 577):‘‘Il est difficile de donner de l’
ensemble une de
´finition pre
´cise, car trop de genres, surtout ceux du
Mio-Plioce
´ne, sont connus d’une façon tre
`s fragmentaire.’’
In addition, recent preliminary phylogenetic analyses have
questioned the monophyly of this subfamily (Fernı
´cola et al., 2002;
Fernı
´cola, 2005).
The Hoplophorinae Hoplophorini (¼Sclerocalyptini) have
traditionally presented a similar prospect, with remarkable taxo-
nomic overestimation; about 8 genera and 26 species have been
traditionally included in this tribe (see, among others, Ameghino,
1889, 1895, 1919; Rovereto, 1914; Castellanos, 1925, 1948, 1951;
Cabrera, 1939, 1944; Scillato-Yane
´and Carlini, 1998a; Zurita, 2002).
Thus, Neosclerocalyptus (¼Sclerocalyptus ¼Hoplophorus ¼Isolinia;
see Paula Couto, 1957; Zurita et al., 2007), one of the two Pleisto-
cene genera of Hoplophorini, is the taxon with greatest proposed
taxonomic overestimation. About 15 species have been attributed
to this taxon (see Mones, 1986), four of which would be Ensenadan
in age (see Ameghino, 1889).
However, a recent revision of this tribe has shown that the
Hoplophorini from the present Argentinian territory are repre-
sented with certainty by 2 genera (Eosclerocalyptus and
Neosclerocalyptus)and7species(Zurita, 2007), and probably an
additional genus, Eonaucum, although little material of the latter
is known (see Scillato-Yane
´and Carlini, 1998a). Furthermore, the
stratigraphic distribution of these genera agrees with the pattern
of morphological changes observed in relation to inferred
climatic changes. The present contribution discusses the species
of Neosclerocalyptus (Ensenadan–Lujanian; early Pleistocene–
early Holocene) of Ensenadan age (early Pleistocene to early-
middle Pleistocene) considered as valid, and assesses their
significance as biostratigraphic, paleobiogeographic and paleo-
environmental indicators.
The nomenclatural and taxonomical schemes used here agree
with the proposals of Paula Couto (1957, 1965) and Zurita et al.
(2007). The chronological and biostratigraphic schemes used here
follow those of Cione and Tonni (2005).
2. Paleontological systematics
Order Cingulata Illiger, 1811
Superfamily Glyptodontoidea Gray, 1869
Family Glyptodontidae Gray, 1869
Subfamily Hoplophorinae Huxley, 1864
Tribe Hoplophorini Huxley, 1864
Genus Neosclerocalyptus Paula Couto, 1957
Synonymy.Sclerocalyptus Ameghino, 1891;Isolinia Castellanos,
1951 [partim] n. sin; Chacus Zurita, 2002 n. sin.
Type species.Glyptodon ornatus Owen, 1845
Stratigraphic and geographic distribution. Early Pleistocene–
early Holocene (Ensenadan–Lujanian) of the Argentinian provinces
Buenos Aires, La Pampa, Co
´rdoba, Mendoza, San Luis, Santa Fe,
Entre Rı
´os, Corrientes, Chaco, Santiago del Estero, Tucuma
´n, For-
mosa and Salta (Zurita et al., 2005). Pleistocene of Uruguay,
Paraguay (Villa Hayes and Boquero
´n Departments) (Zurita, 2007)
and Bolivia (N
˜uapua and Santa Cruz de la Sierra; see Ameghino,
1889; Hoffstetter, 1968; Zurita et al., 2009).
Diagnosis. Small-sized compared with other Pleistocene Glyp-
todontidae. Cephalic shield wide (especially at naso-frontal level),
with subquadrangular outline and formed by numerous osteo-
derms (approximately 80) with ornamentation similar to that of
the dorsal carapace. Dorsal carapace low and elongated, sub-
cylindrical, dorsal outline almost straight, formed by approximately
50–55 transversal rows of osteoderms at the sides of the carapace;
in some cases carapace with slight middle constriction separating
cephalic and caudal halves; antero-lateral margins flaring outwards
as ‘‘wings’’ and formed by very small pentagonal or hexagonal
osteoderms; these osteoderms with thick perforations at each
intersection of the sulcus surrounding the central figure with the
sulci that separate peripheral figures, in the area adjacent to the
cephalic notch. Osteoderms with primitive ornamentation similar
to that of Propalaehoplophorinae, relatively thin and large, formed
by a flat or somewhat concave central figure surrounded by a row of
smaller polygonal figures, 7–10 in the anterior and middle regions
of the carapace, and 10–12 (exceptionally 13 or 14) in the posterior
region (occasionally with an accessory row of peripherals in this
area), with shallow sulci separating the figures from each other.
Lateral carapace osteoderms rectangular and anteroposteriorly
elongated, with a large central figure, and arranged in transversal
rows; this arrangement disappearing toward the dorsum where the
osteoderms become pentagonal or hexagonal withmore developed
peripheral figures. Caudal shield formed by four or five rings (each
with two osteoderm rows) and a cylindrical–conical caudal tube
somewhat flattened dorsoventrally and tapering distally, with two
large terminal dorsolateral figures. Laterally with 5–7 oval figures,
increasingly larger toward the apex; the last four lateral figures
occupying more than half of the tube length. Each central figure
may be surrounded by one or two rows of peripherals, contrasting
with the single peripheral row of Eosclerocalyptus. Skull with highly
developed and pneumatized fronto-nasal sinuses (maximally
developed in Lujanian species), directed ventrally, although not as
much as in Panochthus, and separated from the rest of the skull by
an evident V-shaped notch; orbit closed posteriorly by a postorbital
apophysis. Area anterior to orbital (nasal and frontal) notches
semicircular, a condition associated with increased pneumatization
of the sinuses. Infraorbital foramen projected in a plane passing at
M3 level, differing from Eosclerocalyptus in which this plane passes
through the M3–M4 boundary. Predental region of palate narrower
and more extended anteroposteriorly than in Eosclerocalyptus and
Propalaehoplophorinae, with dental series ending posteriorly to
zygomatic process of squamosal. Paroccipital processes more
developed and more laterally projecting than in Eosclerocalyptus.
Supraoccipital is placed at an angle of 40–50
with respect to
basilar plane. First molariform simple and elongated ante-
roposteriorly; second molariform with incipient lobation; other
molariforms clearly trilobed. Molariforms M6–M8 have bilaterally
symmetrical lobes and anterior wall of first lobe is flat or slightly
convex. Humerus similar to that of Hoplophorus euphractus,
somewhat shorter and more robust, with posterior margin of
deltoid ‘V’ more developed and entepicondylar foramen more
proximal, as in Panochthus. Scapula has sinuous acromion. Pes and
manus tetradactyl due to loss of digit I; digit V is highly reduced.
2.1. Neosclerocalyptus pseudornatus (Ameghino, 1889)
(Fig. 1 A–H)
Synonymy.Hoplophorus pseudornatus Ameghino, 1889;
Sclerocalyptus pseudornatus (Ameghino, 1889).
Lectotype. MACN 1233, fragment of dorsal carapace with 13
osteoderms.
Paralectotype.MACN 13084, distal half of caudal tube.
A.E. Zurita et al. / Quaternary International 210 (2009) 82–92 83
Author's personal copy
Type locality. ‘‘Toscas del Rı
´o de La Plata’’, (caliche duricrusts
outcropping at the margins and bed of the river), Buenos Aires City,
Argentina (34
38
0
S, 58
28
0
W).
Referred materials.CC 20, caudal tube; CC 107, complete skull;
CC 21 caudal tube; CC 167, lateral right portion of dorsal carapace;
MMP 234, right half of skull; MACN 8579, complete skull; MACN
8773, complete skull; MACN 2936, left femur; MACN 8676, right
femur; MACN 5858, right hemimandible; MACN 2262, right
hemimandible; MACN 5007, caudal tube; MACN 13084, caudal tube
(figured by Ameghino, 1889: pl., LXLII, Figs. 1–3); MACN 1798,
caudal tube; MACN 5028, caudal tube; MACN 7075, caudal tube;
MACN 12543, carapace fragment; MACN 1930, large fragment of
right humerus; MACN 1966, right mandibular ramus; MACN 1989,
left humerus; MACN 2209, left radius; MACN 2014, left radius;
MACN 2018, left radius; MACN 2262, left mandibular ramus; MACN
2332, distal half of left femur; MACN 2315, atlas; MACN 2273, right
Fig. 1. Neosclerocalyptus pseudornatus. (A) Skull in frontal view, (B) In dorsal view, (C) Lateral view, (D) Ventral view (MACN 8579), (E) Left hemimandible in lateral view (MACN
5858), (F) Cephalic shield in dorsal view (MACN 8773), (G) Caudal tube in dorsal view, (H) Caudal tube in lateral view (MACN 7075).
A.E. Zurita et al. / Quaternary International 210 (2009) 82–9284
Author's personal copy
tibiofibula; MACN 2276, distal half of caudal tube; MACN 1793,
proximal portion of left ulna; MACN 2019, distal half of left
humerus; MACN 2316, proximal portion of right ulna; MACN 2272,
right tibiofibula; MACN 1836, caudal tube; MACN 2232, distal half
of left femur; MACN 2076, distal half of left femur; MLP 16–144,
distal half of caudal tube.
Diagnosis. Skull with parietoccipital region directed dorsally,
but not as much as in Neosclerocalyptus ornatus, with sagittal crest
similar to that of Eosclerocalyptus proximus. Fronto-nasal sinuses
little pneumatized and laterally expanded, with external bony
lamina recurved in helicoidal fashion (Fig. 1A,C). Anteriormost
portion of nasals is not preserved in any of the specimens. Area
between fronto-nasal sinuses and postorbital processes of frontal
bone more developed transversally and anteroposteriorly than in
N. ornatus,Neosclerocalyptus gouldi Zurita et al. (2008) and Neo-
sclerocalyptus paskoensis, due to lesser pneumatization of the
sinuses (Fig. 1B,C). Zygomatic arches high and straight, similar to
those of E. proximus and N. ornatus and clearly different from those
of E. tapinocephalus (whose zygomatic arches become shorter
toward the orbital notches) (Fig. 1C). Descendent processes of
maxillae very developed at jugal level. Infraorbital foramen is small,
but with extremely thick lower bony margin (Fig. 1D). Foramen
magnum is larger transversally than dorsoventrally, oval in shape.
First upper molariform is elliptical in cross section, with greater
axis parallel to the sagittal plane; M2 with incipient lobation; M3–
M8 clearly trilobed (Fig. 1D). Cephalic shield wide (especially at
naso-frontal level) and formed by numerous osteoderms (approx-
imately 80) with ornamentation similar to that of dorsal carapace
(Fig. 1F). Mandible and caudal tube are morphologically similar to
those of N. ornatus (Fig. 1E,G,H). Dorsal carapace very similar in
shape to that of N. ornatus, that is, low, subcylindrical, elongated
and with almost straight dorsal outline, different from more
globose carapace of E. proximus; in one specimen (CC 167), plate
rows adjacent to caudal notch with accessory rows of peripheral
osteoderms at proximal and distal margins.
Chronological and geographical distribution. Early-middle
Ensenadan (1.07–0.98 Ma) (early Pleistocene) (Fig. 3). ‘‘Toscas del
Rı
´o de La Plata’’ (see above; Buenos Aires City and Olivos) and Mar
del Plata, Buenos Aires province, Argentina (Fig. 4).
Historical and taxonomic aspects. This species was originally
recognized by Ameghino (1889) as H. pseudornatus on the basis of
fragmentary dorsal carapace osteoderms and a proximally broken
caudal tube collected from the sediments of the ‘‘Toscas’’ of Rı
´odeLa
Plata (Buenos Aires). These materials had been figured earlier by
Lydekker (1887) as Hoplophorus sp. a?; some time afterwards,
however, this same author (Lydekker,1894) classified these remains
as Lomaphorus ornatus. In his published response, Ameghino (1895)
again asserted the validity of this species and summarized its main
distinctive characters. Since Ameghino (1889: 808–809) did not
choose a holotype from the type serie (MACN 1233 and MACN
13084), we select (in the context of this taxonomic revision) the
MACN 1233 as the lectotype of this species (see ICZN, 1999, art. 74).
2.2. N. ornatus (Owen, 1845)(Fig. 2A–F)
Synonymy.G. ornatus Owen, 1845;Hoplophorus ornatus (Owen,
1845); Sclerocalyptus ornatus (Owen, 1845).
Holotype. RCS 3606 (ex RCS 554), four dorsal carapace osteo-
derms (Owen, 1845: 119).
Type locality. Near Matanzas river, approximately 32 km south
of Buenos Aires City, Argentina.
Neotype.(MACN?; missing), complete dorsal carapace, caudal
rings and caudal tube (figured by Burmeister, 1871, pl. XVII)
(Lydekker, 1887: 128) (see discussion below)
Type locality: Buenos Aires province, Argentina.
Referred materials.MLP 16–28, complete skeleton and dorsal
carapace (figured by Lydekker, 1894; plates XI, XII and XIII); MMP
4300, complete dorsal carapace and caudal tube; MACN 8091,
dorsoventrally compressed skull; CC 656, skull, mandible and
cephalic shield; MACN 11948, mandibular ramus; MSP 12, almost
complete dorsal carapace, complete caudal rings and caudal tube,
and very fragmentary posterior skull with matching portion of
cephalic shield.
Diagnosis. Hoplophorini slightly larger than N. pseudornatus,
similar in size to N. gouldi. Skull with convex profile, parietocci-
pital region elevated, even more so than in N. pseudornatus, and
marked sagittal crest (Fig. 2A–C). Fronto-nasal sinuses more
defined than in N. pseudornatus, ‘funnel’-shaped, inclined
ventrally, clearly pneumatized and laterally expanded (more so
than in N. pseudornatus and H. euphractus, but not as much as in
N. gouldi and N. paskoensis), separated from each other and from
the frontal and maxillary bones by evident V-shaped cleft; free
margins of nasals curved inward, especially at their middle and
lower portion (Fig. 2A–C). Region between fronto-nasal sinuses
and postorbital processes of frontal less developed transversally
and anteroposteriorly than in N. pseudornatus, due to greater
pneumatization of sinuses. Zygomatic arches high and straight,
very similar to those of N. pseudornatus and E. proximus (Fig. 2C).
Infraorbital foramen larger than that of N. pseudornatus, with
thinner lower bony margin very similar to that of N. gouldi
(Fig. 2D). Foramen magnum similar to that of N. pseudornatus, i.e.,
with oval outline. First molariform simple, elongated ante-
roposteriorly; second molariform with incipient lobation, as in
N. pseudornatus; remaining molariforms trilobed and similar to
those of other species of Neosclerocalyptus (Fig. 2D). Mandible
similar to that of N. pseudornatus and N. gouldi, with ascending
rami extended anteroposteriorly, although not as much as in
E. proximus, and inclined forward. Dorsal carapace elongated, low,
subcylindrical, and with dorsal outline almost completely straight,
with slight narrowing at the middle separating two parts: anterior
low narrow portion, and posterior somewhat higher and broader
portion (Fig. 2E). Some specimens have accessory rows of
peripheral figures at distal and proximal margins of osteoderm
rows adjacent to caudal notch, as in N. pseudornatus.
Chronological and geographic distribution. Middle-late
Ensenadan (0.98–0.40 Ma) (early-middle Pleistocene) (Fig. 3). Mar
del Plata and San Pedro (Buenos Aires province), Granadero
Baigorria (Santa Fe province) (Fig. 4).
Historical and taxonomic aspects. This species was recognized
and figured by Owen (1845: Fig. 5) based on a series of four or five
associated osteoderms from a dorsal carapace collected from
Pleistocene sediments in the vicinity of Matanzas river, some 32 km
southwest from Buenos Aires City (Owen, 1845; see Rusconi, 1930).
Originally, Owen (1845) assigned these remains to genus Glyptodon
Owen, 1839 and recognized the species ‘‘G. ornatus’’, whose type
was deposited in the collections of the ‘‘Royal College of Surgeons’’
(RCS) (Londres), where Owen worked as curator. Although the
precise stratigraphic provenance of the material collected by Owen
(1845) (RCS 3606) cannot be established, diverse authors have
recognized Ensenadan outcroppings at several sectors of Matanzas
river (see Rusconi, 1930,1936). Added to these considerations is the
fact that the RCS collection was greatly damaged during a series of
bombing raids that affected London in 1941. As a consequence, only
175 of the original 5200 catalogued specimens could be rescued
(Currant, pers. comm.). Cave’s (1942) list of the materials that were
not destroyed does not include the type of N. ornatus, and conse-
quently it should be considered lost.
Given that the type material of this Hoplophorini species did not
present relevant diagnostic characters, Lydekker (1887: 128 ‘‘Since
there may be possibly be a doubt as to the identity of the complete
A.E. Zurita et al. / Quaternary International 210 (2009) 82–92 85
Author's personal copy
carapace figured by Burmeister in the ‘‘An. Mus. Buenos Aires’’ under
the present name with the fragment to which the name ‘‘G’’ ornatus
was applied, it will be advisable to regard the former as the type’’)
designated as neotype a very well preserved dorsal carapace with
associated caudal rings and caudal tube that had been figured by
Burmeister (1871; plate XVII). Unfortunately, this material has not
been found in the collections of the Museo Argentino de Ciencias
Naturales ‘‘Bernardino Rivadavia’’, and is thus considered as
missing.
Lastly, in a taxonomic revision, Lydekker (1894: 20–24)
described and illustrated an excellent specimen of this species
(MLP 16–28;Fig. 2A–F), from the Ensenadan of Mar del Plata,
Buenos Aires province, which is morphologically almost identical to
the specimen that he had earlier designated as the neotype.
Fig. 2. Neosclerocalyptus ornatus (MLP 16–28). (A) Skull in frontal view, (B) In dorsal view, (C) In lateral view, (D) In ventral view, (E) Cephalic shield, dorsal carapace, caudal rings
and caudal tube in lateral view, (F) Complete skeleton in lateral view.
A.E. Zurita et al. / Quaternary International 210 (2009) 82–9286
Author's personal copy
Since then, and until now, for a period of 113 years, all the
specialists have directly or indirectly associated this specimen
(MLP 16–28) with the name N. ornatus (¼H. ornatus ¼S. ornatus)
(see, among others, Ameghino, 1895; Richter, 1911; Vinacci, 1939;
Hoffstetter, 1958; Pascual et al., 1966; Paula Couto, 1979;Zurita
et al., 2005).
To paraphrase Ameghino (1895: 845) ‘‘Es un soberbio ejemplar de
un individuo completamente adulto (.) y debera
´ser consultada
preferentemente por los paleonto
´logos porque representa aprox-
imadamente la forma exacta del animal’’ (‘‘It is a superb sample of
a fully grown adult (.) and it should be preferably consulted by
paleontologists because it represents approximately the actual
shape of the animal’’).
3. Other Ensenadan Hoplophorini? problematic issues
As previously mentioned, Neosclerocalyptus is a genus with
remarkable taxonomic overestimation, since most of the species
were recognized based on fundamentally typological taxonomic
criteria (see Giraudo, 1997; Hevia and Romero, 1999). Consequently,
many of the characters used for recognition of new species (e.g.
osteoderm thickness and number of peripheral figures in dorsal
carapace osteoderms; shape, length, degree of flattening and
number of lateral figures in the caudal tube) are currently insuffi-
cient, as their intraspecific variability has not been adequately
evaluated, as demonstrated for other Glyptodontidae Hoplophor-
inae (Perea, 2005).
In this taxonomic context, ‘‘Hoplophorus’’ perfectus Gervais and
Ameghino is a species created in 1880 whose type is a small dorsal
carapace fragment collected from the ‘‘Toscas del rı
´o de La Plata’’,
Buenos Aires City (MACN 1232). Later on, Ameghino figured the
type material (1889, pl. LXIV, Fig. 1) while he also associated new
materials to this taxon without adequate justification, particularly
a caudal tube (MACN 7079) possibly from the Bonaerian deposits,
or even the Lujanian beds, of Luja
´n river at Colonia Salazar
(unpublished.) (Ameghino, 1889, pl. XC, Figs. 1–3). The characters
used by Gervais and Ameghino (1880), for the original identifi-
cation of the species, are insufficient, because the only diagnostic
features mentioned by these authors are larger size of osteoderms
and central figures, and greater number of peripheral figures
(11–13). However, the materials that correspond to the lectotype
(three dorsal carapace osteoderms; MACN 1232) do not display
clear diagnostic characters that could link them to the Pleistocene
genus Neosclerocalyptus (Zurita et al., 2005; Zurita, 2007).
A thorough analysis of these osteoderms indicates that their
morphology does not correspond to the known variability for
dorsal carapaces of Glyptodontidae Hoplophorini. They probably
correspond to the tribe Panochthini (cf. Panochthus intermedius)
given the size proportion between central and peripheral figures
(see Lydekker, 1894).
‘‘Hoplophorus’’ scrobiculatus, a taxon described by Ameghino
(1889), is another recognized Ensenadan species. The story of this
species is quite singular, as Ameghino designated as type material
a dorsal carapace and caudal tube that were deposited in the
Fig. 3. Chronological distribution of the species of Neosclerocalyptus.
Fig. 4. Geographic distribution of Neosclerocalyptus pseudornatus (circle) and N.
ornatus (triangle). 1 Ciudad de Buenos Aires, 2 Olivos, 3 Mar del Plata, 4, San Pedro, 5
Granadero Baigorria.
A.E. Zurita et al. / Quaternary International 210 (2009) 82–92 87
Author's personal copy
collections of the then Museo Nacional de Buenos Aires (today the
Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’),
and which he presented as associated materials. At the time,
Ameghino noted that the small size of the specimen, as well as the
particular morphology of the osteoderms, indicated a supposed
transition to genus Lomaphorus. Thus, the diagnostic characters
supplied by Ameghino (1889: 817) (e.g. number of peripheral
figures surrounding each central figure, slight concavity on dorsal
surface of osteoderms, large penultimate lateral figure of caudal
tube, etc.) are common to the Hoplophorinae Hoplophorini.
Furthermore, he only illustrated three associated osteoderms
(Ameghino, 1889; pl. LXXXV, Fig. 4).
However, six years later, Ameghino (1895: 543–544) stated that
this species should disappear because it had been described based
on a dorsal carapace that was referable to the Lomaphorini
‘‘Lomaphorus compressus’’ and a caudal tube probably assignable to
Neosclerocalyptus, although, as stated above, these materials had
never figured. Unfortunately, the materials on which Ameghino
based his description have not been found in the collections of
Museo Argentino de Ciencias Naturales, and consequently this
taxon should be considered in querenda.
4. Paleobiogeography and biostratigraphy of
Neosclerocalyptus, with emphasis on Ensenadan species
The biogeographical history of the Glyptodontidae Hoplophorini
(late Miocene–early Holocene) is apparently restricted to the
southern cone of South America, where their range extends with
certainty from about 20
Sto38
S latitudinally and from 43
Wto
66
W longitudinally. This contrasts with the case of the Glypto-
dontidae Glyptodontinae, the other characteristic and frequent
taxon for the late Neogene, whose first records date from the late
Miocene of Colombia (Carlini et al., 2008b); the distribution of
glyptodontines during the Pleistocene included South America
from Argentina to Venezuela, Central America, and North America
up to ca. 35
N(Marshall et al., 1984; Tonni and Scillato-Yane
´, 1997;
Rinco
´n, 2006; Carlini et al., 2008a). In addition, this is probably the
only Glyptodontidae subfamily that participated in the Great
American Biotic Interchange (GABI) and underwent cladogenesis,
which is reflected in five described species (Gilette and Ray, 1981;
Flynn et al., 2005; Morgan and White, 2005; White and Morgan,
2005; Carlini et al., 2008a).
Neosclerocalyptus is a typical Pampean genus of Hoplophorinae
Hoplophorini, geographically restricted to the present territories of
Argentina, Bolivia, Paraguay and Uruguay (Zurita et al., 2005;
Zurita, 2007). It was probably well adapted to progressively colder
and more arid environments (Tonni and Fidalgo, 1979; Fidalgo and
Tonni, 1983), as the records of this genus are very scarce in areas
that were relatively humid and warm during most of the Pleisto-
cene (Argentinian Mesopotamia, western Uruguay and southern
Brazil) (Kraglievich, 1932; Scillato-Yane
´et al., 1998; Carlini et al.,
2004; Noriega et al., 2004), but are very abundant in the area that
currently comprises the Pampean region and central-northern
Argentina (Zurita et al., 2004), and they are the only well repre-
sented Hoplophorini in Paraguay. Accordingly, Neosclerocalyptus is
characterized by an evident and progressive increase of pneuma-
tization and lateral expansion of the fronto-nasal sinuses, a feature
that becomes even more evident in the middle and late Pleistocene
species. This increasing pneumatization of the fronto-nasal sinuses
is a probable response to the cold arid or semiarid Pleistocene
environments (see Clapperton, 1993; Tonni et al., 1999a,b; Cione
and Tonni, 2001; Prado et al., 2001; Cione et al., 2003; Tonni et al.,
2003; Zurita et al., 2005).
Latitudinally, the northernmost record of the genus in Argentina
corresponds to Las Lajitas, in Salta province (24
42
0
S) (see Zurita
et al., 2002); in Argentina, the distribution of species of this genus
ranges from 26
41
0
S (Avia Terai, Chaco province) to 38
44
0
S (Bahı
´a
Blanca, Buenos Aires province), and longitudinally from 57
33
´W
(Mar del Plata, Buenos Aires province) to 65
02
0
W (Merlo, San Luis
province; see Chiesa et al., 2000, 2005). Outside Argentina, the
northernmost records correspond to the localities of N
˜uapua (20
52
0
S; 63
04
0
W; see Hoffstetter, 1968) and Santa Cruz de la Sierra
(17
47
0
S, 63
11
0
W; see Zurita et al., 2009), in Bolivia.
Taxonomically, four Neosclerocalyptus species can be recognized
with certainty. As we have mentioned, N. pseudornatus and
N. ornatus are recorded in the Ensenadan Stage (early Pleistocene to
early-middle Pleistocene), while a single species (N. goudi)is
recognized for the Bonaerian Stage (middle Pleistocene). Lastly,
N. paskoensis is an exclusively Lujanian (late Pleistocene–early
Holocene) species (Fig. 3), widely distributed in the present
Argentinian territory between 26
41
0
S (Avia Terai, Chaco province)
and 38
44
0
S (Bahı
´a Blanca, Buenos Aires province) (Zurita, 2007).
Chronologically, the Ensenadan Stage (early Pleistocene to early-
middle Pleistocene; 1.8–0.4 Ma) spans a wide interval of more than
1 million years, during which numerous climatic–environmental
changes that drastically affected faunal composition and diversity
(Cione and Tonni, 1999, 2001; Rabassa et al., 2005; Soibelzon, 2005)
took place within a general climate cooling trend. In this context,
the paleoenvironmental evidence suggests that the Ensenadan
Stage of southern South America was characterized by the defined
predominance of cold arid or semiarid climates interspersed with
brief warmer and more humid pulses (Tonni and Cione, 1994, 1995;
Tonni et al.,1999b; Cione and Tonni, 2001; Soibelzon et al., 2006a).
Specific pulses have been detected at approximately 1.0 Ma (Nabel
et al., 2000) and 0.4 Ma (Soibelzon et al., 2006a). Considering the
entire Pleistocene (ca. 2.6 Ma to 10 ka B.P.), the Ensenadan Stage
seems to have been the period with the highest proportion of
mammals adapted to open arid environments (Bobe Quinteros
et al., 2004). In the current Pampean region, these climatic–envi-
ronmental processes favored the expansion of characteristic
elements from the Central and Patagonian Domains (e.g. Microcavia
australis,Lestodelphys halli,Tolypeutes matacus). One distinctive
feature of this period is the large size attained by some taxa, greater
than that observed during the middle Pleistocene–early Holocene
(e.g. P. intermedius,Glyptodon munizi,Toxodon ensenadensis)
(Scillato-Yane
´and Carlini, 1998b; Cione et al., 2003; Soibelzon et al.,
2006b), as well as the marked proliferation of herbivorous mega-
mammals belonging to several orders (e.g. Notoungulata, Xenar-
thra Phyllophaga, Litopterna; Tonni and Cione, 1994).
In the current Pampean region, the sediments of Ensenadan age
correspond to the Miramar and Ensenada formations (including
Ameghino’s ‘Preensenadan’ stage) and the outcroppings of Vorohue
´
and San Andre
´s formations located north of Mar del Plata
(Soibelzon, 2005).
In this context, N. pseudornatus (ca. 1.07–0.98 Ma) is the oldest
species of the genus, and already begins to show a tendency toward
increased pneumatization and development of the fronto-nasal
sinuses, possibly as a response to climatic–environmental condi-
tions (Zurita et al., 2005; Zurita, 2007). To date, the geographical
distribution of this species is restricted from 34
31
0
S (‘‘Toscas del
Rı
´o de La Plata’’, Buenos Aires City) to 38
S (Mar del Plata), i.e.,
central-eastern Argentina (Fig. 4).
From a stratigraphic perspective, the sediments that correspond
to ‘‘Toscas del Rı
´o de la Plata’’ were initially correlated by Ameghino
(1889) with the uppermost (‘cuspidal’) Ensenadan beds. Unfortu-
nately, as already noted by Ameghino and Kraglievich (1921: 136)
‘‘.los tı
´picos yacimientos del Ensenadense con su rica y variada fauna
esta
´n destinados a desaparecer totalmente en un futuro no lejano
a consecuencia de las modificaciones que experimenta.la antigua
ribera de Buenos Aires, a lo largo de la cual.la bajante del rı
´o ponı
´aal
A.E. Zurita et al. / Quaternary International 210 (2009) 82–9288
Author's personal copy
descubierto las cla
´sicas toscas’’ (the typical Ensenadan fossil beds
with their rich and varied fauna are doomed to complete disap-
pearance in the near future due to the modifications suffered by.
the old coastline of Buenos Aires, along which.low levels of the
river have uncovered the typical ‘tosca’ [caliche duricrusts] rocks).
These sediments are currently considered to represent a period
between the Jaramillo and Olduvai events, corresponding to Chron
1 (reversed magnetic polarity). In this sense, the presence of the
Notoungulata Mesotherium cristatum, considered as a guide fossil
for the upper Ensenadan Stage, supports this temporal assignation
(Tonni and Cione, 1994), and so does the presence of the bird
Pseudoseisura sp. nov.(Tonni et al., 1999b). Chronologically, this
implies that these sediments could be dated between 1.07 Ma and
0.98 Ma (early-middle Ensenadan Stage) (see Soibelzon et al.,
2008).
From a paleoenvironmental viewpoint, the micromammals of
this age (1.0 Ma; middle Ensenadan Stage) indicate clear predom-
inance of cold arid climatic conditions, interspersed with some
brief humid periods (Tonni et al., 1999b). In an independent study,
Bonadonna and Alberdi (1987) detected an important cold arid
pulse (ca. 1.0 Ma) that implied some faunal changes. Thus, the
temporal distribution of this Hoplophorini species coincides
partially with the ‘‘Great Patagonian Glaciation’’, which took place
between 1.168 and 1.016 Ma (Rabassa et al., 2005). The same fossil
beds from which the remains of N. pseudornatus were collected
(‘‘Toscas del Rı
´o de La Plata’’), have also yielded remains of Lama
guanicoe (Muller) (Menegaz and Ortiz Jaureguizar, 1995; Menegaz,
2000), a clear indicator of dry–cold climates, along with M. crista-
tum Serres, 1867, and probably also ‘Brasilic’ fauna such as Tapirus
Brisson (the southernmost record of this genus), Calomys Water-
house, and Procyonidae, which indicate more tropical and humid
climates (Bond et al., 1995; Tonni and Cione, 1995; Bond, 1999;
Soibelzon et al., 2005). The possible coexistence of taxa with
different ecological requirements and current allopatric distribu-
tion could be interpreted as an indicator of the occurrence of
heterogeneous environments (Tonni et al., 1998). However, given
the lack of high resolution biostratigraphic studies, the possibility
that the taxa indicative of warmer climates and those suggestive of
colder drier climates were not actually coeval, but the result of
time-averaging sediments, cannot be dismissed.
Lastly, the specimen from Mar del Plata (MMP 234)was
collected from Miramar Formation (see Isla and Dondas, 2001),
which was assigned by Cione and Tonni (1995, 1999) to the Ense-
nadan Stage sensu lato (s.l.). The paleomagnetic studies made by
Orgeira (1987) suggest a Matuyama s.l. paleomagnetic age, greater
than 0.7 Ma and less than 2.41 Ma.
N. ornatus is a species restricted to the final Ensenadan Stage
(0.98–0.40 Ma) (Fig. 3). Its geographical distribution is wider than
that of N. pseudornatus, encompassing central and central-eastern
Argentina (Buenos Aires and Santa Fe provinces), from 32
53
´S
(Granadero Baigorria, Santa Fe province) to 38
S (Mar del Plata,
Buenos Aires province) (Fig. 4). Stratigraphically, the unquestion-
able records of N. ornatus in Buenos Aires province correspond to
two specimens (MLP 16–28 and MACN 8091) collected from the
cliffs situated north of Mar del Plata, and possibly from the Miramar
Formation (see Kraglievich, 1952). These sedimentary sequences
have been little studied compared to those located south of Mar del
Plata, which have been the subject of intense paleontological,
paleomagnetic and sedimentological studies (Bidegain et al., 2003).
Magnetostratigraphic (Bidegain et al., 1998, 2003) and
biostratigraphic (Tonni et al., 1998) studies of Ensenadan levels
indicate that these sediments are somewhat more recent than
those from the ‘‘Toscas del Rı
´o de La Plata’’, and even though no
absolute datings have been made, their age could range between
0.98 and 0.40 Ma (middle-late Ensenadan Stage; Tonni et al., 1998;
Bidegain et al., 2003). Another record, represented by a complete
dorsal carapace associated with a caudal tube (MMP 4300), has
been unquestionably collected from the Miramar Formation in Mar
del Plata (see Isla and Dondas, 2001). The sediments from this unit
were originally nominated by Ameghino (1908) as the Ensenada
Formation, and by Frenguelli (1928) as part of the Ensenadan
Stage. It was formally recognized by Kraglievich (1952), who
designated it as the Miramar Formation. This formation, which
underlies the San Andre
´s Formation and overlies the Arroyo Seco
Formation (Bidegain et al., 1998), is 1–3 m thick and primarily
composed of fluvio-lacustrine sediments, including several litho-
logical types: ‘‘.conglomerados de fenoclastos de limo endurecido
cementados por limo arcilloso verdoso o pardo grisa
´ceo. Lentes de
limo muy arcilloso verde o azul y camadas lenticulares de acarreo
fluvial compuestas por pequen
˜os fragmentos rodados de rocas.en su
seccio
´n superior comprende bancos loe
´sicos pardos cubiertos por una
costra calca
´rea.’’ (‘‘.conglomerates of hardened silt phenoclasts
cemented by greenish or grayish brown clayey silt. Blue or green
highly clayey silt lenses and fluvial-transported lenticular bedding
formed by small rock pebbles.at its upper section includes
brown loessic banks covered by a calcareous crust.’’) (Kraglievich,
1952: 18; see also Isla and Dondas, 2001). Early on, Orgeira (1987)
and Tonni et al. (1992) correlated this unit with the Matuyama
Chron (2.48–0.73 Ma). Later, Cione and Tonni (1995, 1999)
assigned this formation to the Ensenadan Stage s.l. (ca. 1.8–0.4 Ma).
Thus, the same levels that yielded these three N. ornatus spec-
imens (MMP 4300,MLP 16–28 and MACN 8091), contain an
assemblage of vertebrates (Lestodelphys halli,Tympanoctomys bar-
rerae, Thinocoridae) that suggests persistence of the cold arid
climate that existed during the Sanandresian–Ensenadan stages
(Tonni et al., 1998). This could be correlated with a global cooling
event that took place in the 0.80–0.50 Ma lapse (Tonni et al., 1998).
Along these lines, Verzi et al. (2002) have noted the presence of the
octodontid Tympanoctomys cordubensis (Ameghino, 1889) in what
is currently the territory of Co
´rdoba province (near the city capital)
as well as in Buenos Aires province, in sediments dated at 0.90–
0.78 Ma. Tympanoctomys is currently considered as the South
American rodent most adapted to xeric conditions (Ojeda et al.,
1996), and its presence demonstrates the existence of cold arid
climate during that lapse. Similarly, the sediments that are close to
the transition between the Matuyama and Brunhes Chrons (ca.
0.78 Ma) indicate a shift toward colder and more arid conditions
(Soibelzon et al., 2006a).
Another specimen comes from the Ensenadan beds at San
Pedro, Buenos Aires province (MSP 12). Diverse paleomagnetic and
magnetostratigraphic studies (Bobbio et al.,1986; Nabel et al.,1993;
Nabel, 1993) have indicated that these sequences correspond to the
upper section of the Ensenada Formation (late Ensenadan Stage)
and the Buenos Aires Formation (Bonaerian Stage), and noted that
the Brunhes–Matuyama boundary (0.73 Ma) was found at the
uppermost part of the Ensenada Formation. The lithological char-
acterization includes silty clayey sediments in its basal sector (unit I)
and silty-sandy sediments with light brown or yellowish gray
coloration near the top of the formation (units II–V). The entire
sequence is characterized by high content of volcanic glass, possibly
of Eolian origin, that becomes especially evident within the levels
closer to the Brunhes–Matuyama boundary (Nabel et al., 1993).
From a sedimentological standpoint, a series of studies of the
uppermost section of the Ensenada and Buenos Aires formations at
the city of La Plata, Buenos Aires province, have shown alternance
of loess and paleosols throughout the sequence, indicating cyclic
climatic changes from cold and arid or semiarid (loess) to warmer
and more humid (paleosols) (Tonni et al., 1999b).
A series of paleomagnetic studies performed on sediments from
the vicinity of Baradero, Buenos Aires province (see Nabel et al.,
A.E. Zurita et al. / Quaternary International 210 (2009) 82–92 89
Author's personal copy
1993, 1993), show a significant increase in amounts of volcanic
glass toward the upper part of the Ensenada Formation, near the
Brunhes–Matuyama boundary (0.78 Ma). On the other hand, the
base of the Brunhes polarity zone shows predominantly loessic
sedimentation (Nabel et al., 2000). Primarily, both phenomena
would be correlated with increased aridity and cold, in accordance
with available paleontological evidence (Tonni et al., 1999b). The
taxa characteristic of warm humid environments (e.g. Tapiridae,
Procyonidae and Echimyidae) which are recorded during the early
and middle Ensenadan Stage (Nabel et al., 2000), disappear from
the fossil record later on, as taxa adapted to more arid environ-
ments (Microcavia,Reithrodon,Zaedyus and Tolypeutes;Tonni and
Cione, 1994) are recorded. However, the paleosol located at the
Brunhes–Matuyama boundary might basically indicate some
climatic stability and the presence of a conspicuous plant cover
(Nabel et al., 2000; Voglino and Pardin
˜as, 2005). The study of
a sequence attributable to the uppermost Ensenadan Stage (ca.
0.78 Ma), at Ramallo, in northern Buenos Aires province, agrees
with this model, as it shows a transition from warm humid to cold
arid conditions (e.g. Lestodelphys and Microcavia), immediately
above the Brunhes–Matuyama boundary (ca. 0.73 Ma) (Voglino and
Pardin
˜as, 2005).
Coincidentally, at lower latitudes (Tarija, Bolivia), MacFadden
(2000) has noted that the uppermost section of the Tolomosa
Formation (ca. 1.1–0.70 Ma) shows a change of faunal composition
in the transition from arid to warmer and more humid environ-
mental conditions (but see Coltorti et al., 2007).
Outside Buenos Aires province, a fourth record (CC 656)was
exhumed from the coastal cliffs of the Parana
´River at Granadero
Baigorria, Santa Fe province. In the 1960s decade, this material was
assigned by Castellanos (unpublished) to the ‘‘Lower Pleistocene’’
(¼‘‘Belgranense’’, ‘‘Belgranian’’ Stage). Currently, the ‘‘Belgranian’’
Stage in the sense of Ameghino (1889) and Castellanos correspond
to the lapse between the end of the Ensenadan and the beginning of
the Bonaerian stages (middle Pleistocene) (Cione and Tonni, 1995).
The provenance of this specimen possibly corresponds to sedi-
ments from the Rosario Formation, of Ensenadan age, which makes
up much of the coastal cliffs of Parana
´River from Rosario to Puerto
San Martı
´n. The sediments from this formation are characterized by
predominance of hard silts, reddish brown and green in color, with
scarce calcium carbonate concretions and manganese nodules
(Iriondo, 1987; Parent et al., 2002).
5. Conclusions
a) Four species of genus Neosclerocalyptus have been recognized
and described for the Ensenadan Stage: Neosclerocalyptus
scrobiculatus,Neosclerocalyptus perfectus,N. pseudornatus and
N. ornatus;however, this work demonstrates that only two of
them can be considered valid. Both of these species are bio-
stratigraphically significant because their biochrons are clearly
identified and confined. N. pseudornatus holds the earliest
record, with a biochron restricted to the early-middle Ensena-
dan Stage, between 1.07 and 0.98 Ma. The other Ensenadan
species, N. ornatus, is more recent, with a stratigraphic range
restricted to the late Ensenadan Stage, around 0.98–0.40 Ma.
b) Anatomically speaking, the major differences between the
two taxa are located in the skull. In N. pseudornatus, the fronto-
nasal sinuses are less pneumatized and less laterally expanded
than in N. ornatus, and their external wall exhibits evident
‘‘scrolled’’ shape due to the strong curvature of the free margins
of the nasal bones toward the sagittal plane. In addition, the
parieto-occipital region is not inclined upwards as markedly as
in N. ornatus. On the other hand, the fronto-nasal sinuses of
N. ornatus are better defined and clearly more pneumatized and
laterally expanded, with a characteristic ‘‘funnel’’ shape; the
infraorbital foramina of all specimens are larger than those of
N. pseudornatus.
c) From a paleobiogeographical perspective, the range of both
species is restricted to the current Pampean region. Thus far,
N. pseudornatus has been recorded in the ‘‘Toscas del Rı
´odeLa
Plata’’ (Olivos and Buenos Aires City) and in Mar del Plata.
Likewise, N. ornatus is recorded in San Pedro and Mar del Plata
(Buenos Aires province) and Granadero Baigorria (Santa Fe
province).
d) The climatic–environmental evidence provided by diverse
indicators suggests that these species lived under the arid/
semiarid and cold conditions characteristic of most of the
Pleistocene, and that they were probably well adapted to such
environments. In this context, the increased pneumatization and
lateral expansion of the fronto-nasal sinuses is interpreted here
as a probable response to this type of environments. In agree-
ment with this interpretation, the records of Neosclerocalyptus
from the middle and late Pleistocene are very abundant in Eolian
sediments of the Pampean region and central-northern
Argentina, where cold arid or semiarid climatic–environmental
conditions occurred during most of the Pleistocene; in contrast,
its records are very scarce in areas that experienced warmer and
more humid conditions during the same lapse (e.g. Argentinian
Mesopotamia, Uruguay and Brazil).
Acknowledgments
The authors wish to express their gratitude to Dr. E.P. Tonni and
Dr. J. Rabassa for their thorough reviews and helpful suggestions.
Finally, we also thank the curators of the different collections we
have studied for granting access to the materials. This work was
partially funded by grants PICT R 074 and PICTO-UNNE 00164 from
ANCyT.
References
Ameghino, F., 1889. Contribucio
´n al conocimiento de los mamı
´feros fo
´siles de la
Repu
´blica Argentina. Actas de la Academia Nacional de Ciencias, Co
´rdoba 6,
1–1027.
Ameghino, F., 1891. Mamı
´feros y aves fo
´siles argentinas. Especies nuevas, adiciones
y correcciones. Revista Argentina de Historia Natural 1 (4), 240–259. Buenos
Aires.
Ameghino, F., 1895. Sur les e
´dente
´s fossiles de l
´Argentine (Examen critique,
re
´vision et correction de l‘ ouvrage de M.R. Lydekker ‘‘The extinct Edentates of
Argentina’’). Revista del Jardı
´n Zoolo
´gico de Buenos Aires 3, 97–192.
Ameghino, F., 1908. Las formaciones sedimentarias de la regio
´n Litoral de Mar del
Plata y Chapalmalal. Anales del Museo Nacional de Buenos Aires 3 (10),343–428.
Ameghino, C., 1919. Sobre mamı
´feros fo
´siles del Piso Araucanense de Catamarca y
Tucuma
´n. I Reunio
´n Nacional de la Sociedad Argentina de Ciencias Naturales,
Actas, 151–152. La
´minas III y VII.
Ameghino, C., Kraglievich, L., 1921. Descripcio
´n del Megatherium gallardoi C. Ameg.
descubierto en el Pampeano Inferior de la Ciudad de Buenos Aires. Anales del
Museo Nacional de Historia Natural de Buenos Aires, 135–156.
Bidegain, J.C., Martı
´nez, G.A., Osterieth, M.L., Van Velzen, A., 1998. Magneto-
estratigrafı
´a de la secuencia Cenozoica tardı
´a de Camet (Norte de Mar del Plata),
provincia de Buenos Aires. V Jornadas Geolo
´gicas y Geofı
´sicas Bonaerenses,
Actas, 235–238.
Bidegain, J.C., Osterrieth, M., Martı
´nez, G., Rico, Y., 2003. Geologı
´a y registros
magne
´ticos en la localidad de Camet: correlacio
´n con perfiles complementarios
de Arroyo La Tapera y Santa Clara del Mar. I Jornadas de Geologı
´a del Cua-
ternario, Paleomagnetismo y Magnetismo ambiental, 1–11. edicio
´n electro
´nica
de Actas.
Bobbio, M.L., Devicenzi, S.M., Orgeira, M.J., Valencio, D.A., 1986. La magneto-
estratigrafı
´a del ‘‘Ensenadense’’ y ‘‘Bonaerense’’ de la Ciudad de La Plata
(excavacio
´n nuevo teatro Argentino): su significado geolo
´gico. Revista de la
Asociacio
´nGeolo
´gica Argentina 41 (1–2), 7–21.
Bobe Quinteros, R., Behresmeyer, A.K., Carrasco-Ormaza
´bal, G., 2004. Paleoclima y
evolucio
´n faunı
´stica en el Plio-Pleistoceno de Africa y Ame
´rica del Sur.
Ameghiniana 41 (4), 641–649.
Bonadonna, F.P., Alberdi, M.T., 1987. The N/Q boundary at 1.64 Ma? Mediterranean
Series Geology 6, 115–130.
A.E. Zurita et al. / Quaternary International 210 (2009) 82–9290
Author's personal copy
Bond, M., 1999. Quaternary native ungulates of southern South American. A
synthesis. In: Rabassa, J., Salemme, M. (Eds.), Quaternary of South America and
Antarctic Peninsula, vol. 12, pp. 177–205. Rotterdam.
Bond, M., Cerden
˜o, E.P., Lo
´pez, G., 1995. Los Ungulados nativos de Ame
´rica del Sur. In:
Alberdi,M.T.,Leone, G., Tonni,E.P.(Eds.), Evolucio
´nclima
´ticay biolo
´gicade la regio
´n
Pampeana durante los u
´ltimos cinco millones de an
˜os. Un ensayo de correlacio
´n
con el Mediterra
´neo occidental. Monografı
´as, vol.12. CSIC, Madrid, pp. 259–275.
Bondesio, P., Rabassa, J., Pascual, R., Vucetich, M.G., Scillato-Yane
´, G.J., 1980. La
Formacio
´n Collo
´n-Cura
´de Pilcaniyeu Viejo y sus alrededores (Rı
´o Negro,
Repu
´blica Argentina). Su antigu
¨edad y las condiciones ambientales segu
´nsu
distribucio
´n, su litogene
´sis y sus vertebrados. II Congreso Argentino de Pale-
ontologı
´a y Bioestratigrafı
´a y I Congreso Latinoamericano de Paleontologı
´a,
Buenos Aires, Actas 3, 85–99.
Bostelmann,E.T., Rinderknecht, A., Pereira, J., 2008. Primerosregistros de Glyptatelinae
Cuaternarios(Mammalia, Xenarthra)para el Cono Sur de Sudame
´rica. III Congreso
Latinoamericano de Paleontologı
´a de Vertebrados, Neuque
´n, resu
´menes, 29.
Burmeister, H., 1871. Descripcio
´n del ge
´nero Hoplophorus. Monografı
´a de los Glip-
todontes en el Museo Pu
´blico de Buenos Aires. Anales del Museo Pu
´blico de
Buenos Aires 2 (2), 157–226.
Cabrera, A., 1939. Sobre vertebrados fo
´siles del Plioceno de Adolfo Alsina. Revista
del Museo de La Plata (Nueva Serie) 2 (6), 3–35. La Plata.
Cabrera, A., 1944. Los Gliptodontoideos del Araucaniano de Catamarca. Revista del
Museo de La Plata (Nueva Serie) 3 (15), 5–76. La Plata.
Carlini, A.A., Scillato-Yane
´, G.J., 1999. Evolution of Quaternary Xenarthrans (Mam-
malia) of Argentina. In: Rabassa, J., Salemme, M. (Eds.), Quaternary of South
America and Antarctic Peninsula, vol. 12, pp. 149–175. Rotterdam.
Carlini, A.A., Vizcaı
´no, S.F., Scillato-Yane
´, G.J., 1997. Armored Xenarthrans: a unique
taxonomic and ecologic assemblage. In: Kay, R.F., Cifelli, R.L., Flynn, J.J.,
Madden, R.H. (Eds.), Vertebrate Paleontology of the Miocene Honda Group,
Republic of Colombia, vol. 13. Smithsonian Institution Press , pp. 213–226.
Carlini, A.A., Zurita, A.E., Gasparini, G.M., Noriega, J.I., 2004. Los mamı
´feros del Pleis-
toceno de la Mesopotamia argentina y su relacio
´n tanto con aquellos del Centro-
Norte de la Argentina, Paraguay, sur de Bolivia, como con los del sur de Brasil y
oeste de Uruguay: Paleobiogeografı
´a y Paleoambientes. Revista del Instituto
Superior de Correlacio
´nGeologı
´a(Miscela
´neas) 12,5–12. San Miguel de Tucuma
´n.
Carlini, A.A., Zurita, A.E., Aguilera, O., 2008a. North American glyptodontines
(Xenarthra, Mammalia) in the Upper Pleistocene of northern South America.
Palaeontologische Zeitschrift 82 (2), 139–152.
Carlini, A.A., Zurita, A.E., Scillato-Yane
´, G.J., Sa
´nchez, R., Aguilera, O., 2008b. New
glyptodont from Codore Formation (Pliocene), Falco
´n State, Venezuela, its
relationship with the Asterostemma problem and the Glyptodontinae. Palae-
ontologische Zeitschrift 82 (2), 125–138.
Castellanos, A., 1925. Descripcio
´n de un tubo caudal de Sclerocalyptus matthewi n. sp
(descubierto en el Pampeano inferior del valle de los Reartes, Sierras de
Co
´rdoba). Revista de la Universidad Nacional de Co
´rdoba 10–12, 1–54.
Castellanos, A., 1948. Um novo Esclerocaliptino. In: Notas Preliminares e Estudos,
vol. 47. Departamento Nacional da Produçao Mineral. Divisao de Geologı
´ae
Mineralogı
´a, Rio de Janeiro, pp. 1–13.
Castellanos, A., 1951. Un nuevo ge
´nero de esclerocalyptino (‘‘Isolinia’’) descubierto
en el Araucaniano del valle de los Reartes (sierras de Co
´rdoba). Revista de la
Asociacio
´n Geolo
´gica Argentina 6 (2), 95–100.
Castellanos, A., 1953. Descripcio
´n de restos de Paraglyptodon uquiensis n. sp. de
Uquı
´a (Senador Pe
´rez), Jujuy. Memorias del Museo de Entre Rı
´os 32 (Paleon-
tologı
´a), 1–32. Parana
´.
Cave, A.J.E., 1942. Museum Royal Collegue of Surgeons of England. Scientific Report
for the year 1940–1941, 3–16. London.
Chiesa, J., Lucero, N., Strasser, E., 2005. Sclerocalyptinae en la Depresio
´n de Conlara.
San Luis, Argentina, XXI Jornadas Argentinas de Paleontologı
´a de Vertebrados,
Plaza Huincul, resu
´menes, pp. 12–13.
Chiesa, J., Tognelli, G., Strasser, E., Prado, J., 2000. Paleomamı
´feros y paleoambiente
cuaternario del este de la Depresio
´n de Conlara, San Luis, Argentina. Ame-
ghiniana 37 (4), 22R.
Cione, A.L., Tonni, E.P., 1995. Bioestratigrafı
´a y cronologı
´a del Cenozoico de la regio
´n
Pampeana. In: Alberdi, M.T., Leone, G., Tonni, E.P. (Eds.), Evolucio
´n biolo
´gica y
clima
´tica de la regio
´n Pampeana durante los u
´ltimos cinco millones de an
˜os. Un
ensayo de correlacio
´n con el Mediterra
´neo occidental. Monografı
´as, vol. 12.
CSIC, Madrid, pp. 47–74.
Cione, A.L., Tonni, E.P., 1999. Biostratigraphy and chronological scale of upper-most
Cenozoic in the Pampean Area, Argentina. In: Rabassa, J., Salemme, M. (Eds.),
Quaternary of South America and Antarctic Peninsula, vol. 12, pp. 23–51.
Rotterdam.
Cione, A.L., Tonni, E.P., 2001. Correlation of Pliocene to Holocene southern South
American and European vertebrate-bearing units. Bolletino della Societa
´Pale-
ontologica Italiana 40 (2), 167–173.
Cione, A.L., Tonni, E.P., 2005. Bioestratigrafı
´a basada en mamı
´feros del Cenozoico
Superior de la provincia de Buenos Aires, Argentina. In: XVI Congreso Geolo
´gico
Argentino, Relatorio, 183–200. La Plata.
Cione, A., Tonni, E.P., Soibelzon, L., 2003. The broken zig-zag: Late Cenozoic large
mamad and tortoise extinction in South America. Revista del Museo Argentino
de Ciencias Naturales, Buenos Aires 5 (1), 1–19.
Clapperton, C., 1993. Nature of environmental changes in South America at the
Last Glacial Maxim um. Palaeogeograp hy, Palaeoclimatology, Palaeoecology
101, 189–208.
Coltorti, M., Abbazzi, L., Ferreti, M., Lacumin, P., Paredes Rı
´os, F., Pellegrini, M.,
Pieruccini, P., Rustioni, M., Tito, G., Rook, L., 2007. Last Glacial Mammals in South
America: a new scenario from the Tarija Basin (Bolivia). Naturwissenschaften
94, 288–299.
Downing, K.F., White, R., 1995. The cingulates (Xenarthra) of the Leisey Shell Pit
local fauna (Irvingtonian), Hillsborough County, Florida. Bulletin of the Florida
Museum of Natural History 37 II (12), 375–396.
Fernı
´cola, J.C., 2005. Ana
´lisis filogene
´tico de la familia Glyptodontidae Gray. Ame-
ghiniana 42 (4), 27R.
Fernı
´cola, J.C., Farin
˜a, R., Vizcaı
´no, S.F., 2002. Ana
´lisis filogene
´tico de la familia
Glyptodontidae Burmeister, 1879. In: I Congreso Latinoamericano de Paleon-
tologı
´a de Vertebrados, Santiago de Chile, resu
´menes, 32–33.
Fidalgo, F., Tonni, E.P., 1983. Geologı
´a y paleontologı
´a de los sedimentos encauzados
del Pleistoceno tardı
´o y Holoceno en Punta Hermengo y Arroyo Las Brusquitas
(Partido de General Alvarado y General Pueyrredo
´n, Provincia de Buenos Aires).
Ameghiniana 20 (3–4), 281–296.
Flynn, J., Kowallis, B.J., Nun
˜ez, C., Carranza-Castan
˜eda, O., Millar, W.E., Swisher, C.C.,
Lindsay, E., 2005. Geochronology of Hemphillian–Blancan Aged Strata, Guana-
juato, Mexico, and implications for timming of the Great American Biotic
Interchange. Journal of Geology 113, 287–307.
Frenguelli, J., 1928. Observaciones geolo
´gicas en la regio
´n costera de la provincia de
Buenos Aires. Anales de la Facultad de Ciencias de la Educacio
´n (Universidad
Nacional del Litoral) 2, 1–145.
Gervais, H., Ameghino, F., 1880. Los mamı
´feros fo
´siles de la Ame
´rica del Sur. Sabih e
Igon, Paris – Buenos Aires, p. 225.
Gilette, D.D., Ray, C.E., 1981. Glyptodonts of North America. Smithsonian Contri-
butions to Paleobiology 40, 1–251.
Giraudo, A., 1997. El Concepto de especie. Parte I. Natura Neotropicalis 28 (2),
161–169. Santa Fe.
Hevia, G.M., Romero, J.M., 1999. Sistema
´tica y Paleontologı
´a. In: Mele
´ndez, B. (Ed.),
Tratado de Paleontologı
´a, vol. 5. Consejo Superior de Investigaciones Cientı
´ficas,
Madrid, pp. 137–188.
Hoffstetter, R., 1958. Xenarthra. In: Piveteau, J. (Ed.), Traite
´de Pale
´ontologie, vol.
6(2), pp. 535–636.
Hoffstetter, R., 1968. N
˜uapua, un gisement de verte
´bre
´s ple
´istoce
`nes dans le Chaco
Bolivien. Bulletin du Muse
´um National d’Histoire Naturelle, Paris, (2
Se
´rie) 40
(4), 823–836.
International Commission on Zoological Nomenclature, 1999. International Code of
Zoological Nomenclature Adopted by the XX General Assembly of the Inter-
national Union of Biological Sciences. International Trust for Zoological
Nomenclature, London, pp. 1–306.
Iriondo, M., 1987. Geomorfologı
´a y Cuaternario de la provincia de Santa Fe
(Argentina). D‘ Orbignyana 4, 1–54. Corrientes.
Isla, F.I., Dondas, A., 2001. Facies fluviales del Pleistoceno de Mar del Plata,
Argentina. Revista de la Asociacio
´n Geolo
´gica Argentina 56 (3), 259–267.
Kraglievich, L., 1932. Nuevos apuntes para la geologı
´a y paleontologı
´a uruguayas.
Anales del Museo de Historia Natural de Montevideo 2 (3), 257–320.
Kraglievich, L., 1952. El perfil geolo
´gico de Chapalmalal y Miramar, Provincia de
Buenos Aires. Revista del Museo de Ciencias Naturales y Tradicionales de Mar
del Plata 1, 8–37.
Lydekker, R.,1887. Catalogue of the fossil Mammalia in the British Museum. Natural
History 5, 1–345. London.
Lydekker, R.,1894. Contribution to knowledge of the fossil vertebrates of Argentina.
2. The extinct Edentates of Argentina. Anales del Museo de La Plata (Paleon-
tologı
´a) 3, 1–118.
MacFadden, B., 2000. Middle Pleistocene climate change recorded in fossil mammal
teeth from Tarija, Bolivia, and upper limit of the Ensenadan Land-Mammal Age.
Quaternary Research 54, 121–131.
Marshall, L., Berta, A., Hoffstetter, R., Pascual, R., Reig, R., Bombin, M., Mones, A.,
1984. Mammals and Stratigraphy: Geochronology of the Continental Mammal-
Bearing Quaternary of South America. Palaeovertebrata, Me
´moire Extraordi-
naire, Montpellier, pp. 1–76.
McKenna, M.C., Bell, S.K., 1997. Classification of Mammals. Above the Species Level.
Columbia University Press, New York, pp. 1–631.
Menegaz, A.N., 2000. Los Came
´lidos y Ce
´rvidos del Cuaternario del Sector
Bonaerense de la Regio
´n Pampeana. Tesis Doctoral (Ine
´dita), Universidad
Nacional de La Plata, p. 240.
Menegaz, A.N., Ortiz Jaureguizar, E., 1995. Los Artioda
´ctilos. In: Alberdi, M.T.,
Leone, G., Tonni, E.P. (Eds.), Evolucio
´n biolo
´gica y clima
´tica de la regio
´n Pampeana
durante los u
´ltimos cinco millones de an
˜os. Un ensayo de correlacio
´n con el
Mediterra
´neo occidental. Museo Monografı
´as, vol. 12. CSIC, Madrid, pp. 311–337.
Mones, A., 1986. Paleovertebrata Sudamericana. Cata
´logo sistema
´tico de los verte-
brados fo
´siles de Ame
´rica del Sur. Parte I. Lista preliminar y Bibliografı
´a. Courier
Forschungsinstitut Senckenberg 82, 1–62.
Morgan, G.S., White, R.S., 2005. Miocene and Pliocene vertebrates from Arizona.
New Mexico Museum of Natural History and Science Bulletin 29, 115–136.
Nabel, P., 1993. The Brunhes–Matuyama boundary in Pleistocene sediments of
Buenos Aires province, Argentina. Quaternary International 17, 79–85.
Nabel, P., Camilion, M.C., Machado, G.A., Spiegelman, A.T., Mormeneo, L., 1993.
Magneto y litoestratigrafı
´a de los sedimentos pampeanos en los alrededores de
la ciudad de Baradero, provincia de Buenos Aires. Revista de la Asociacio
´n
Geolo
´gica Argentina 48 (3–4), 193–206.
Nabel, P.E., Cione, A.L., Tonni, E.P., 2000. Environmental changes in the Pampean
area of Argentina at the Matuyama–Brunhes (C1r-C1n) Chrons boundary.
Palaeogeography, Palaeoclimatology, Palaeoecology 162, 403–412.
Noriega, J.I., Carlini, A.A., Tonni, E.P., 2004. Vertebrados del Pleistoceno tardı
´odela
cuenca del arroyo Ensenada (Departamento Diamante, provincia de Entre Rı
´os,
A.E. Zurita et al. / Quaternary International 210 (2009) 82–92 91
Author's personal copy
Argentina). Revista del Instituto Superior de Correlacio
´n Geologı
´a (Miscela
´neas)
12, 71–76. San Miguel de Tucuma
´n.
Ojeda, R.A., Gonnet, J.M., Borghi, C.E., Giannoni, S.M., Campos, C.M., Dı
´az, G.B., 1996.
Ecological observations of the red vizcacha rat, Tymponoctomys barreae in
desert habitats of Argentina. Mastozoologı
´a Neotropical 3, 183–191.
Orgeira, M.J., 1987. Estudio paleomagne
´tico de sedimentos del Cenozoico tardı
´oen
la costa atla
´ntica bonaerense. Revista de la Asociacio
´n Geolo
´gica Argentina 42
(3–4), 362–376.
Owen, R., 1839. Description of a tooth and part of the skeleton of the Glyptodon,
a large quadruped of the Edentate order, to which belongs the tessellated bony
armour figured by Mr. Clift in his memoir on the remains of the Megatherium,
brought to England by Sir Woodbine Parish, F.G.S. Proceedings of the Geological
Society of London 3, 108–113.
Owen, R., 1845. Descriptive and illustrated catalogue of the fossil organic remains of
Mammalia and Aves contained in the Museum of the Royal College of Surgeons
of London, England, 1–391.
Parent, H., Polare, M., Lattuca, F., 2002. Estratigrafı
´a. In: Albert, O.A., Lattuca, F.,
Parent, H. (Eds.), Geologı
´a y Geomorfologı
´a de la Cuenca del Arroyo del Medio,
Provincia Santa Fe (Argentina), 72(1–2). Boletı
´n del Instituto de Fisiografı
´ay
Geologı
´a, Rosario, pp. 22–35.
Pascual, R., Ortega Hinojosa, E.J., Gondar, D.G., Tonni, E.P., 1966. Vertebrata. In:
Borrello, A.V. (Ed.), Paleontografı
´a Bonaerense. Vertebrata I. Comisio
´nde
Investigaciones Cientı
´ficas de la provincia de Buenos Aires, p. 202. Buenos Aires.
Paula Couto, C., 1957. So
ˆbre um gliptodonte do Brasil, vol. 165. Boletim Divisa
˜ode
Geologia e Mineralogia, Rio de Janeiro, pp. 1–37, Fig. 1–30.
Paula Couto, C., 1965. So
ˆbre a nomenclatura de dois gliptodontes, vol. 125. Div. Geol.
Min., D.N.P.M. Bol., Rio de Janeiro, pp. 1–6.
Paula Couto, C., 1979. Tratado de Paleomastozoologı
´a. Academia Brasileira de
Ciencias, Rio de Janeiro, p. 590.
Perea, D., 2005. Pseudoplohophorus absolutus n. sp. (Xenarthra, Glyptodontidae),
variabilidad en Sclerocalyptinae y redefinicio
´n de una biozona del Mioceno
Superior de Uruguay. Ameghiniana 42 (1), 175–190.
Politis, G.G., Gutierrez, M.A., 1998. Gliptodontes y cazadores-recolectores de la
regio
´n Pampeana (Argentina). Latin America Antiquity 9 (2), 111–134.
Prado, J.L., Alberdi, M.T., Azanza, B., Sanchez, B., 2001. Climate and changes in
mammal diversity during the late Pleistocene–Holocene in the Pampean region
(Argentina). Acta Palaeontologica Polonica 46 (2), 261–276.
Rabassa, J., Coronato, A.M., Salemme, M., 2005. Chronology of the Late Cenozoic Pata-
gonianglaciations and their correlationwith biostratigraphic units of the Pampean
region (Argentina). Journal of South American Earth Sciences 20 (1-2), 81–103.
Richter, J., 1911. Ueber Hoplophorus.Palaeontographica 57 (6), 257–283. plates,
22–23, Fig. 1-6.
Rinco
´n, A.D., 2006. Los Xenarthra Cingulata de Venezuela y Trinidad. In: IX Con-
greso Argentino de Paleontologı
´a y Bioestratigrafı
´a, resu
´menes, Co
´rdoba, 113.
Rovereto, C., 1914. Los estratos araucanos y sus fo
´siles. Anales del Museo Nacional
de Historia Natural de Buenos Aires, 1–250.
Rusconi, C., 1930. Observaciones geopaleontolo
´gicas en el sur de Villa Lugano
(Capital Federal). Physis X, 109–126.
Rusconi, C., 1936. Distribucio
´n de los vertebrados fo
´siles del Piso Ensenadense.
Boletı
´n de la Academia Nacional de Ciencias de Co
´rdoba 33, 183–215.
Scillato-Yane
´, G.J., 1977. Sur quelques Glyptodontidae nouveaux (Mammalia,
Edentata) du De
´se
´adien (Oligoce
´ne infe
´rieur) de Patagonie (Argentine). Bulletin
du Muse
´um National d
´Histoire Naturelle 487, 249–260.
Scillato-Yane
´, G.J., 1986. Los Xenarthra fo
´siles de Argentina (Mammalia, Edentata).
In: VII Congreso Argentino de Paleontologı
´a y Bioestratigrafı
´a, Mendoza, Actas,
151–155.
Scillato-Yane
´, G.J., Carlini, A.A., 1998a. Nuevos Xenarthra del Friasense (Mioceno
Medio) de Argentina. Stvdia Geologica Salmantisencia 34, 43–67.
Scillato-Yane
´, G.J., Carlini, A.A., 1998b. Un gigantesco gliptodonte en los alrededores
de la ciudad de La Plata. Museo 2 (11), 45–48. Facultad de Ciencias Naturales y
Museo de La Plata (UNLP).
Scillato-Yane
´, G.J., Carlini, A.A., Vizcaı
´no, S.F., Ortiz Jaureguizar, E., 1995. Los
Xenarthros. In: Alberdi, M.T., Leone, G., Tonni, E.P. (Eds.), Evolucio
´n biolo
´gica y
clima
´tica de la regio
´n Pampeana durante los u
´ltimos cinco millones de an
˜os. Un
ensayo de correlacio
´n con el Mediterra
´neo occidental. Monografı
´as, vol. 12.
CSIC, Madrid, pp. 183–209.
Scillato-Yane
´, G.J., Tonni, E.P., Carlini, A.A., Noriega, J.I., 1998. Nuevos Hallazgos de
Mamı
´feros del Cuaternario en el Arroyo Toropı
´, Corrientes, Argentina. Aspectos
Bioestratigra
´ficos, Paleoambientales y Paleozoogeograficos. In: VI Congreso
Latinoamericano de Geologı
´a y X Congreso Nacional de Geologı
´a Econo
´mica,
Actas 1, 263–268.
Scott, W.B., 1903–1904. Mammalia of Santa Cruz beds. I. Edentata. Reports of the
Princeton University Expeditions to Patagonia 1896–1899, 5: 1–226, plates.
1–35, Fig. 1–32 (1903b); 227–364, plates. 36–63, Fig. 33–38 (1904a). Princeton.
Soibelzon, E., 2005. Los mamı
´feros del Ensenadense (Pleistoceno inferior-medio)
del Este de la regio
´n Pampeana: asociaciones faunı
´sticas, bioestratigrafı
´ay
diversidad. Una aproximacio
´n a su estudio. In: XVI Congreso Geolo
´gico
Argentino, Actas, La Plata, 233–238.
Soibelzon, L.H., Tonni, E.P., Bond, M., 2005. The fossil record of South American
short-faced bears (Ursidae, Tremarctinae). Journal of South American Earth
Sciences 20, 105–113.
Soibelzon, E., Carlini, A.A., Tonni, E.P.,Soibelzon, H.L., 2006a. Chaetophractus vellerosus
(Mammalia: Dasypodidae) in the Ensenadan (Early-Middle Pleistocene) of the
southeastern Pampean region (Argentina). Palazoogeographical and paleo-
climatic aspects. Neues Jahrbuch fu
¨r Geologie und Palaontologie 12, 734–748.
Soibelzon, E., Tonni, E.P., Bidegain, J.C., 2008. Cronologı
´a, magnetoestratigrafı
´ay
caracterizacio
´n bioestratigra
´fica del Ensenadense (Pleistoceno inferior-medio)
en la ciudad de Buenos Aires, Argentina. Revista de la Asociacio
´n Geolo
´gica
Argentina 63 (3), 421–429.
Soibelzon, E., Zurita, A.E., Carlini, A.A., 2006b. Glyptodon munizi Ameghino (Mam-
malia, Cingulata, Glyptodontidae): redescripcio
´n y anatomı
´a. Ameghiniana 43
(2), 377–384.
Tonni, E.P., Cione, A.L., 1994. Los mamı
´feros y el clima en el Pleistoceno y Holoceno
de la provincia de Buenos Aires. Jornadas de Arqueologı
´a e Interdisciplinarias,
127–142. Buenos Aires: Programa de Estudios Prehisto
´ricos, Actas.
Tonni,E.P., Cione,A.L., 1995.Los mamı
´feros comoindicadores de cambios clima
´ticos en
el Cuaternariode la regio
´n pampeanade la Argentina. In: Argollo,J., Mourguiart,Ph
(Eds.), Cambios cuaternarios en Ame
´rica del Sur, pp. 319–326. La Paz.
Tonni, E.P., Fidalgo, F., 1979. Consideraciones sobre los cambios clima
´ticos durante el
Pleistoceno tardı
´o-reciente en la Provincia de Buenos Aires. Aspectos ecolo
´gicos
y zoogeogra
´ficos relacionados. Ameghiniana 15 (1–2), 235–253.
Tonni, E.P., Alberdi, M.T., Prado, J.L., Bargo, S., Cione, L.A., 1992. Changes of mammal
assemblage in the Pampean region (Argentina) and their relation with the Plio-
Pleistocene boundary. Palaeogeography, Palaeoclimatology, Palaeoecology 95,
179–194.
Tonni, E.P., Cione, A.L., Figini, A.J., 1999a. Predominance of arid climates indicated by
mammals in the Pampas of Argentina during the Late Pleistocene and Holo-
cene. Palaeogeography, Palaeoclimatology, Palaeoecology 147, 257–281.
Tonni, E.P., Nabel, P., Cione, A.L., Etchichury, M., Tofalo, R., Scillato-Yane
´, G.J., San
Cristo
´bal, J., Carlini, A.A., Vargas, D., 1999b. The Ensenada and Buenos Aires
formations (Pleistocene) in a quarry near La Plata, Argentina. Journal of South
America Earth Sciences 12, 273–291.
Tonni, E.P., Pardin
˜as, U.J., Verzi, D., Noriega, J.I., Scaglia, O., Dondas, A., 1998.
Microvertebrados pleistoce
´nicos del Sudeste de la Provincia de Buenos Aires
(Argentina): bioestratigrafı
´a y paleoambientes. V Jornadas Geolo
´gicas y
Geofı
´sicas Bonaerenses, Buenos Aires, Actas, 73–83.
Tonni, E.P., Scillato-Yane
´, G.J., 1997. Una nueva localidad con mamı
´feros pleistocenos
en el Norte de la Argentina. Aspectos paleozoogeogra
´ficos. In: VI Congreso da
Associacao Brasileira de Estudos do Quaternario e Reuniao sobre o Quaternario
da America do Sul, Anais, Curitiba, Brasil, 345–348.
Tonni, E.P., Huarte, R., Carbonari, J.E., Figini, A.J., 2003. New radiocarbon chronology
for the Guerrero Member of the Luja
´n Formation (Buenos Aires, Argentina):
palaeoclimatic significance. Quaternary International 109–110, 45–48.
Verzi, D.H., Tonni, E.P., Scaglia, O., San Cristo
´bal, J.O., 2002. The fossil record of the
desert-adapted South American rodent Tympanoctomys (Rodentia, Octodonti-
dae). Palaeoenvironmental and biogeographic significance. Palaeogeography,
Palaeoclimatology, Palaeoecology 179, 149–158.
Vinacci, E.L., 1939. Estudio comparativo del esqueleto apendicular de Glyptodontia.
Tesis Doctoral (ine
´dita) n
0024, Universidad Nacional de La Plata, Argentina,
102 pp. þ12 la
´ms.
Vizcaı
´no, S.F., Rinderknetcht, A., Czerwonogora, A., 2003. An enigmatic Cingulata
(Mammalia, Xenarthra) from the Late Miocene of Uruguay. Journal of Verte-
brate Paleontology 23 (4), 981–983.
Voglino, D., Pardin
˜as, U.F., 2005. Roedores sigmodontinos (Mammalia: Rodentia:
Cricetidae) y otros micromamı
´feros pleistoce
´nicos del norte de la provincia de
Buenos Aires (Argentina): reconstruccio
´n paleoambiental para el Ensenadense
cuspidal. Ameghiniana 42 (1), 143–158.
White, R.S., Morgan, G.S.., 2005. Arizona Blancan vertebrate faunas in regional
perspective. In: McCord, R.D. (Ed.), Vertebrate Palaeontology of Arizona, Mesa
Southwest Museum Bulletin vol. 11, 117–138.
Zurita, A.E., 2002. Nuevo gliptodonte (Mammalia, Glyptodontoidea) del Cuaternario
de la provincia de Chaco, Argentina. Ameghiniana 39 (2), 175–182.
Zurita, A.E., 2007. Sistema
´tica y evolucio
´n de los Hoplophorini (Xenarthra, Glyp-
todontidae, Hoplophorinae. Mioceno tardı
´o-Holoceno temprano). Importancia
bioestratigra
´fica, paleobiogeogra
´fica y paleoambiental. Tesis Doctoral (Ine
´dita),
Universidad Nacional de La Plata, p. 363.
Zurita, A.E., Carlini, A.A., Scillato-Yane
´, G.J., Tonni, E.P., 2004. Mamı
´feros extintos
del Cuaternario de la provincia del Chaco (Argentina) y su rela cio
´n con
aquellos del este de la regio
´n pampeana y de Chile. Revista Geolo
´gica de Chile
31 (1), 65–89.
Zurita, A.E., Min
˜o-Boilini, A.R., Soibelzon, E., Carlini, A.A., Paredes Rı
´os, F., 2009. The
diversity of Glyptodontidae (Xenarthra, Cingulata) in the Tarija Valley (Bolivia):
systematic, biostratigraphic and paleobiogeographic aspects of a particular
assemblage. Neues Jahrbuch fu
¨r Geologie und Palaontologie 251/52, 225–237.
Zurita, A.E., Carlini, A.A., Scillato-Yane
´, G.J., 2008. A new species of Neo-
sclerocalyptus Paula Couto, 1957 (Xenarthra, Glyptodontidae, Hoplophorinae)
from the middle Pleistocene of the Pampean region, Argentina. Geodiversitas
30 (4), 779–791.
Zurita, A.E., Scillato-Yane
´, G.J., Carlini, A.A., Alonso, R., 2002. Sclerocalyptus evidens
Ameghino 1889 (Mammalia, Xenarthra, Glyptodontidae) en el Pleistoceno de
Salta, Argentina. In: VIII Congreso Argentino de Paleontologı
´a y Bioestratigrafı
´a,
Corrientes, resu
´menes, 58.
Zurita, A.E., Scillato-Yane
´, G.J., Carlini, A.A., 2005. Palaeozoogeographic, biostrati-
graphic and systematic aspects of the genus Sclerocalyptus Ameghino 1891
(Xenarthra, Glyptodontidae) of Argentina. Journal of South American Earth
Sciences 20, 120–129.
Zurita, A.E., Scillato-Yane
´, G.J., Mones, A., 2007. Aspectos Nomenclaturales de la
Familia Glyptodontidae (Mammalia, Xenarthra): el caso de Sclerocalyptus
Ameghino, Hoplophorus Lund y la Tribu Sclerocalyptini. Ameghiniana 44 (1),
241–244.
A.E. Zurita et al. / Quaternary International 210 (2009) 82–9292
