Historical Biology

An International Journal of Paleobiology

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A new fossil condor (Aves, Cathartiformes)

from the Early Pliocene of Catamarca province,
Argentina

Federico J. Degrange, Ricardo A. Bonini, Sergio M. Georgieff & Lucía M.

Ibañez

To cite this article: Federico J. Degrange, Ricardo A. Bonini, Sergio M. Georgieff & Lucía M.
Ibañez (06 Dec 2023): A new fossil condor (Aves, Cathartiformes) from the Early Pliocene of
Catamarca province, Argentina, Historical Biology, DOI: 10.1080/08912963.2023.2288612

To link to this article: https://doi.org/10.1080/08912963.2023.2288612

Published online: 06 Dec 2023.

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HISTORICAL BIOLOGY
https://doi.org/10.1080/08912963.2023.2288612

A new fossil condor (Aves, Cathartiformes) from the Early Pliocene of Catamarca
province, Argentina
a b c,d c
Federico J. Degrange , Ricardo A. Bonini , Sergio M. Georgieff and Lucía M. Ibañez
a
Centro de Investigaciones en Ciencias de la Tierra (CICTERRA), Universidad Nacional de Córdoba, CONICET, Córdoba, Argentina; bINCUAPA CONICET,
Universidad Nacional del Centro de la provincia de Buenos Aires, Olavarría, Argentina; cIESGLO, Universidad Nacional de Tucumán, San Miguel de
Tucumán, Argentina; dCONICET – CCT NOA Sur, Argentina

ABSTRACT ARTICLE HISTORY

Condors are New World vultures (Cathartiformes, Cathartidae) with broad wings and short tails. The fossil Received 23 June 2023
record of condors (Aves, Cathartiformes) in South America is scarce, and particularly in Argentina most of the Accepted 23 November 2023
remains assigned to condors have been debated. Here, we present a new fossil species of Cathartidae found KEYWORDS
in the Early Pliocene of Catamarca province, Argentina, that belongs to the extant genus Vultur. The new Cathartids; Vultur; Mio-
species, Vultur messii sp. nov. is considered to be the only valid fossil species of this iconic genus of birds. The pliocene; Catamarca;
presence of a fossil Vultur species, recovered from deposits older than 4.8 Ma, coincides with the rise of the Andalhuala Formation
pampean hills and the Andes, and the beginning of the regional aridification. The existence of incipient
mountain chains may also indicate that the centre of origin of this genus is linked to the Andes mountain,
which in turn indicates an earlier arrival and diversification of the condors in South America.

Introduction Degrange et al. 2021; Jones et al. 2021; Degrange 2022). This enrich­
ment in species diversity, especially by the end of the Cenozoic, has
Fossil bird remains from Catamarca province are, in general, scarce.
been related to the high diversity in the megafauna with the condors as
With the exception of the well-represented Phorusrhacidae
the most frequent representatives in the Cathartidae fossil record of
(Rovereto 1914; Patterson and Kraglievich 1960; Vezzosi and
South America. Despite this past diversity, only two species remain
Noriega 2017), most birds from this region are known only from
today: the California condor Gymnogyps californianus that inhabits
fragmentary remains. To date, only three non-phorusrhacid taxa
open grassland, woodland regions and coastlines of southwestern
have been recorded. These taxa include the accipitrid Geranoaetus
U.S.A. and northwestern Mexico; and the Andean condor Vultur
sp., the teratornithid Argentavis magnificens Campbell and Tonni
gryphus, that is distributed along western South America, mainly in
1980, and the basal palaeolodid Megapaleolodus peiranoi (Agnolín
high mountains.
2009) (Campbell 1995; Agnolín 2006, 2009). All of these taxa come
The Andean condor is one of the most recognisable birds and
from the Late Miocene-Early Pliocene Andalhuala Formation
certainly one of the most iconic birds of South America, being of
(Tambussi et al. 2023), and represent three different avian guilds:
great significance for several indigenous tribes. Here, we present
aerial predator, scavenger, and wader, which are added to the
terrestrial predator avian guild represented by the phorusrhacids. a new record of a new condor species, coming from the Andean
An additional species coming from the same Formation but from region. Only two remains of Cathartidae have been recorded for the
Tucumán province corresponds to the terrestrial rheid Argentinean Andean region: Dryornis hatcheri from the Santa Cruz
Ophistodactylus kirchneri (Noriega et al. 2017). Formation in western Santa Cruz province, and a fragmentary
Cathartidae (or ‘New World vultures’) are a group of birds distal humerus recorded from the Toro Negro Formation in La
characterised by a soaring flight and carrion-based trophic habits Rioja province which was considered to be related to Dryornis
(Hertel 1994) that occur throughout North and South America. (Rodríguez Brizuela 2004; Degrange et al. 2021; Degrange 2022).
Although there are some osteological differences, their general The Cathartiformes described here stems from San Fernando Norte
morphology has been considered conservative. Based on general locality, 40 km north of Belén city, in the homonymous department,
external morphology, two morphotypes have been recognised: the Catamarca province. The remains were unearthed from Andalhuala
condors and the vultures (sensu Hertel 1992 and Stuchhi et al. Formation (Late Miocene-Early Pliocene) (Figure 1).
2015). Condors are larger and have broad wings and short tails,
while vultures are smaller birds with short wings and rounded long Materials and methods
tails.
Cathartidae has a very abundant fossil record, reaching its highest Unless indicated otherwise, osteological terminology follows
diversity in the Pleistocene, with most of the species having extremely Baumel and Witmer (1993). Myological terminology follows
large geographic ranges (Noriega and Areta 2005; Noriega and Tonni Vanden Berge and Zweers (1993).
2007; Degrange et al. 2021). The knowledge about the fossil record of Institutional Abbreviations: AMNH, American Museum of
this group in South America has improved in the past years (e.g. Natural History, Ornithology Collection, New York, U.S.A..; CIT-O,

CONTACT Federico J. Degrange fjdino@gmail.com Centro de Investigaciones en Ciencias de la Tierra (CICTERRA), Universidad Nacional de Córdoba, CONICET, Av.
Vélez Sársfield 1611, Córdoba X5016GCA, Argentina
© 2023 Informa UK Limited, trading as Taylor & Francis Group

Published online 06 Dec 2023

2 F. J. DEGRANGE ET AL.

Figure 1. Geographic and stratigraphic provenance of Vultur messii sp. nov. A, South America map showing the location of Catamarca province, Argentina; B, location map
showing the outcrops of Andalhuala Formation from which the holotype of V. messii sp. nov. comes, and its relation with two dated tuffs in this area; C, stratigraphic section
outcropping in San Fernando Norte locality showing the stratigraphic position of the fossil-bearing level (indicated with a star).

Colección Osteológica de Aves of the Centro de Investigaciones en fossiliferous levels of the Andalhuala Formation (Bonini et al.
Ciencias de la Tierra (CICTERRA), Córdoba, Argentina; MCH, 2017). The fossil location corresponds to deposits older than
Museo Arqueológico ‘Condor Huasi’, Belén, Argentina; OUVC, 4.8 Ma.
Ohio University Vertebrate Collections, Athens, U.S.A.; USNM, The fossil was collected in reddish brown mudstones between
Smithsonian National Museum of Natural History, Washington D. a 5 YR 7/4 siltstone, strike and dip 125°/10° and magnetic suscept­
C., U.S.A.. ibility of 1.64 (Sample 28), and a 5 YR 6/4 siltstone, with a magnetic
Material examined: Cathartes aura CIT-O134, OUVC 10,285; susceptibility of 2.37 (Sample 29). The paleoenvironment corre­
Cathartes burrovianus AMNH 1264; Coragyps atratus CIT-O344; sponds to alluvial floodplains close to the main channel-belt
Gymnogyps californianus AMNH 1937, 32152, USNM 13,823, (Bonini et al. 2017).
492447; Sarcoramphus papa AMNH 1893, 2136, 3901, 3902, 3903; The mineralogical composition of the clay-sized fraction, deter­
Vultur gryphus CIT-O263, USNM 18,485. Observations of mined by X-ray diffraction, corresponds to interstratified illite-
Geronogyps reliquus and Breagyps clarki were made from the origi­ smectite, poor crystallinity illite, chlorite and quartz. The smectite
nal figures and descriptions in the published literature (see Miller is mainly represented by beidellite, a tetrite (Moore and RC 1997),
1910, Howard 1974; Campbell 1979). whose layer charge cannot be neutralised by the Li+ used when
applying the Greene-Kelly (1955) method.
Illite-smectite interlayers are the most common interlayers and
Geology, age and paleoenvironmental settings are widely distributed. The presence of beidellite is rare, except in
The remains of the new Cathartiformes came from continental edaphic environments, according to Brindley (1980), so we can
sedimentary rocks of the Andalhuala Formation outcropping in infer soil development for these levels. However, the presence of
San Fernando Norte, Villavil-Quillay Basin (VQB) into the isolated illite may indicate that a mesodiagenesis stage has been
Northwestern Pampean Range geological province (Georgieff reached. Chlorite and quartz would be inherited materials.
et al. 2017). Two tuff outcroppings in Puerta de Corral Quemado, Majority elements were determined to calculate the chemical
in the VQB, constrained this unit between ca. 7.14 and 3.66 Ma Late alteration index (CIA), obtaining values of 60.76 for Sample 28 and
Miocene (Messinian) to Early Pliocene (Zanclean) (Latorre et al. 61.60 for Sample 29. Both correspond to intermediate degrees of
1997). In San Fernando, two tuffs dated at 4.72 ± 0.08 Ma and 4.79 alteration (Nesbitt and Yong 1982), indicating semi-arid to tem­
± 0.15 Ma, indicated Early Pliocene (Zanclean) age for the plate climatic conditions.
 HISTORICAL BIOLOGY 3

Systematic palaeontology Gymnogyps californianus, however this opening is much larger in

the fossil species, and similar to Geronogyps and Breagyps. However,
Class Aves Linnaeus, 1758
in Breagyps, the foramen is more proximally located, while in
Subclass Neognathae Pycraft, 1900
Geronogyps is more distally located, and in Gymnogyps, there is
Order Cathartiformes Coues, 1884
a deep furrow proximal to the foramen. Medially to this foramen,
Family Cathartidae Lafresnaye, 1839
Genus Vultur Linnaeus, 1758 there is a marked curved ridge that bounds the foramen. This ridge
is straighter in Vultur, Coragyps, and Cathartes. When viewed
proximally, medially to the crista trochanteris, a marked protuber­
Type species
ance is located, a feature observed only in Vultur among cathartids
Vultur gryphus Linnaeus, 1758
(this protuberance, although present is much less developed and
barely noticeable in Gymnogyps). The crista trochanteris is mark­
Included species
edly bended medially, similar to Breagyps and Geronogyps. The
Type species and V. messi sp. nov.
caudal angle between the lateral margin of the crista trochanteris
and the caudal aspect of the facies articularis antitrochanterica is
Vultur messii sp. nov. Figure 2 obtuse, and larger than that of Vultur (much smaller, and acute in
Cathartes). The scar for the insertion of the m. iliotrochanteris
caudalis is narrow and poorly marked (contrary to Vultur). The
Diagnosis impressio obturatoriae has a marked oval shape. The scar of the
insertion of the m. iliofemoralis internus has an oval shape, very
MCH 363 is assigned to genus Vultur (a genus that until the similar to that of Vultur. The linea intermuscularis cranialis is
discovery of the new material was monotypic) based on the pre­ poorly marked.
sence of a single foramen and a marked protuberance located The distal area of the femur is more badly preserved, heavily
medially to the crista trochanteris, an impressio lig. cruciati crania­ crushed and lacking a portion of the condylus medialis. The fossa
lis widely extended and a condylus lateralis that gradually transi­ poplitea is deep and wide (shallow in Coragyps and Gymnogyps),
tions towards the diaphysis. It differs from Vultur gryphus in the with at least two noticeable large pneumatic foramina. This fossa is
following features: the presence of a much larger unique foramen, bounded laterally by the condylus lateralis, which when viewed
a caudal deep pit bounded cranially by the narrower facies articu­ laterally has a gradual transition towards the diaphysis as in
laris antitrochanterica, a markedly medially deflected crista tro­ Vultur (abrupt transition in Coragyps and Cathartes). The scar of
chanteris, a rounded and markedly large scar of origin of origin of the m. gastrocnemius pars intermedia is rounded and large
the m. gastrocnemius pars intermedia, an impressio (smaller and oval in Vultur). The impression lig. cruciati cranialis is
ansae m. iliofibularis facing caudally, and a wide and shallow sulcus very wide and extended, as in Vultur (shallow and narrow in the
patellaris. smaller Cathartes and Coragyps). The
tuberculum m. gastrocnemialis lateralis is low, large, with a heart-
Holotype shaped scar (oval in Vultur, rounded in Coragyps and Cathartes).
The impressio ansae m. iliofibularis is small and oval, distally
MCH 363, right femur lacking part of the diaphysis. located in the aforementioned tuberculum, and faces caudally (cau­
dolaterally in V. gryphus, laterally in Geronogyps and Breagyps). As
Etymology in Vultur, the linea intermuscularis caudalis extends distally later­
ally on the shaft, bounding a scar located right proximally over the
Honouring the Argentinean football player Lionel Andrés Messi, crista lateralis sulci patellaris (laterally located in Cathartes and
one of the best of our times, who, being captain and best player, Coragyps). This crista is stout and wide, bounding laterally a wide
together with an outstanding team won the FIFA World Cup in but shallow sulcus patellaris (narrow in Geronogyps and deeper in
Qatar 2022. Vultur gryphus, in Gymnogyps it is wide and deep, and present
a quadrangular contour when viewed distally). The impressio lig.
Procedence collateralis lateralis is shallow. The fovea tendinis m. tibialis crania­
lis is shallow as in Cathartes (deep in Vultur). The sulcus inter­
Andalhuala Formation (Late Miocene-Early Pliocene). S27°17’34.2’ condylaris is shallow (deep in Gymnogyps).
W66°54’40.4’, San Fernando Norte, Belén Department, Catamarca
province (Figure 1). The fossil comes from older deposits than 4.8
Ma, corresponding to the Early Pliocene. Discussion and conclusions
The stout shape of the femur, along with the rounded and modestly
cranially projected crista trochanteris, with a pneumatic foramen
Description
medially located allow us to assign MCH 363 to Cathartidae.
The caput femoris is stout, slightly constricted (not constricted in Among them, the fossil record of condors in South America is
Vultur and Cathartes) with a marked and deep fovea lig. capitis. As particularly rich. Apart from Geronogyps reliquus, regrettably, the
in Vultur, the facies articularis acetabularis is poorly extended fossil species have no skeletal elements in common with Vultur
distally (not extended in Cathartes and Coragyps). The facies articu­ messii. This restricts direct comparisons mostly to differences in
laris antitrochanterica is narrow, bounding caudally a deep pit, size. From the Pisco Formation in Peru, two species have been
absent in other cathartics (in Gymnogyps there is a long furrow, recovered: Kuntur cardenasi from the late Miocene of Laguna
endowed with pneumatic foramina, laterally bounded by a marked, Seca locality, and Perugyps diazi from the late Miocene-early
caudally directed tubercle absent in Vultur). Medial to the stout Pliocene of Montemar and Sacaco Sur localities (Stucchi and
crista trochanteris (stouter in Gymnogyps), a large pneumatic fora­ Emslie 2005; Stucchi et al. 2015). Kuntur is ~ 12% larger than
men is placed (as is typical of Cathartidae). Having a unique fora­ Vultur gryphus and V. messii; while Perugyps is only ~ 7% larger,
men is also observed in Vultur gryphus, Breagyps clarki and but both species show several osteological differences with Vultur
4 F. J. DEGRANGE ET AL.

Figure 2. Holotype of Vultur messii sp. nov., MCH 363, right femur (A), compared with that of Vultur gryphus CIT-O 263 (B) in cranial, caudal, lateral, medial, proximal and
distal views. Abbreviations: cl, condylus lateralis; ct, condylus medialis; faa, facies articularis antitrochanterica; flc, fovea lig. capitis; fp, fossa poplitea; ftc, fovea
tendinis m. tibialis cranialis; gi, scar of m. gastrocnemius pars intermedia; ial, impressio ansae m. iliofibularis; icl, impressio lig. collateralis lateralis; ifi, scar
of m. iliofemoralis internus; ilcc, impressio lig. cruciatum cranialis; io, impressio obturatoriae; itc, scar of m. iliotrochanteris caudalis; p, caudal pit; pf, pneumatic foramen;
sf, sulcus fibularis; si, sulcus intercondylaris; sp, sulcus patellaris; tf, trochanter femoris; tgl, tuberculum m. gastrocnemius lateralis. The black arrow points at the
characteristic medial protuberance of Vultur. Scale = 1 cm.`.

gryphus (Stucchi and Emslie 2005; Stucchi et al. 2015): Kuntur cathartids (Emslie 1998). Plesiovultur nevesi and Wingegyps cartellei
regarding its tarsometatarsus, which shows a mosaic of features are condors which stem from Minais Gerais in Brazil (Alvarenaga
from Vultur, Geronogyps and Breagyps; and Perugyps by being and Olson 2004; Alvarenga et al. 2008). Wingegyps coming from the
particularly different regarding its mandible, coracoid and hind Pleistocene is much smaller than Vultur, and is more closely related
limbs. Aizenogyps toomeyae is condor found in the late Pliocene to Gymnogyps; meanwhile Plesiovultur coming from the late
of Florida, U.S.A.. Although this species has a similar size to the Pleistocene–early Holocene is ~11% smaller than Vultur, and is
Andean condor (and, hence V. messii), it has been noted that it is more similar to Sarcoramphus (which is considered to be a taxon
much more sturdier showing osteological differences to all the intermediate between condors and vultures, and not a true condor;
 HISTORICAL BIOLOGY 5

Stucchi et al. 2015). Pampagyps imperator from the late Pleistocene addition to the existence of incipient mountain chains, may also
of Buenos Aires province (Agnolín et al. 2017) is ~14% smaller than indicate that the centre of origin of this genus is linked to the Andes
Vultur. Hadrogyps coming from the middle Miocene of California mountain.
(Emslie 1988a), and Pliogyps from the middle Miocene-middle It was suggested that the condor-type cathartids have a North
Pliocene of Kansas and Florida (Tordoff 1959; Becker 1986) have American origin and that they may have become established in South
been considered to be related to condors, but not true condors America as early as the Late Miocene-Early Pliocene (Emslie 1988b;
(Stucchi and Emslie 2005). Dryornis pampeanus coming from the Stucchi and Emslie 2005; Stucchi et al. 2015). However, Agnolín
Pliocene of Chapadmalal and Monte Hermoso localities (Buenos (2016), based on the presence of fossil cathartids in the Oligocene
Aires province), and D. hatcheri coming from the early Miocene of of Brazil (Alvarenga 1985), and various condor records in the late
Lago Pueyrredón (Santa Cruz province) have been shown to be Miocene of Peru and the Pliocene of Argentina, stated that a South
more closely related to vultures than to condors (Degrange et al. American origin for the group should not be discarded. Moreover,
2021; Degrange 2022). Emslie (1988a,b) indicated that Gymnogyps radiated in North
Geronogyps reliquus has been recorded in the Pleistocene of America, while Vultur did so in South America. The presence of
Buenos Aires and Entre Ríos provinces in Argentina, and in the a fossil Vultur species, found in deposits older than 4.8 Ma, indicates
Talara Tar Seeps in Peru (Campbell 1979; Tonni and Noriega 1998; an earlier arrival and diversification of the condors in South America.
Noriega and Tonni 2007). Geronogyps is about the size of
Gymnogyps californianus, and although several differences in the
humerus discriminates this species from Vultur, the femur shown Acknowledgments
also several differences: the medial protuberance and the caudal pit We thank M. Armella, C. Madozzo, I. Tapia, J. Baez, A. Nuñez Blasco,
are absent, the trochanter femoris is more projected proximally J. M. Robledo, Á. Miño-Boilini, and G. Schmidt for their assistance in the
than in Vultur, the sulcus patellaris is narrower, the impressio field. AMNH ornithological collection staff, and L. Witmer (Ohio University
ansae m. iliofibularis faces laterally, and the sulcus fibularis is wider. Heritage College of Osteopathic Medicine, Athens, USA) are thanked for allow­
Breagyps clarki coming from the late Pleistocene of California ing access to Accipitriformes specimens. I. Tapia (CICTERRA) repaired and
photographed the fossil with her customary skill. D. Ksepka generously provided
has been phylogenetically related to Vultur gryphus (Emslie 1988b). photos of Gymnogyps californianus. We are grateful to the Editor G. Dyke, two
Nevertheless, although size difference between Breagyps and Vultur anonymous reviewers and D. Brandoni who provided helpful feedback on the
seems to be insignificant (Breagyps is only ~ 9% smaller) (Howard manuscript. Finally, our warmest acknowledgement goes to Lionel Andrés Messi
1974; Hertel 1992), Breagyps lacks the characteristic protuberance and the whole Argentinean football team for bringing joy to our whole country.
of the femur, and the medial proximal foramen is more proximally
located than in Vultur. Moreover, the crista trochanteris is more Disclosure statement
bent proximo-medially in Breagyps the impressio
ansae m. iliofibularis faces laterally, the fossa patellaris is wider, No potential conflict of interest was reported by the author(s).
and the trochlea fibularis is shallower.
Lastly, Gymnogyps differs from V. messii (and Vultur gryphus) by Funding
having a furrow proximally located to the unique pnematic foramen,
the protuberance so marked in Vultur is barely appreciable, a furrow This a contribution of PICT 2019-771, 2019-03412, and PUE 2016 - CONICET -
CICTERRA.
caudodistally located to the facies articularis antitrochanterica, the
impressio obturatoriae are located more proximally on the diaphysis,
a wider sulcus patellaris with a quadrangular contour when viewed ORCID
distally, a shallower fossa poplitea, a narrower trochlea fibularis, and
Federico J. Degrange http://orcid.org/0000-0002-9463-4893
a deeper sulcus intercondylaris. Ricardo A. Bonini http://orcid.org/0000-0002-1402-6530
The Andean condor nowadays inhabits an area from northern Sergio M. Georgieff http://orcid.org/0000-0002-5608-4819
Colombia and northwestern Venezuela to southernmost Argentina Lucía M. Ibañez http://orcid.org/0000-0003-1313-1379
Andean region where carrion is available, descending to the sea-level
in Peru, Chile and southermost Argentina, where it feeds of carrion
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