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Introduction
Condors and vultures constitute the family
Vulturidae with five extant genera and seven
species, restricted to the Americas.
At least one extinct genus (Breagyps Miller and
Howard, 1938) and four extant genera (Gymnogyps,
Sarcoramphus, Cathartes, and Coragyps) are recorded
in the Pleistocene of North America, whereas two ex-
tinct genera (Geronogyps Campbell, 1979, and Win-
gegyps Alvarenga and Olson, 2004) and the five ex-
tant genera (Vultur, Gymnogyps, Sarcoramphus,
Cathartes, and Coragyps) are recorded for the
Pleistocene of South America (Miller, 1910; Howard,
1962; Brodkorb, 1964, Campbell, 1979; Tonni and
Noriega, 1998; Tambussi and Noriega, 1999;
Alvarenga and Olson, 2004).
Several authors claim that the fundamental differ-
ence between condors and vultures is the greater size
of the former, but this division is mainly based on cra-
nial osteology (Miller and Howard, 1938; Fisher, 1944;
Emslie, 1988; Alvarenga and Olson, 2004) apparently
with phylogenetic basis. The earliest records for this
family are in the Middle or Upper Eocene of France
(Mourer-Chauviré, 2002), whereas in South America
the earliest representative is Brasilogyps Alvarenga,
1985, from the Upper Oligocene or Lower Miocene of
southeast of Brazil. In North America, the earliest
records for this family date from the Pliocene, al-
though there are contradictory references for Eocene
records (Olson, 1985; Emslie, 1988; Mourer-Chauviré,
2002). It is possible that the family originated in the
Old World, to later become fixed only in the
Americas, especially in South America.
It is perfectly plausible to think that the diversity
of the Vulturidae could have been much greater dur-
AMEGHINIANA (Rev. Asoc. Paleontol. Argent.) - 45 (3): 613-618. Buenos Aires, 30-09-2008 ISSN 0002-7014
©Asociación Paleontológica Argentina AMGHB2-0002-7014/08$00.00+.50
1Museu de História Natural de Taubaté. Rua Juvenal Dias de
Carvalho, 111. CEP 12070-640 Taubaté, SP, Brazil.
halvarenga@uol.com.br; rmigotto@yahoo.com.br
2Departamento de Zoologia, Instituto de Biociências,
Universidade de São Paulo. C.P. 11294, CEP 05422-970, São Paulo,
SP, Brazil. grrbrito@yahoo.com.br; ehofling@usp.br
3Laboratório de Estudos Evolutivos Humanos, Departamento de
Genética e Biologia Evolutiva, Instituto de Biociências,
Universidade de São Paulo, C.P. 11461, CEP 05422-970 São Paulo,
SP, Brazil. alexhubbe@yahoo.com
Pleistovultur nevesi gen. et sp. nov. (Aves: Vulturidae) and
the diversity of condors and vultures in the South
American Pleistocene
Herculano ALVARENGA1, Guilherme R.R. BRITO2, Rafael MIGOTTO1;2, Alex HUBBE3y
Elizabeth HÖFLING2
Key words. Vulturidae. Pleistocene. South America. Condor. Pleistovultur nevesi. Caves.
Palabras clave. Vulturidae. Pleistoceno. América del Sur. Condor. Pleistovultur nevesi. Cavernas.
Abstract. Pleistovultur nevesi, a new genus and species of a large Vulturidae is described from the Upper
Pleistocene or beginning of the Holocene, based on a complete and well preserved right tibiotarsus from
the Cuvieri cave deposits, one of the hundreds of caves of the Lagoa Santa region in Minas Gerais state,
Brazil. Also described here is a distal half of a left tibiotarsus from Gruta dos Brejões, a cave in Morro do
Chapéu, Bahia state, Brazil, which probably belongs to another not-yet-known genus and species of the
Vulturidae. The described material, added to prior literature, is proof of the appreciable diversity of the
Vulturidae during the Pleistocene in South America.
Resumen. PLEISTOVULTUR NEVESI GEN. ET SP. NOV. (AVES: VULTURIDAE) Y LA DIVERSIDAD DE CÓNDORES Y BUITRES
EN EL PLEISTOCENO DE AMÉRICA DEL SUR. Se describe un nuevo género y especie de Vulturidae de gran
tamaño para el Pleistoceno Tardío o comienzos del Holoceno, en base a un completo y bien preservado
tibiotarso procedente de los depósitos de la caverna Cuvieri, una de los centenares de cavernas presentes
en la región de la Lagoa Santa en el estado de Minas Gerais, Brasil. Se describe asimismo la mitad distal
de un tibiotarso izquierdo de otro gran Vulturidae de Gruta dos Brejoes, otra caverna en Morro do
Chapéu, Estado da Bahia, Brasil, que probablemente pertenezca a otro aún desconocido género y especie.
El material descrito, sumado al registro édito previo, es una prueba de la apreciable diversidad de vultúri-
dos durante el Pleistoceno de América del Sur.
H. Alvarenga, G.R.R. Brito, R. Migotto, A. Hubbe y E. Höfling
614
ing the Pleistocene in South America in the context
of its megafauna, as occurs in Africa currently with
Old Word vultures (Accipitridae). In corroboration
of this hypothesis, we herein describe a new genus
and species of a large Vulturidae from the Upper
Pleistocene or Lower Holocene of Brazil, based on a
complete right tibiotarsus, collected in a cave of the
Matozinhos region, Minas Gerais state, Brazil (figure
1). We also describe and compare another incomplete
tibiotarsus from the Gruta dos Brejões, in Morro do
Chapéu, Bahia state, reinforcing the diversity of these
birds during the South American Pleistocene.
Material and methods
One of the studied specimen is a complete right
tibiotarsus, from the Cuvieri cave, Matozinhos,
Minas Gerais State, Brazil, initially broken and cov-
ered by carbonatic crust; it was previously treated
with acetic acid and afterwards mechanically cleaned
under a stereomicroscope, and then restored (figure
2.2). The bone is relatively well preserved, apparent-
ly from an adult individual, based on the bone sur-
face texture when observed through the microscope,
without any indication of a thick periosteum. Erosion
is present on the edges of the proximal joint surfaces
and on the lateral and medial edges of the condyles,
which compromise a sharp precision in the measure-
ments of the proximal and distal width. It is housed
in the Museu de História Natural de Taubaté (MH-
NT-VT-5238). It was compared with homologous
bones of extant skeletons of all species of the South
American Vulturidae (appendix 1), especially with
the genera Vultur and Sarcoramphus because of their
similar size. Comparisons to Gymnogyps were done
through high resolution photographs, from several
views, taken by one of the authors (H.A.) in the Los
Angeles County Museum, California (specimen B
1372). Comparisons with Breagyps were done only
from the descriptions and illustrations of Howard
(1974) whereas comparisons with the genus Gerono-
gyps were undertaken by means of the photographs
and descriptions of Campbell (1979). We also exam-
ined and compared a slightly damaged distal half of
a left tibiotarsus, that was collected by Castor
Cartelle in the cave of Brejões, Municipality of Morro
do Chapéu, Bahia, Brazil (figure 1), housed as MCL-
1795 in the Museu de Ciências of the Pontificia
Universidade Católica, Minas Gerais, Brazil; this ma-
terial came from the same locality as that of the type
of Wingegyps cartellei (Alvarenga and Olson, 2004).
The terminology used herein is mainly according to
Howard (1929) and, in some cases, Baumel and
Witmer (1993). The measurements were done with
0.1mm/precision Mitutoyo calipers.
Systematic paleontology
Family VULTURIDAE
Pleistovultur gen. nov.
Type species. Pleistovultur nevesi sp. nov.
Diagnosis. Tibiotarsus with an almost oval-shaped
articular surface for the head of the fibula, similar to
Sarcoramphus (different from that of Vultur distally
expanded); there is a crest between this joint and the
fibular crest (similar in Breagyps, absent in Vultur,
and tenuous in Sarcoramphus). The distal part of the
outer cnemial crest is sharp (similar in Vultur,
Gymnogyps, and Breagyps ) and reaches the level of
the fibular crest whereas in Sarcoramphus it is thick,
rounded, and shorter. The distal opening of the ten-
dinal groove is rounded (similar in Vultur, Gymno-
gyps, and Breagyps; oval in Sarcoramphus and Gerono-
gyps). The lateral condyle in anterior view is very
high and its axis is parallel to the diaphysis (it is
much shorter in Sarcoramphus; in Vultur, Gymnogyps,
and Breagyps the lateral condyle is wider and oblique
to the lateral side of the diaphysis). There is no fora-
AMEGHINIANA 45 (3), 2008
Figure 1. Map showing the locality of (A) Matozinhos (Gruta
Cuvieri), MG, Brazil, (19°28’36’’S, 44°00’41’’W), the type locality
of Pleistovultur nevesi and (B) Gruta dos Brejões in Morro do
chapéu, BA, Brazil, (11º00’30’’S, 41°26’07’’W), the type locality of
Wingegyps cartellei and also the left tibiotarsus MCL-A- 1795 /
mapa indicando la localidad de (A) Matozinhos (Gruta Cuvieri), MG,
Brasil (19°28´36´´S, 44°00´41´´W), localidad tipo de Pleistovultur
nevesi y (B) Gruta dos Brejoes en Morro do Chapéu, BA, Brasil
(11°00’30’’S, 41°26’07’’W), localidad tipo de Wingegyps cartellei y
también del tibiotarso izquierdo MCL-A-1795.
Pleistovultur nevesi gen. et sp. nov. (Aves:Vulturidae) 615
AMEGHINIANA 45 (3), 2008
Figure 2. The right tibiotarsus of Vultur gryphus MHNT 591; Pleistovultur nevesi gen. et sp. nov. holotype MHNT-VT-5238; Sarcoram-
phus papa MHNT 1787 respectively, in anterior view (1, 2 and 3); the proximal end in medial view (4, 5 and 6); the distal end in anterior
view (7, 8 y 9) / tibiotarso derecho de Vultur gryphus MHNT 591; Pleistovultur nevesi gen. et sp. nov. holotipo MHNT-VT-5238; y Sarco-
ramphus papa MHNT 1787 respectivamente en vista anterior (1, 2 y 3); extremidad proximal en vista lateral (4, 5 y 6); y extremidad distal en vista
anterior (7, 8 y 9). Scale bar / escala 2 cm.
H. Alvarenga, G.R.R. Brito, R. Migotto, A. Hubbe y E. Höfling
616
men proximal to the lateral condyle (in anterior view,
and at the same level of the tendinal bridge); all
Vulturidae show a foramen at this place, but it is
more conspicuous in Vultur. In lateral view, the late-
ral condyle of Pleistovultur appears to be more round-
ed than in Vultur and Sarcoramphus, however impor-
tant erosion at its edges may give a false impression.
Etymology. Pleisto from Pleistocene + vultur .
Pleistovultur nevesi sp. nov.
Figures 2.2, 5, 8
Holotype. A complete right tibiotarsus; MHNT-VT-5238 (figure
2.2).
Type locality. Brazil, Minas Gerais State, Muni-
cipality of Matozinhos, Gruta Cuvieri (19º28’36’’S,
44º00’41’’W), elevation ca. 812m (figure 1).
Horizon and age. The holotype of Pleistovultur was
discovered inside the Gruta Cuvieri, Municipality of
Matozinhos, Minas Gerais, Brazil, one of the hun-
dreds of caves in the region north/northeast of the
city of Belo Horizonte, many of which were explored
and studied in the first half of the 19th Century by
the Dane Peter Wilhelm Lund. Unfortunately, due to
previous work done in this cave, the taphonomic
conditions of the tibiotarsus holotype of Pleistovultur
nevesi could not be duly defined, thus making it dif-
ficult to precisely determine its age. Nevertheless, the
knowledge of the associated fauna, present in sever-
al adjacent caves and previously dated (Laming-
Emperaire et al., 1975; Cartelle, 1999), permits at-
tributing an Upper Pleistocene or Early Holocene age
for this entire fauna, including Pleistovultur. In the
same Cuvieri cave, Neves and Pilo (2003) dated a
ground sloth Scelidodon cuvieri, and obtained a radio-
carbon age of 9.990 + 40ybp; also, from the same
cave, a second specimen of Scelidodon was dated of
12.510 ( 70ybp (Neves, pers. com.).
Etymology. Nevesi is in honor to Walter Neves, an-
thropologist from Laboratório de Estudos Evolutivos
Humanos of the Departamento de Genética e
Biologia Evolutiva, Instituto de Biociências, Uni-
versidade de São Paulo, responsible for collecting
and forwarding the material for our study.
Diagnosis. The same for the genus.
Description. A large Vulturidae with a tibiotarsus about
25% larger than that of Sarcoramphus papa and about
11% smaller than that of Vultur gryphus (figure 2.1, 2.2
and 2.3). Measurements of the holotype in table 1.
Vulturidae gen. et sp. indet.
Figure 3
A distal half of a left tibiotarsus (MCL-A-1795),
from the Gruta dos Brejões, Bahia, Brazil (figure 3)
(11°00’30’’S, 41°26’07’’W), the same locality of the type
specimen of Wingegyps cartellei (Alvarenga and Olson,
2004), also was studied and compared; unfortunately
the condyles are quite damaged, thus prejudicing a
better diagnosis. The very wide supratendinal bridge,
the oval distal opening of the tendinal groove, and the
very straight line of the medial border of the linea ex-
AMEGHINIANA 45 (3), 2008
Table 1. Measurements of the tibiotarsus of Pleistovultur nevesi gen. et sp. nov. compared to Pleistocene and extant large vultures (mm)
/ medidas del tibiotarso de Pleistovultur nevesi gen. et sp. nov. comparado con otros grandes buitres del Pleistoceno y reciente (mm).
Pleistovultur nevesi gen. et sp. nov. (Aves:Vulturidae) 617
tensoria, exclude this specimen from Pleistovultur,
Vultur, Gymnogyps, Breagyps, and Sarcoramphus, even
though by size it is very near to Pleistovultur. When
compared with the illustration of Geronogyps (in
Campbell, 1979), the supratendinal bridge and the
proximal and distal openings are very different. So,
we believe that this vulture cannot be assigned to any
known genus of Vulturidae; however, better material is
needed in order to merit a new name.
Discussion
During the Pleistocene, South America possessed
a rich mammalian megafauna, comparable to that
living in Africa today. Carrion-eating birds, repre-
sented by the family Vulturidae, analogous in the
food pyramid to the Old World vultures (Accipi-
tridae), almost certainly needed to feed on large car-
casses. Thus, the disappearance of the megafauna
certainly leads to extinction for many of the
Vulturidae. We believe it to be absolutely foreseeable
that there were an appreciable number of genera and
species of the Vulturidae in the South American
Pleistocene, which appears to be confirmed by recent
reports. Campbell (1979) described the presence of
the genera Gymnogyps and Geronogyps on the coast of
Peru, and also reported another possible of the genus
Sarcoramphus (Sarcoramphus? fisheri). Tonni y Noriega
(1998) and Tambussi and Noriega (1999) described
the presence of Geronogyps and Vultur gryphus from
the Pleistocene of the south of Buenos Aires
Province. Alvarenga (1998) identified the presence of
Vultur gryphus in the Pleistocene/Holocene from the
caves of Minas Gerais, Brazil. Alvarenga and Olson
(2004) described Wingegyps, a small condor of the
Pleistocene/Holocene from caves in Bahia and Minas
Gerais, Brazil. The tibiotarsus MCL-A-1795 certainly
represents an additional genus, which awaits better
material in order to be described. Winge (1888), who
studied numerous birds from caves of the same re-
gion as Pleistovultur, described (from Lapa Escri-
vania) fragments of humerus, ulna and coracoid of “a
much bigger Vulturidae than Gypargus papa (=
Sarcoramphus papa)” besides other bones attributed to
the extant Sarcoramphus papa from the Lapa do Bau.
These bones belonging to a Vulturidae larger than
Sarcoramphus, have been recently cleaned and re-
prepared by Olson and Emslie (pers. com.); unfortu-
nately they do not present structures for a good tax-
onomic diagnosis; it is possible that this material can
be attributed to Pleistovultur. Oluf Winge also per-
ceived the distinction of the fossil specimens that
Alvarenga and Olson (2004) subsequently described
as Wingegyps cartellei. The general panorama of vul-
tures from the South American Pleistocene now
seems to begin to be understood.
Acknowledgements
We are grateful to W. Neves (IB-USP) for making the fossil
tibiotarsus used in this work available for our study; to C. Cartelle
(PUC de Belo Horizonte) for letting us compare the specimen
MCL-A 1795; to Felipe Curcio (IB-USP) for his help on Fig. 1; to
Storrs L. Olson (Smithsonian Institution, Washington DC) and
Steven Emslie (UNC) for important personal communications; to
J.I. Noriega (CICyTTP-CONICET, Diamante) and C. Tambussi
(D.P.V. Museo de La Plata, Argentina) for some important criti-
cism and corrections in the manuscript. Our special gratitude to
Walter Hartwig from Touro University, Ca, for revision of the
manuscript and English corrections. We are also grateful to the
Conselho Nacional de Pesquisas e Desenvolvimento (CNPq) for
the scholarships of R.M. (130053/2006-6) and G.R.B. (140359/2004-
4) and the grant of E.H. (303926/85-6-RV), and also to the
Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPE-
SP) for the scholarship of A.H. (06/51406-1) and for the research
grant to Walter Neves (04/013216).
References
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AMEGHINIANA 45 (3), 2008
Figure 3. The distal half of a left tibiotarsus of a Vulturidae gen et
sp. indet (MCL-A-1795), coated with ammonium chloride, from
Morro do Chapéu, BA, Brazil, in anterior view / mitad distal de
tibiotarso izquierdo Vulturidae gen et sp. indet (MCL-A-1795), cubier-
to con cloruro de amonio, proveniente de Morro do Chapéu, BA, Brasil,
en vista anterior. Scale bar / Escala 2 cm.
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Recibido: 30 de abril de 2007.
Aceptado: 30 de julio de 2008.
Appendix 1. List of skeletons of extant Vulturidae used to com-
pare with Pleistovultur nevesi gen. et sp. nov.: MHNT= Museu de
História Natural de Taubaté; AZ= Collection of birds of the
Departamento de Zoologia, Instituto de Biociências, Universidade
de São Paulo / lista de los esqueletos de buitres vivientes que se usaron
para comparar con Pleistovultur nevesi gen. et sp. nov.: MHNT=
Museu de História Natural de Taubaté; AZ= Collection of birds of the
Departamento de Zoologia, Instituto de Biociências, Universidade de São
Paulo.
Vultur gryphus: two skeletons: MHNT- 591 and AZ-579.
Sarcoramphus papa: four skeletons: MHNT-775, 804, 903 and 1787.
Cathartes melambrotus: one skeleton: MHNT-1213
Cathartes aura: three skeletons: MHNT-97, 790 and 794
Cathartes burrovianus: one skeleton: MHNT-714
Coragyps atratus: seven skeletons: MHNT- 46, 285, 625, 750, 786,
985 and 1264.
AMEGHINIANA 45 (3), 2008
