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A new late Miocene condor (Aves, Cathartidae) from

Peru and the origin of South American condors
a b c c
Marcelo Stucchi , Steven D. Emslie , Rafael M. Varas-Malca & Mario Urbina-Schmitt
a
Asociación para la Investigación y Conservación de la Biodiversidad, Av. Vicús 538, Lima,
33, Perú,
b
Department of Biology and Marine Biology, University of North Carolina Wilmington, 601 S.
College Road, Wilmington, North Carolina, 28403, U.S.A.
c
Departamento de Paleontología de Vertebrados, Museo de Historia Natural Javier Prado,
Universidad Nacional Mayor de San Marcos, Av. Arenales 1256, Lima, 14, Perú
Published online: 24 Jul 2015.
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To cite this article: Marcelo Stucchi, Steven D. Emslie, Rafael M. Varas-Malca & Mario Urbina-Schmitt (2015): A new late
Miocene condor (Aves, Cathartidae) from Peru and the origin of South American condors, Journal of Vertebrate Paleontology

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 Journal of Vertebrate Paleontology e972507 (5 pages)
Ó by the Society of Vertebrate Paleontology
DOI: 10.1080/02724634.2015.972507
A NEW LATE MIOCENE CONDOR (AVES, CATHARTIDAE) FROM PERU AND THE ORIGIN
OF SOUTH AMERICAN CONDORS
MARCELO STUCCHI,*,1 STEVEN D. EMSLIE,2 RAFAEL M. VARAS-MALCA,3 and MARIO URBINA-SCHMITT3;
1
Asociaci

on para la Investigaci

on y Conservaci

on de la Biodiversidad, Av. Vic

us 538, Lima 33, Per

2
Department of Biology and Marine Biology, University of North Carolina Wilmington, 601 S. College Road, Wilmington, North
Carolina 28403, U.S.A.;3Departamento de Paleontolog
ıa de Vertebrados, Museo de Historia Natural Javier Prado, Universidad
Nacional Mayor de San Marcos, Av. Arenales 1256, Lima 14, Per
u
http://zoobank.org/urn:lsid:zoobank.org:pub:BC9A886A-7271-44D2-BA93-4492FF406A4A
Cathartids or New World vultures are characterized by their
great ability at soaring flight, scavenging habits, and, for condors,
their large body size, including one of the greatest wingspans
Downloaded by [Rafael M. Varas-Malca] at 23:21 24 July 2015
among living birds. This group currently includes seven species
in five genera, divided into two groups: condors (Gymnogyps,
Vultur) and the smaller vultures (Coragyps, Cathartes). The fifth
genus, represented by the King Vulture (Sarcoramphus papa), is
considered to be a morphological intermediate between vultures
and condors (Fisher, 1944); however, some authors include it
among condors (Emslie, 1988a). Hertel (1992) also divided the
species of this family in two groups, but according to their body
size and the shape of their body and wings. The California Con-
dor (Gymnogyps californianus), the Andean Condor (Vultur gry-
phus), and the King Vulture are recognized by their large body
size, relatively long broad wings, and short tails; these Recent
species are well adapted to soaring and have been observed soar-
ing at altitudes higher than those of the small vultures in
Cathartes and Coragyps. Meanwhile, vultures have relatively
short, rounded wings and long tails, making them more maneu-
verable and better adapted for a more forested habitat.
The oldest fossil record of this family is a vulture from late Oli-
gocene deposits of S~ao Paulo, Brazil (Alvarenga, 1985). To date,
seven extant and 14 extinct species and genera have been recog-
nized in both North and South America (Emslie, 1988a; Stucchi,
2008). Here, we describe a new genus and species of a condor
coming from the late Miocene of Laguna Seca, a new locality in
the Pisco Formation (Ica region, Peru), based on a partial right
tarsometatarsus. This new taxon provides additional information
on the evolution and origin of South American condors.
Institutional Abbreviations—FMNH, Field Museum of Natu-
ral History (Chicago, U.S.A.); MUSM, Museo de Historia Natu-
ral Javier Prado, Universidad Nacional Mayor de San Marcos
(Lima, Peru); USNM, National Museum of Natural History,
Smithsonian Institution (Washington, DC, U.S.A.).
Osteological terminology follows Baumel and Witmer (1993).
The description of this new fossil condor is based on comparisons
with published measurements and figures of previously described
species (see Table 1) as well as museum skeletal material of the
extant species Gymnogyps californianus (USNM 492447), Vultur
gryphus (FMNH 104700, 104696, MUSM 14479), and Sarcoram-
phus papa (FMNH 338795). We also use information from Miller
(1910), Fisher (1944), Howard (1974), Campbell (1979), Becker
(1986), Emslie (1988a, 1988b, 1998), Su
arez and Emslie (2003), and
Stucchi and Emslie (2005). We recognize New World vultures and
*Corresponding author.
Color versions of one or more of the figures in this article can be found
online at www.tandfonline.com/ujvp.
SHORT COMMUNICATION
u, aicb.peru@gmail.com;
condors as within Order Cathartiformes, rather than as Cathartidae
within Accipitriformes, following Remsen et al. (2014).
SYSTEMATIC PALEONTOLOGY
AVES Linnaeus, 1758
CATHARTIFORMES Remsen, Cadena, Jaramillo, Nores,
Pacheco, P
erez-Em
an, Robbins, Stiles, Stotz, and Zimmer, 2014
CATHARTIDAE Lafresnaye, 1839 (sensu Remsen, Cadena,
Jaramillo, Nores, Pacheco, P

erez-Em
an, Robbins, Stiles, Stotz,
and Zimmer, 2014)
KUNTUR, gen. nov.
(Fig. 1)
Type and Only Known Species—Kuntur cardenasi, sp. nov.
Diagnosis—As for the species, by monotypy.
Etymology—‘Kuntur’ is the original term from which the word
‘condor’ is derived. It originates from quechua, the main native
language of Peru. Gender is masculine.
KUNTUR CARDENASI, gen. et sp. nov.
Holotype—MUSM 2423, a right tarsometatarsus that pre-
serves proximally both cotylae and the hypotarsus and distally
all three trochleae, but the distal shaft and fossa metatarsi I are
damaged through dorsoplantar crushing (Fig. 1). It was discov-
ered by Mario Urbina-Schmitt.
Measurements—Total length: 146 mm, proximal width:
32.8 mm, distal width: 34.6 mm, shaft width: ca. 15.8 mm. Addi-
tional measurements and ratios are available in Table 1.
Etymology—In honor of Santiago de C
ardenas, the first Peru-
vian naturalist to study condors in Peru in the 18th century
(C
ardenas, 1937 [1762]; Stucchi, 2012).
Locality and Age—MUSM 2423 was collected from a newly
identified vertebrate-bearing locality within the Ocucaje area
(Ica, Peru). Laguna Seca (LAG) is located about 30 km south-
east of the Ocucaje district, on the east margin of the Ica River
(Fig. 2). Two different depositional environments are exposed in
sequential layers. The upper layer contains sediments deposited
in ancient lagoons—from which the name of the locality is deri-
ved—of Recent-Pliocene age (DeVries, pers. comm., October,
2010). The underlying beds are marine sediments of the middle
Miocene–upper Pliocene Pisco Formation (see Muizon and
DeVries, 1985), from which MUSM 2423 was derived.
The lower marine sediments at LAG have yielded a diverse
marine faunal assemblage including sharks, manta rays (Mylio-
batis sp.) seabirds (penguins, Pelecaniformes indet.), cetaceans,
phocid seals, and mollusks. The lamniform sharks Isurus hastalis
and Carcharocles megalodon that are present occur elsewhere in
 Stucchi et al.—New late Miocene condor (e972507-2)

TABLE 1. Comparative measurements (in mm) of fossil and extant condor tarsometatarsi.

Kuntur Perugyps Vultur Gymnogyps Gymnogyps Gymnogyps

cardenasi diazi gryphus californianus(extant) californianus (fossil) howardae
Dimension MUSM 2423 MUSM 204 (n D 16) (n D 17) (n D 32)
Total length »146.0 »141.0 119.0–138.4 111.3–123.7 116.0–133.5 118.5–122.0
Proximal width 32.8 28.7 27.2–30.3 23.5–28.6 25.3–30.5 25.8–29.9
Distal width 34.6 »40.0 29.7–33.4 27.0–31.4 29.4–34.9 29.5–34.0
Trochlea tarsometatarsi II width 10.6 — 8.2–10.0 7.8–9.2 7.9–10.4 —
Trochlea tarsometatarsi II depth 14.5 — 13.6–16.0 12.0–13.4 12.8–15.8 —
Trochlea tarsometatarsi III width 13.5 »11.3 11.4–13.1 10.3–12.1 11.3–13.4 10.6–12.1
Trochlea tarsometatarsi III depth 19.2 »14.7 16.5–19.6 14.1–16.0 15.5–17.9 —
Trochlea tarsometatarsi IV width 9.5 — 6.4–8.7 6.7–7.5 6.7–8.3 —
Trochlea tarsometatarsi IV depth 15.1 — 14.0–16.4 13.0–16.0 14.0–17.2 —
TL/PW 3.82 4.91 4.44 4.35 4.34 4.46
TL/DW 4.22 3.52 4.06 3.99 3.85 3.84

Gymnogyps kofordi Gymnogyps Aizenogyps Breagyps clarki Geronogyps

Dimension (n D 4) varonai toomeyae (n D 12) reliquus
Total length 122.7 — 127.9 113.1–128.4 121.8–124.5
Proximal width 27.1–29.3 25.7 29.2 26.0–29.1 25.4–26.3
Distal width 29–31.6 33.0 32.8 28.2–31.0 29.0–30.0
Trochlea tarsometatarsi II width 8.5–9.8 9.2–10.0 9.3 7.0–8.8 —
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Trochlea tarsometatarsi II depth 12.3–13.8 13.1–13.4 14.8 12.5–14.5 —

Trochlea tarsometatarsi III width 10.5–11.0 11.4–11.7 12.4 11.0–12.5 10.6–11.6
Trochlea tarsometatarsi III depth 15.0–16.5 16.5–16.6 17.0 16.4–17.9 —
Trochlea tarsometatarsi IV width 7.0–8.0 7.6–8.3 7.8 6.3–7.3 —
Trochlea tarsometatarsi IV depth 14.0–15.5 14.9–16.3 15.2 12.6–15.3 —
TL/PW 4.19 — 4.38 4.37 4.77
TL/DW 3.88 — 3.90 4.01 4.17

Measurements of Perugyps diazi from Stucchi and Emslie (2005); Vultur gryphus, Gymnogyps californianus, Gymnogyps kofordi, and Breagyps clarki
from Emslie (1988a); Gymnogyps varonai from Su
arez and Emslie (2003); Aizenogyps toomeyae from Emslie (1998); and Gymnogyps howardae and
Geronogyps reliquus from Campbell (1979). For the ratios of V. gryphus, G. californianus, G. howardae, G. kofordi, B. clarki, and Ge. reliquus, the
means of measurements published by these authors were used.

upper Miocene beds (Ehret et al., 2012). Similarly, the pontopor-
iid dolphin Brachydelphis jahuayensis (MUSM 2425) is else-
where restricted to the El Jahuay level (9.95 Ma; Muizon and
Bellon, 1986; Lambert and Muizon, 2013). In addition, an inde-
terminate penguin species (MUSM 2424; Spheniscus sp. 3; see
Stucchi, 2007) also occurs in El Jahuay, Aguada de Lomas, and
Montemar localities (9.95 Ma–7.30 Ma; Ehret et al., 2012). Fur-
thermore, the phocid seal material (MUSM 2251) represents an
undescribed Acrophoca-like species. Moreover, the presence of
the mollusk Chione suggests that the deposit is at least 10 Ma,
because it is identified elsewhere in 14–10 Ma deposits (DeVries,
pers. comm., October 2010). Collectively, these fossil occur-
rences point to at least late Miocene age, but we could not
exclude an older age. According to this, a correlation with the El
Jahuay level (9.95 Ma, Tortonian) in the Sacaco area (Peru) is
possible. Further investigation regarding the geology and geo-
chronology of the locality is ongoing.
Diagnosis—The specimen is recognized as belonging to a
species in Cathartidae in having a deep fossa infracotylaris
dorsalis and sulcus extensorius, a large eminentia intercotyla-
ris, and a crista plantaris medialis that extends distally
approximately one-third the length of the shaft and by the
shape and relative position of the trochleae (sensu Emslie,
1998). The specimen MUSM 2423 is identified as represent-
ing a species of condor that is distinguished from smaller vul-
tures (Cathartes, Coragyps) and Sarcoramphus by its large
size and prominent eminentia intercotylaris. The fossil speci-
men differs from tarsometatarsi of other fossil and extant
condors (the species in Gymnogyps, Breagyps, Geronogyps,
Perugyps, Vultur, Aizenogyps, and Hadrogyps) by (1) shaft
long and relatively narrow, with total length and proximal
width exceeding all known species; (2) distinct fossa infraco-
tylaris dorsalis and sulcus extensorius on the proximal half of
shaft, with upper third relatively larger and deeper than in
all other species, and with minute foramina vascularia proxi-
malia; (3) hypotarsus in plantar view forms a plate that is
deeper than wide; and (4) trochlea metatarsi III with proxi-
modistally long anterior articular surface that tapers proxi-
mally, with the proximal edge nearly reaching the height of
the canalis interosseus distalis.
Description and Comparisons—Only one other condor is
known from the late Miocene of Peru, Perugyps diazi Stucchi
and Emslie, 2005, found also in outcrops of the Pisco Formation
(Montemar locality, 6.0–4.5 Ma; see Stucchi and Emslie, 2005).
The specimen here described differs from a right tarsometatarsus
(MUSM 204) referred to Perugyps by its greater length, greater
proximal width (Table 1), its shaft not as stout and flaring out-
ward proximally, and relatively larger anterior articular surface of
trochlea metatarsi III that narrows proximally (Fig. 1G). Also, in
Kuntur cardenasi, the canalis interosseus distalis is separated
from the foramen vasculare distale rather than having a proximal
origin within the foramen vasculare distale as is usual (Fig. 1A,
H). The following characters from Campbell (1979) were com-
pared between Kuntur and other fossil and living genera of con-
dors: (1) surface of trochlea metatarsi II small anteriorly in
Geronogyps (late Pleistocene, Peru), Breagyps (late Pleistocene,
California, U.S.A.; Miller, 1910), and Kuntur (Fig. 1A), moderate
to large in Aizenogyps (Plio-Pleistocene, Florida, U.S.A.; Emslie,
1998), Perugyps, and Gymnogyps, large in Vultur; (2) trochlea
metatarsi II extending distally to near the distal-most point of
trochlea metatarsi III in Geronogyps, does not project as far dis-
tally in Aizenogyps, Breagyps, Gymnogyps, Kuntur (Fig. 1A),
Perugyps, and Vultur; (3) trochlea metatarsi II with rounded pos-
terior edge in Geronogyps, tapers in width proximally in Kuntur
(Fig. 1C) and Vultur, similar shape but directed plantar-proximad
in Gymnogyps; (4) trochlea metatarsi IV of moderate size and
dorsal surface sharply delimited proximally in Vultur, moderate
Downloaded by [Rafael M. Varas-Malca] at 23:21 24 July 2015

Stucchi et al.—New late Miocene condor (e972507-3)

 Stucchi et al.—New late Miocene condor (e972507-4)

FIGURE 2. Location of Laguna Seca local-

ity, late Miocene, Pisco Formation (Ica, Peru).
Downloaded by [Rafael M. Varas-Malca] at 23:21 24 July 2015

to large but barely delimited proximally in Gymnogyps, large and
not well-defined proximally in Aizenogyps, Breagyps, Gerono-
gyps, and Kuntur (Fig. 1A); (5) trochlea metatarsi IV with a dis-
tinct plantar-proximal corner to plantar edge in Geronogyps,
rounded in Kuntur (Fig. 1D) and Vultur, rounded to prominent
proximal projection in Gymnogyps; and (6) groove on trochlea
metatarsi IV anteriorly, moderately wide in Geronogyps, Gymno-
gyps, and Kuntur (Fig. 1D), very wide in Vultur.
Additional relevant characters from Emslie (1988a) include
(1) posterior protrusion of cotyla lateralis small to moderate in
Breagyps, Cathartes, Coragyps, Kuntur, Sarcoramphus, and Vul-
tur, large in Gymnogyps; (2) internal rim of cotyla medialis low
in Breagyps, Kuntur, and Vultur, high and rounded in Cathartes,
Coragyps, and Sarcoramphus; Gymnogyps presents both charac-
ter states; (3) crista plantaris medialis short, extending distally
less than one-third the distance on the shaft in all genera, except
Sarcoramphus, in which it extends up to half the distance distally
on the shaft; (4) proximal end of the fossa infracotylaris dorsalis
deeply excavated in all genera except Gymnogyps, in which both
deep and shallow conditions are present; and (5) sulcus extensor-
ius extends to midlength of the shaft in Breagyps, Coragyps,
Kuntur, Perugyps, and Vultur, but is longer and extends farther
down the shaft in Cathartes and Sarcoramphus; Gymnogyps
presents both states.
DISCUSSION
Morphological comparisons of the tarsometatarsus of Kun-
tur with those of other genera of Cathartidae indicate that the
new genus shares three of the above characters noted by
Campbell (1979) with Vultur (2, 3, 5), three with Geronogyps
(1, 4, 6), and minimally three with Breagyps (1, 2, 4; other fea-
tures were not verified in this genus). Of the five additional rel-
evant characters taken from Emslie (1988a) and described
above, we found that Kuntur shares all with Vultur and Brea-
gyps; five other characters of the tarsometatarsus were listed
by Emslie (1988a), but two characters were not comparable
due to fossil deterioration, and three are shared among all
taxa, so they were not considered. These comparisons suggest
that Kuntur is affiliated more with fossil and extinct South
American condors (i.e., Perugyps, Geronogyps, Gymnogyps
howardae, and Vultur) than with North American species
except for Breagyps, which was considered of intermediate
morphology between Gymnogyps and Vultur by Fisher (1944).
Emslie (1988a) found Breagyps to be more closely related to
Vultur, an interpretation previously inferred by Miller (1910)
and Howard (1974). Furthermore, Hadrogyps aigialeus Emslie,
1988 (middle Miocene, 13–15 Ma, California, U.S.A.), and
Pliogyps charon Becker, 1986 (late Miocene, 9 Ma, Florida,
U.S.A.), are proportionally more robust than K. cardenasi,
with more widely separated foramina vascularia proximalia, a
wider fossa infracotylaris dorsalis, and a larger trochlea meta-
tarsi II (Becker, 1986; Emslie, 1988b). In addition, two other
fossils, MUSM 1260 (a fragmentary tibiotarsus) and MUSM
1455 (an ungual phalanx), attributed by Stucchi (2008) to an
indeterminate species of Cathartidae, are similar in age to
Kuntur. These specimens cannot be compared with Kuntur
because they do not include a tarsometatarsus.
Origin of South American Condors
The presence of condor-like vultures H. aigialeus and P.
charon from the middle and late Miocene, respectively, of North
America still supports a North American origin for condors.
Although the earliest record of Cathartidae is the late Oligocene
Brasilogyps faustoi Alvarenga, 1985, from S~ao Paulo (Brazil), it
was a vulture that shared similar morphology and body size with
Coragyps (Alvarenga, 1985; Emslie, 1988b), thus precluding it
from consideration as an ancestral condor. Conversely, H. aigia-
leus and P. charon are considered to be basal condors despite
their medium size, which is smaller than that of S. papa; see
Emslie (1988b).

FIGURE 1. Holotype of Kuntur cardenasi (MUSM 2423). A, dorsal view, and its interpretative line drawing; B, plantar view, and its interpretative
line drawing; C, internal view; D, external view; E, proximal view; F, distal view; G, detail of proximal half to show the fossa infracotylaris dorsalis and
the sulcus extensorius; H, detail of distal fragment to show separation of foramen vasculare distale from canalis interosseus distalis (white arrows).
Numbers indicate anatomical structures as follows: 1, cotyla lateralis; 2, eminentia intercotylaris; 3, cotyla medialis; 4, fossa infracotylaris dorsalis; 5,
sulcus extensorius; 6, foramen vasculare distale; 7, canalis interosseus distalis; 8, trochlea metatarsi IV; 9, trochlea metatarsi III; 10, trochlea metatarsi
II; 11, crista plantaris medialis; 12, fossa parahypotarsalis medialis. Diagonal lines indicate broken surfaces; dotted areas indicate those filled with sedi-
ment. Scale bars equal 20 mm. (Color figure available online at www.tandfonline.com/UJVP.)
 Stucchi et al.—New late Miocene condor (e972507-5)
All Peruvian fossil condors from the Miocene, the new taxon
Kuntur cardenasi, P. diazi, and an indeterminate cathartid
(MUSM 1260 and MUSM 1455), indicate an earlier arrival of
condors to South America than previously thought, perhaps by
following the coastal corridors on the western side of the conti-
nent (Stucchi and Emslie, 2005; Stucchi, 2008). However, differ-
ences in size between the larger species of Perugyps, Kuntur, and
the North American condor-like vultures (Hadrogyps, Pliogyps)
suggest that an increase in body size occurred rapidly once ances-
tral condors reached South America, whereas large dimensions
in North American condors did not appear until Aizenogyps
evolved in the late Pliocene. This evidence suggests that both
condor groups acquired their large body size independently
(Stucchi, 2008). This hypothesis still leaves a possible link with
Brasilogyps and ancestral condors, but Emslie (1988a) found
more affinities between North and South American condors and
Sarcoramphus rather than the smaller vultures Cathartes and
Coragyps, both considered more primitive. Age, body size, and
similarities between Kuntur, Breagyps, Geronogyps, and Vultur
do not challenge this view because these latter genera are youn-
ger, and a possible relationship of condors with Brasilogyps
requires discovery of additional fossil material.
Downloaded by [Rafael M. Varas-Malca] at 23:21 24 July 2015
Ancient occurrence of condors along the Peruvian coast in the
late Miocene is reflected in the habits of coeval local species
from that time to the present, as shown by late Pleistocene con-
dors from Talara Tar Seeps, Peru (Geronogyps and Gymnogyps;
Campbell, 1979), as well as the extant Andean Condor (Stucchi,
2013), the only condor currently living in this region.
ACKNOWLEDGMENTS
We thank T. J. DeVries and K. Nick for sharing information
on the locality age. R. Salas-Gismondi (MUSM), S. Olson and J.
Dean (USNM), and D. Willard and J. Bates (FMNH) allowed
access to collections under their care. We also thank editor T.
Worthy as well as two anonymous reviewers for their useful sug-
gestions to improve the manuscript.
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Submitted April 20, 2014; revisions received August 20, 2014; accepted
September 25, 2014.
Handling editor: Trevor Worthy.
Citation for this article: Stucchi, M., S. D. Emslie, R. M. Varas-Malca,
and M. Urbina-Schmitt. 2015. A new late Miocene condor (Aves, Cathar-
tidae) from Peru and the origin of South American condors. Journal of
Vertebrate Paleontology. DOI: 10.1080/02724634.2015.972507.