[Papers in Palaeontology, 2021, pp.
1–16]
PHYLOGENETIC AFFINITIES AND MORPHOLOGY OF
THE PLIOCENE CATHARTIFORM DRYORNIS
PAMPEANUS MORENO & MERCERAT
by FEDERICO J. DEGRANGE 1 , CLAUDIA P. TAMBUSSI 1 ,
MATÍAS L. TAGLIORETTI 2 , 3 and FERNANDO A. SCAGLIA 2
1
Centro de Investigaciones en Ciencias de la Tierra (CICTERRA), UNC, CONICET, Avenida V
elez S
arsfield 1611, X5016GCA, C
ordoba, Argentina;
fjdino@gmail.com, tambussi.claudia@conicet.gov.ar
2
Museo Municipal de Ciencias Naturales Lorenzo Scaglia, Av. Libertad 3099, B7600HJB, Mar del Plata, Argentina; paleomat@gmail.com, feroscaglia@gmail.com
3
Facultad de Ciencias Exactas y Naturales, Instituto de Geolog
ıa de Costas y del Cuaternario (CIC-UNMdP), Universidad Nacional de Mar del Plata, Dean Funes
3250, B7602AYJ, Mar del Plata, Argentina
Typescript received 12 October 2020; accepted in revised form 18 December 2020
Abstract: The fossil bird Dryornis pampeanus Moreno &
Mercerat, is reinterpreted after examination of new referred
material (humerus, coracoid, fragments of ulna, radius, sca-
pula, sternum and tibiotarsus) from the Pliocene Chapad-
malal Formation of Argentina. The current diagnosis is
emended in the light of important considerations that cast
doubt on the previous attribution of the taxon to condors.
The phylogenetic position of D. pampeanus was tested in a
series of maximum parsimony analyses that included all
seven living Cathartiformes and 207 osteological characters.
The phylogenetic analyses placed D. pampeanus as the sister
taxon of extant vultures. An estimation of 26 kg for the
mass, positions D. pampeanus as the largest cathartiform to
DRYORNIS pampeanus Moreno & Mercerat, 1891 was estab-
lished based on distal portions of the right humerus and
right femur that were supposedly from the same individ-
ual. This association was later amended, assigning the
femur to the Phorusrhacidae Mesembriornis milneedwardsi
(Patterson & Kraglievich 1960; Brodkorb 1967; Tambussi
& Noriega 1999). This material was collected from the
early Pliocene Monte Hermoso Formation, exposed on
the Buenos Aires Atlantic coast of Argentina. This is an
important unit because it is the type locality of the
Montehermosan South American Land Mammal Age
(SALMA), which includes diverse vertebrate remains
(Deschamps et al. 2012; Tomassini et al. 2013). This spe-
cies also possibly occurs in slightly more recent sediments
corresponding to the Toro Negro Formation exposed in
La Rioja province (Rodr
ıguez Brizuela 2004). Its assign-
ment, however, could not be corroborated because the
material could not be located.
According to the general features of the humerus, Dry-
ornis was considered a cathartid condor related to Vultur
© 2021 The Palaeontological Association
have ever lived. The presence of this taxon in both the
Monte Hermoso and Chapadmalal Formations not only
extends the stratigraphic range of the species, but also sup-
ports the idea that they were partially contemporaneous dur-
ing the early Pliocene. The dependence of the vultures on
ephemeral carrion suggests that they have especially large
ranges. The sites from which the lectotype and new material
were recovered (Monte Hermoso and Chapadmalal, respec-
tively) are only 400 km apart, suggesting that the two sites
were at least partly contemporaneous.
Key words: Cathartidae, Chapadmalal, Pliocene, Dryornis,
scavenger, vulture.
by Tonni (1980), although the diagnosis of the species
needs to be verified, largely because of the fragmentary
nature of the lectotype. In the Cenozoic, condors have
lived in both Americas since the Oligocene (Wetmore
1927; Alvarenga et al. 2008). Now, they do not live in the
Pampean region, although during the Pleistocene they
were more diverse in that area, and consisted of at least
six different species (Tambussi & Noriega 1996, 1999;
Tonni & Noriega 1998; Alvarenga et al. 2008, Agnol
ın
et al. 2017). There is also now only one extant species in
South America, the Andean condor Vultur gryphus, which
is limited to mountain areas or semiarid environments,
suggesting a great reduction in population and diversity,
and a westward shift in the distribution of this lineage
of birds today. The Andean condor is a large bird
(~12 500 g, Dunning 2008) that is found in high and
semi-arid areas.
Cathartids or New World vultures also include another
morphotype, known as vultures, which includes medium-
sized birds (between 935 and 2100 g) with short wings
doi: 10.1002/spp2.1361 1
2 PAPERS IN PALAEONTOLOGY
and rounded long tails. Condors and vultures are obligate
scavengers, known for their efficient soaring flight and
acute eyesight and smell (Holland et al. 2017).
New material that has been positively assigned to
D. pampeanus (MMP 5731) allows us to significantly
expand the information on this species and to explore,
for the first time, its phylogenetic position within
cathartiforms. The material comes from Palaeosol 6 of
the Chapadmalal Formation at La Estafeta Beach, close
to Mar del Plata City on the Atlantic coast of Argen-
tina (Fig. 1). A late Pliocene age (Piacenzian) is
assigned to these sediments (3.3 Ma according to the
available dating, Schultz et al. 1998). This formation
contains a diverse terrestrial bird fauna assemblage
(Tambussi 2011; Degrange et al. 2015) that includes
tinamids, phorusrhacids, eagles, charadriiforms, and pas-
seriforms. Dryornis is the only registered scavenger bird
so far.
The aim of the present paper is to review the morphol-
ogy and phylogenetic position of D. pampeanus based on
the newly collected specimen, and to discuss the relation-
ships between the avian assemblages of the Monte Her-
moso and Chapadmalal Formations.
GEOLOGICAL SETTING
Z
arate (2005) proposed an age range of 5 or 4.5 Ma to
3.2 Ma for the Chapadmalal Formation, and a range of
5.28 Ma to 5 or 4.5 Ma for the Monte Hermoso Forma-
tion (Deschamps et al. 2012; Tomassini & Montalvo
2013; Tomassini et al. 2013). It is noteworthy that there
is no absolute dating for the Monte Hermoso Formation
and only one date for the Chapadmalal Formation in the
upper levels (3.27
Ma, Schultz et al. 1998). Pascual
et al. (1965) and Pascual & Odreman Rivas (1973)
claimed that faunal differences between Chapadmalalan
and Montehermosan age are minimal, and therefore that
they should both be considered in the same SALMA,
retaining Montehermosan for the time span encompassed
by Monte Hermoso and Chapadmalal Formations. Tonni
et al. (1992) and Cione & Tonni (1995a, b, 2005) dis-
cussed the validity of the SALMA and established two
biozones for the Monte Hermoso Formation (the Try-
dodon gaudryi Biozone and the Neocavia depressidens Bio-
zone) and one biozone for the Chapadmalal Formation
(the Paraglyptodon chapalmalensis Biozone). Later, after
further fieldwork, Tomassini et al. (2013) proposed a sin-
gle biozone for the Monte Hermoso Formation (the
Eumysops laeviplicatus Biozone), which became the basis
of the Montehermosan Stage (Cione et al. 2015). New
discoveries continue to add more entries to the list of
fauna for the Chapadmalalan age in its type section
(Taglioretti et al. 2014; Zamorano et al. 2014; Degrange
et al. 2015; Zurita et al. 2016a; present work), and a revi-
sion of some taxa is improving the knowledge for the
Chapadmalalan fauna, and introducing critical modifica-
tions (Cruz et al. 2016; Zurita et al. 2016b; Beck &
Taglioretti 2019). Moreover, recent studies indicate that
most taxa are present in both the Chapadmalal and
Monte Hermoso Formations (Tomassini et al. 2013;
Zurita et al. 2014), indicating that faunal similarities are
closer than previously assumed. Further studies will be
required to assess this faunal similarity and explain
whether the differences are due to palaeobiogeographical
or to temporal factors, or both.
MATERIAL AND METHOD
Unless otherwise indicated, osteological terminology fol-
lows Baumel & Witmer (1993).
Body size estimation
The estimation of body mass (BM, in g) was carried out
following Field et al. (2013), using the maximum diameter
of the coracoid humeral articulation facet (HAF, in mm):
Ln BM ¼ 2:44 ðln HAFÞ þ 2:00
Phylogenetic analysis
The phylogenetic relationships of D. pampeanus within
Cathartiformes were evaluated using the character–taxon
matrix of Brito (2008), which includes 207 morphological
characters, and 70 living taxa (see Degrange et al. 2021).
The position of Dryornis was explored in two ways, by
including and excluding the extinct Pampagys imperator
Agnol
ın et al. (2017). Anseranas semipalmata was selected
as the outgroup. All missing data were scored as ‘?’. A
parsimony analysis was performed using TNT (Tree anal-
ysis using New Technology) version 1.1 (Goloboff &
Catalano 2016). For the analysis, no implied weighting
was used because its use is not exempt from criticism
(Farris 1969, 1989; Goloboff 1993; Kluge 1997). Only 4
characters of the 39 of the matrix built by Emslie (1988)
could be encoded, which is why it was discarded for this
work. Falconidae, Ardeidae and Pelecaniformes were not
included in the analysis. One hundred random Wagner
trees were generated with 100 additional replicates, using
the TBR (tree bisection and reconnection) algorithm and
saving 100 trees per replicate, collapsing the trees after
the search. The strict consensus tree was generated from
the trees obtained.
 DEGRANGE ET AL.: MORPHOLOGY AND PHYLOGENY OF DRYORNIS 3

FIG. 1. A, location map; and B,

A
stratigraphic log for Dryornis pam-
peanus (MMP 5731). Abbreviations:
AAL, Arroyo Lober
ıa Alloformation;
APLL, Playa Los Lobos Alloforma-
tion; APSC, Playa San Carlos
Alloformation; P5–8, paleosol 5–8.

Institutional abbreviations. MLP, Museo de La Plata, Buenos SYSTEMATIC PALAEONTOLOGY

Aires, Argentina; MMP, Museo Municipal de Ciencias
Naturales Lorenzo Scaglia, Mar del Plata, Buenos Aires, The most recent molecular analysis showed that Cathartidae is
Argentina. related to accipitrids (Hackett et al. 2008; Jarvis et al. 2014;
4 PAPERS IN PALAEONTOLOGY
Prum et al. 2015), while another suggested that they may belong
to their own order Cathartiformes (Johnson et al. 2016). This
later proposal is in agreement with that of the South American
Classification Committee, following a suggestion by Van Remsen
(Proposal 361, 29 September 2008).
AVES Linnaeus, 1758
NEOGNATHAE Pycraft, 1900
CATHARTIFORMES Coues, 1894
CATHARTIDAE Lafresnaye, 1839
Genus DRYORNIS Moreno & Mercerat, 1891
Type species. Dryornis pampeanus Moreno & Mercerat, 1891
Dryornis pampeanus Moreno & Mercerat, 1891
Figures 2–6
Lectotype. MLP 20-169, distal portion of right humerus (Fig. 2).
Referred new specimen. MMP 5731, right pectoral girdle and par-
tial arm found articulated: complete right humerus, proximal ends
of the right ulna and radius, right coracoid, right incomplete sca-
pula, fragment of sternum, fragment of tibiotarsus.
Additional assigned material. MMP 4679, distal extremity of left
ulna; MLP 90-X-1-1, incomplete right ulna lacking the distal
extremity, and proximal portion of right radius (see Tambussi &
Noriega 1999).
Emended diagnosis. The largest Cathartidae so far registered (ex-
clusive features denoted by *). Sternum highly pneumatized*.
Absence of a marked fossa, ventral to the cotyla scapularis and
medial to the facies articularis humeralis. The crista articularis of
the facies externa presents a sharp edge, smoother than extant
Cathartids*. Presence of a distinctive oval foramen n.
FIG. 2. Dryornis pampeanus MLP 20-169, lectotype. Distal portion of a right humerus in: A, cranial; B, distal; C, lateral; D, medial;
and E, caudal view. Scale bar represents 1 cm.
supracoracoidei medial in position*. Presence of foramina in the
impressio m. sternocoracoidei. The facies articularis humeralis of
the scapula is slightly concave. Small foramina in the base of the
caput humeri. Elongated and poorly marked scar of the m.
humerotriceps. The stout epicondylus ventralis has a flat edge*.
Condylus ventralis circular. The facies articularis radialis of the
ulna is curved and deep. Incisura radialis is triangular in shape,
deep and pneumatic. Presence of a blunt tubercle laterodorsally
to the incisura tendinosa*. The cotyla humeralis of the radius is
quadrangular in shape. Tuberculum bicipitale radii very well-
developed*.
Occurrence. La Estafeta Beach, Mar del Plata, Buenos Aires Pro-
vince, Argentina (Fig. 1). Chapadmalal Formation, lowest part
of Playa Los Lobos Alloformation (Palaeosol 6 of Z
arate & Fas-
ano 1989; levels 9–10 of Kraglievich 1952), 3.3 Ma (Schultz et al.
1998), Piacenzian, late Pliocene. S38°90 48.38″, W57°370 50.84″.
Description
Comparisons with Pleistovultur nevesi Alvarenga et al., 2008,
Kuntur cardenasi Stucchi et al., 2015, Perugyps diazi Stucchi &
Emslie, 2005 and Wingegyps cartellei Alvarenga & Olson, 2004
are not possible due to the lack of corresponding elements for
each specimen.
Measurements of the new Dryornis pampeanus material are
provided in Table 1. The Dryornis body mass estimation is
26 161 g (HAF, 28.48 mm), which makes this the largest of the
cathartiforms previously registered; and 57% greater than the
condor Vultur grypus and 94% greater than the turkey vulture
Cathartes aura according to the measurements in Dunning
(2008).
Scapula. The facies articularis humeralis is oval and slightly con-
cave (concave in Cathartes, flat in Coragyps and Vultur). Medial
to it, there is a small foramen pneumaticum. The labrum glenoi-
dale is poorly marked, as in Vultur. The facies articularis cora-
coidea is elongated and flat. The tuberculum coracoideum is
 DEGRANGE ET AL.: MORPHOLOGY AND PHYLOGENY OF DRYORNIS
poorly developed. The facies articularis clavicularis is oval in
shape, and wider than in Vultur. The acromion is poorly devel-
oped. The collum scapulae is wide. The corpus scapulae is
curved. The caudal extremity is missing, so the length of the
corpus cannot be measured. The facies lateralis and costalis are
convex (Fig. 3A–D).
Sternum. Only a small fragment of the sternum has been pre-
served (Fig. 3E), which presents only the rectangular shape of
the processus costalis. The processi are flat with rounded edges,
showing a large pneumatization cranially, similar to the condi-
tion observed in smaller cathartids, not in Vultur.
Coracoideum. The os coracoideum is markedly larger than that
of Vultur. The cotyla scapularis is subcircular and shallow, with
thick margins that forms a lip distally. The facies articularis
humeralis is flat, oval, and broad (Fig. 4). Ventral to the cotyla
and medial to the facies, in Vultur there is a marked fossa con-
taining pneumatic foramina. This feature is absent in Dryornis
(as in Coragyps and Cathartes). The labrum glenoidale is
conspicuous, rounded (sharp in Cathartes and Coragyps), and lat-
erally projected. The smooth convex facies articularis clavicularis
does not project beyond the very well-developed processus
acrocoracoideus. The impressio lig. acrocoracohumeralis is
medial, conspicuous, more elongated than in Vultur, deep and
situated proximad to the facies articularis humeralis. Proximo-
lateral to the impression, an elongated impressio coraco-
brachialis can be observed. The processus procoracoideus is
very broad, stout but poorly developed medially (similar to
Vultur and Geronogyps). The subcapital crest (following Elza-
nowski et al. 2012) is prominent, acute and borders laterally a
deep pneumatic fossa, which is also bounded dorsally by the
facies articularis clavicularis. This fossa is larger and deeper
than that of Vultur. The distinctive oval foramen n. supracora-
coidei (absent in Geronogyps reliquus Campbell, 1979) is medial
in position. The sulcus m. supracoracoidei is wide dorsoven-
trally. Although there is a small portion missing, it is possible
to establish that the corpus coracoidei is stout, although pro-
portionally elongated compared with Vultur and Gymnogyps.
The linea intermuscularis ventralis forms a sharp crest poorly
developed and laterally placed, although more medially than in
Geronogyps. The impressio m. sternocoracoidei is wide and
shallow, highly pneumatic, ventrally delimited by a stout border
(crista articularis sternalis of the facies interna), absent in Vul-
tur, present in Cathartes, and poorly developed in Coragyps.
The processus lateralis is short, although stout. The facies artic-
ularis sternalis is widened medially. The angulus medialis is
very marked. The crista articularis of the facies externa presents
a sharp edge, smoother than in extant cathartids. The margo
supra-angularis is narrow and sharp.
Humerus. It is markedly larger than that of Vultur. The caput
humeri is robust, wide, oval in shape, very projected proxi-
mally (Fig. 5), and aligned with the planum intertuberculare.
The sulcus lig. transversus is restricted to the medial margin,
forming a deep scar. The tuberculum dorsale is stout, conical
and proximolaterally directed (proximally directed in Gerono-
gyps, see Campbell 1979; Noriega & Tonni 2007). The crista m.
5
supracoracoidei is an elongated scar, very well-marked. The
crista deltopectoralis is robust, laterocranially directed, although
not that much as in Coragyps. The impressio coracobrachialis
is not marked. Distally and bordered by the crista deltopec-
toralis, the elongated m pectoralis scar can be noticed. Medially
to it, the narrower m. coracobrachialis cranialis scar is located.
Distally to the m. pectoralis scar, a stout and prominent tuber-
cle is placed. The intumescentia humeri is flat. The crista
bicipitalis is stout. The fossa pneumotricipitalis is wide and
deep, dorsally delimited by a stout caudally projected tubercu-
lum ventrale, typical of Cathartidae. The crus dorsale et ven-
trale fossae are robust and very marked. An elongated and
poorly marked scar corresponding to that of the m. humerotri-
ceps is seen ventral to the crus ventrale fossae (this condition
is observed in all the cathartids examined). In caudal view, dis-
tal to the caput humeri, pneumatic foramina can be observed
in two groups: one distal to the caput humeri and the tubercu-
lum dorsale and the other between the caput and the tubercu-
lum ventrale. Although the degree of pneumatization in this
area is higher than that observed in Vultur and Gymnogyps,
there are fewer foramina in the second group than are seen in
Coragyps and Cathartes. In these latter two taxa, a large fossa
contains those foramina, a feature absent in Vultur, Gymnogyps,
Sarcoramphus and Dryornis. The incisura capitis humeri is oval
and deep. The corpus humeri is robust, long and curved, convex
laterally when viewed cranially and sigmoid when viewed later-
ally, describing an ‘S’. The fossa m. brachialis is elliptical, wide,
deep and pneumatic, as in Vultur (greatly pneumatic in Cath-
artes and Coragyps), less extended laterally as in the lectotype
(Tambussi & Noriega 1999). Identically to the lectotype, the
condylus dorsalis is stout, elongated, medioproximally directed
(Figs 2, 5), and more developed cranially than Vultur. The
condylus ventralis is circular (slightly elliptical in Vultur and
Geronogyps), as in the lectotype. The incisura intercondylaris is
shallow. The epicondylus dorsalis is subquadrangular. The epi-
condylus ventralis is stout, subconical, and with a flat edge
(rounded in Cathartes, Coragyps and Vultur). The tuberculum
supracondylare ventrale is a very well-marked flat scar. The proces-
sus flexorius is poorly developed. The fossa olecrani is oval in shape
and deep, as in the lectotype. The sulcus humerotricipitalis is shal-
low, but very well-marked, slightly shallower than the sulcus scapu-
lotricipitalis. This sulcus is clearly delimited by stout ridges.
Ulna. The proximal extremity is slightly larger than that of
Vultur (Fig. 6). The olecranon is stout, rounded (more pointed
in Coragyps and Cathartes), narrower than in Pampagyps
(Agnol
ın et al. 2017), and less projected proximally than in
Geronogyps. The scar of the m. humerotriceps is poorly marked.
The cotyla dorsalis is elongated and slightly convex, meanwhile
the cotyla ventralis is circular and concave. The crista intercoty-
laris is curved, low and poorly marked. The processus cotylaris
dorsalis is flat, medially projected (more projected than in
Pampagyps, but less than in Geronogyps), with a strong oval
depression at its base (Tambussi & Noriega 1999). Laterally, the
conspicuous and elliptical impressio m. scapulotricipitis is
noted. The facies articularis radialis is curved, with rounded
smooth edges, bounded dorsally by a strong edge (absent in
Vultur), and is deeper than that of Vultur (contra Tambussi &
6 PAPERS IN PALAEONTOLOGY

FIG. 3. A–E, scapula; and F–G, fragmentary sternum of Dryornis pampeanus MMP 5731. A–D, scapula in: A, dorsal; B, ventral;
C, medial; and D, lateral view. E, extremitas cranialis scapulae in proximal view. F–G, sternum in: F, lateral; and G, dorsal view.
Abbreviations: acr, acromion; fac, facies articularis acrocoracoidea; fah, facies articularis humeralis; pc, processus costalis. Scale bars
represent 1 cm.

Noriega 1999). The incisura radialis is triangular in shape, very
well-marked, deep and pneumatic as in Vultur and Geronogyps
(poorly pneumatic in Pampagyps and Cathartes, non-pneumatic
in Coragyps). The trochlea humeroulnaris is elliptical and poorly
marked. The stout papillae remigales caudales are elongated
proximodistally, more than in Vultur, and similar to Coragyps;
situated more medial to the midline than in Vultur (Tambussi
& Noriega 1999). Regarding the extremitas distalis ulnae, the
depressio radialis is smooth, oval and shallow (flat in Vultur
and Coragyps). The incisura tuberculi carpalis is very well-
 DEGRANGE ET AL.: MORPHOLOGY AND PHYLOGENY OF DRYORNIS 7

FIG. 4. Coracoid of Dryornis pampeanus MMP 5731. A, ventral; B, medial; C, lateral; and D, dorsal view. E, distal view. F, proximal
view. Abbreviations: am, angulus medialis; c, corpus coracoidei; ce, crista epimarginalis; ci, impressio coracobrachialis; csb, crista sub-
capitalis; csc, cotyla scapularis; fac, facies articularis clavicularis; fah, facies articularis humeralis; fas, facies articularis sternalis; fns, fora-
men n. supracoracoidei; ila, impressio lig. acrocoracohumeralis; ims, impressio m. sternocoracoidei; le, labrum externum; li, labrum
internum; pa, processus acrocoracoideus; ppr, processus procoracoideus; sms, sulcus m. supracoracoidei. Scale bars represent 1 cm.

marked. The tuberculum carpale is prominent, bearing a unique
pneumatic foramen on its dorsal aspect (double in Geronogyps).
The sulcus intercondylaris is poorly preserved. The incisura
tendinosa is wide and very well-marked. Laterodorsally, there is
a blunt tubercle present (more prominent than that of extant
cathartids). The condylus ventralis ulnaris is wide, as in Vultur.
The condylus dorsalis ulnaris has a labrum condyli dorsalis
well-projected ventrally, as in Vultur.
Radius. Only the proximal region of MMP 5731 has been
preserved (Fig. 6K–O), being not quite different from that
of MLP 90-X-1-1. The cotyla humeralis is quadrangular in
8 PAPERS IN PALAEONTOLOGY
 DEGRANGE ET AL.: MORPHOLOGY AND PHYLOGENY OF DRYORNIS 9

FIG. 5. Humerus of Dryornis pampeanus MMP 5731. A, cranial; B, medial; C, caudal; and D, lateral view. E, distal view. F, proximal
view. Abbreviations: cbp, crista bicipitalis; cd, condylus dorsalis; cdp, crista deltopectoralis; ch, caput humeri; cv, condylus ventralis;
ed, epicondylus dorsalis; ev, epicondylus ventralis; fmb, fossa m. brachialis; fo, fossa olecrani; fp, fossa pneumotricipitalis; ic, incisura
capitis; ih, intumescentia humeri; ii, incisura intercondylaris; mcc, m. coracobrachialis cranialis scar; mp, m. pectoralis scar; pf,
processus flexorius; sht, sulcus humerotricipitalis; slt, sulcus lig. transversus; sst, sulcus scapulotricipitalis; t, distal tubercle; td,
tuberculum dorsale; tsv, tuberculum supracondylare ventral; tv, tuberculum ventrale. Scale bars represent 1 cm.

shape (sub-oval in Vultur). Distal to the cotyla, on the dor-
sal surface there is a large pneumatic foramen in Geronogyps
(Campbell 1979). This feature is absent in Dryornis. The
facies articularis ulnaris has a smooth lateral edge that is
straight ventrally. The tuberculum bicipitale radii is placed
immediately next to the cotyla, and it is very well-developed
(Tambussi & Noriega 1999). Ventrally, a strong oval scar
(typical of Cathartidae) can be observed. The cross-section
of the corpus radii is oval.
Tibiotarsus. Only a small fragment assigned tentatively to the left
tibiotarsus has been preserved. The crista fibularis is flat, slanted
caudally, short and sturdier than that of Vultur.
PHYLOGENETIC ANALYSIS
Phylogenetic analysis yielded four most parsimonious
trees (MPTs) of 628 steps, with a consensus of 639 steps
(consistency index (CI) = 0.36; retention index (RI) =
0.85) (Fig. 7A). Dryornis is placed as sister group of
Coragyps + Cathartes. This relationship is based on one
synapomorphy: the presence of an oval-shaped foramen
n. supracoracoidei. Condors have a circular foramen,
located more medially on the corpus coracoidei. Dryornis
pampeanus is characterized by having two autapomor-
phies: the presence of foramina in the impressio m. ster-
nocoracoidei, and the presence of small foramina in the
base of the caput humeri.
We also performed a second analysis with the addition
of the fragmentary specimen of Pampagyps imperator,
which resulted in 28 MPTs of 634 steps. The consensus
tree has 654 steps (CI = 0.35; RI = 0.85) (Fig. 7B). This
consensus tree shows a polytomy at the base of Catharti-
formes composed of Pampagys, Dryornis, Gymnogyps, Sar-
coramphus, Vultur and Cathartes + Coragyps.
DISCUSSION
As mentioned in the description, a number of morpho-
logical features of the new material support the inclusion
of this taxon among the vultures, although it undoubtedly
shares some characteristics with true condors. The phylo-
genetic analysis presented here indicates that Dryornis
pampeanus is not a condor as originally proposed, but a
giant vulture. It is located at the base of the clade that
brings together all living vultures (Coragyps + Cathartes),
as their sister taxon. An estimated mass of c. 26 kg posi-
tions Dryornis as the largest Cathartidae that ever existed,
at almost double the mass of the largest extant cathartid,
the Andean condor Vultur gryphus, and c. 10-fold more
than the mass of the largest living vulture, Coragyps. Its
basal position in the clade of the vultures suggests that
the body size had a notable decrease in the history of the
vultures (Fig. 7A). Other extinct soaring birds also
reached large sizes, such as Pelagornis chilensis Mayr &
Rubilar-Rogers, 2010, which was estimated at 16–29 kg,
and the Teratornithidae Argentavis magnificens Campbell
& Tonni, 1980, which was estimated at 70 or 80 kg
(Campbell & Tonni 1981; Vizca
ıno & Fari~ na 1999;
Palmqvist & Vizca
ıno 2003; Chatterjee et al. 2007).
Among extant flying birds, the Kori bustard Ardeotis kori
is the heaviest, reaching 19 kg (Collar 1996).
Condors are the most frequent representatives in the
Cathartidae fossil record of South America. Condors
achieved a remarkable diversity during the Pleistocene,
which was possibly sustained by the high availability of
large mammals that would have served as food. This is in
contrast to what happened in North America, where both
vultures (Breagyps, Cathartes and Coragyps) and condors
(Hadrogyps, Pliogyps, Aizenogyps, Geronogyps and Gymno-
gyps) were equally diverse.
Another group of soaring scavengers that originated
in South America is the extinct Teratornithidae (Miller
1925; Fisher 1945; Howard 1962). This group of birds
has been recorded from the Oligocene to the Pleis-
tocene in the Americas (Tambussi & Degrange 2013).
Although highly diversified in the Pleistocene of North
America, its fossil record in South America is quite
scarce and is restricted to Taubatornis campbelli, the
oldest and smallest teratorn from the late Oligocene to
early Miocene of S~ao Paulo, in Brazil (Olson & Alvar-
enga 2002); Argentavis magnificens, the largest teratorn,
from the late Miocene of Salinas Grandes de Hidalgo,
in La Pampa province (Cenizo et al. 2012) and from
the Andalhuala Formation in Catamarca province
(Campbell 1995); an undetermined species from the
early late Miocene of Villarino, Buenos Aires province
(De Mendoza & Picasso 2019); and fragmentary
postcranial remains from the Pleistocene of Buenos
Aires province (Cenizo et al. 2016). Putative remains
also come from the late Pleistocene of Ecuador
10 PAPERS IN PALAEONTOLOGY

FIG. 6. Ulna and radius of Dryornis pampeanus MMP 5731 (A–E, K–O) and MMP 4679 (F–J). A–E, proximal fragment in: A, dorsal;
B, ventral; C, medial; and D, lateral view; E, proximal view. F–J, distal fragment in: F, dorsal; G, ventral; H, medial; and I, lateral view;
J, distal view. K–O, radius in: K, dorsal; L, ventral; M, medial; N, lateral; and O, distal view. Abbreviations: cd, cotyla dorsalis; cdu,
condylus dorsalis ulnaris; ch, cotyla humeralis; ci, crista intercotylaris; cv, cotyla ventralis; cvu, condylus ventralis ulnaris; dr, depressio
radialis; ims, impressio m. scapulotricipitis; it, incisura tendinosa; itc, incisura tuberculi carpalis; lcd, labrum condyli dorsalis; mht, m.
humerotriceps scar; ol, olecranon; pcd, processus cotylaris dorsalis; tbr, tuberculum bicipitale radii; tc, tuberculum carpale. Scale bars
represent 1 cm.
 DEGRANGE ET AL.: MORPHOLOGY AND PHYLOGENY OF DRYORNIS
TABLE 1. Measurements of the new specimen MMP 5731,
referred to Dryornis pampeanus.
Element Measurement MMP 5731
(mm)
Scapula Proximal width 31.1
Facies articularis humeralis length 16.1
Facies articularis humeralis height 10.5
Coracoid Facies articularis humeralis length 11.6
Facies articularis humeralis height 28.48
Lateromedial width of the shaft 12.6
Dorsoventral width of the shaft 10.1
Distal width 36.0
Humerus Proximal width 64.7
Distal width 56.3
Total length 339.7
Lateromedial width of the shaft 25.4
Dorsoventral width of the shaft 24.8
Ulna Proximal width 38.7
Distal width* 29.2
Radius Proximal width 16.7
*Measured on MMP4679.
(Campbell & Tonni 1980, 1983). Currently, the only
record in the Chapadmalalan is a fragmentary radius
from Bajada de Las Palomas, Buenos Aires province,
which was tentatively assigned to Teratornithidae
(Agnol
ın 2016). The phylogenetic position of this fam-
ily is still controversial. They were traditionally consid-
ered as related to Cathartidae (Brodkorb 1964; Olson
1985; Mayr 2009), and even Emslie (1988) proposed
Teratornithidae as the sister group of Cathartidae
based on a phylogenetic analysis. Although alternative
relationships with other orders such as Pelecaniformes,
Ciconiiformes and even Procellariiformes have been
proposed (Campbell & Tonni 1980; Cenizo et al. 2012;
De Mendoza & Picasso 2019), this taxon is currently
considered as incerti ordinis (Su
arez & Olson 2009; De
Mendoza & Picasso 2019).
The ability to soar is primarily determined by environ-
mental factors such as the presence of upward air cur-
rents of deflection and thermal convection. The coastal
marine location of Monte Hermoso and Mar del Plata,
plus the proximity to two mountain ranges in the Pam-
pean region (Tandilia and Ventania) in Pliocene times,
make it likely that the environmental conditions were
suitable for gliding flight. Assuming that the Monte Her-
moso and Chapadmalal Formations, at least at some
point, would have been contemporaneous, it can be
inferred that both localities, currently located 400 km
apart, would have been part of the Dryornis distribution
area. As an example, the Andean condor has a home
range of 10 000 km2 (Mandel et al. 2008), while the
11
turkey vulture has a home range of c. 900 km2 (Houston
et al. 2011; Dodge et al. 2014).
Scavenging is a well-documented tactic for food acquisi-
tion in diverse terrestrial animals, and the Cathartidae are
a quintessential example of scavengers. They are true obli-
gate scavengers, in that they are dependent on finding car-
rion on which to feed. Carrion can refer to any size of
animal carcass, and it is not unreasonable to imagine that
there was a high availability of vertebrates of small to large
size. Carcass availability is the second (and a strong) limi-
tation on the distribution and home range of scavengers
(Tyrberg 2008; Guti
errez-C
anovas et al. 2020). Vizca
ıno
et al. (2004) estimated Chapadmalalan vertebrate faunal
biomass distribution across three main categories, in which
12–17.6% of animals were between 10 and 100 kg, and
22–32.4% of animals were above 100 kg. This fauna pro-
vided plentiful carrion resources for scavenging without
the need to travel far, and therefore the area in which the
scavenger was able to live is relatively small (Dodge et al.
2014). In recent savanna biomes, the high number and
diversity of herbivores is mirrored by the great variety of
predators and scavengers. The Chapadmalalan biome had
a great diversity of herbivores but a very low variety of
predators (Prevosti et al. 2011) and scavengers. Taking
into account that the record of Teratornithidae in the Cha-
padmalalan is fragmentary and controversial, at the
moment Dryornis pampeanus is the only obligate scavenger
bird known in this faunal assemblage. A very interesting
fact is that a bird such as the black-headed vulture Cor-
agyps atratus may ingest c. 3 kg of carrion daily. Clearly, it
is impossible to know what the needs of a bird with an
estimated mass of 26 kg were, and whether the scavenging
activity of this bird on its own was sufficient for the
removal of the available carcasses.
CONCLUSION
Based on our phylogenetic analysis, and contrary to pre-
vious suggestions, Dryornis is more closely related to vul-
tures than to condors. The present body mass estimation
indicates that Dryornis is the largest of the Catharti-
formes, and suggests that body mass tends to decrease in
the vulture lineage.
The flight capability of Dryornis and the action of
external factors, such as thermal flows, would have
allowed travel between the two sites, which would have
been only 400 km apart. The existence of this taxon as a
common taxon in the Monte Hermoso and Chapadmalal
Formations reduces the faunal differences between the
two formations and reopens an old biostratigraphical
controversy. The absence of absolute dating for the Monte
Hermoso Formation and for the lower levels of the Cha-
padmalal Formation keep the controversy open, and new
12 PAPERS IN PALAEONTOLOGY

FIG. 7. Phylogenetic relationships of Dryornis pampeanus. A, phylogenetic analysis without Pampagyps imperator. B, phylogenetic
analysis with P. imperator. Note that the inclusion of P. imperator results in a polytomy at the base of Cathartiformes. Bremer support
values are shown above the branches of Cathartiformes, and mass (in kg) is given with the silhouettes.

biostratigraphical, palaeobiogeographical and palaeoecologi-
cal studies are required to determine whether the faunal
differences are due to geographical or ecological factors,
or to temporal asynchrony, or a combination of them.
Dryornis pampeanus was the only scavenger in the Cha-
padmalalan assemblage. It is striking that so far it is the
only single scavenger species to have been recognized in
an area with such a high available biomass. One possible
answer is that the carcass removal activity was undertaken
by other, non-avian, scavengers. Another explanation
could simply be that the fossil record is incomplete. There
has been much discussion about how communities func-
tion in terms of carrion consumption (Guti
errez-C
anovas
et al. 2020). This allows us to affirm that Dryornis was a
key dominant scavenger, but much remains to be learned
before we can comment on how the Chapadmalal com-
munities functioned.
Acknowledgements. Marcelo Reguero (MLP) and Paul Sweet
(AMNH) are thanked for granting access to the collections
under their care. We also thank Guilherme Renzo Rocha Brito
and Emilia Sferco for their assistance in preliminary phylo-
genetic analyses; and Editor Philip Mannion, and Federico
Agnol
ın and an anonymous reviewer for their helpful com-
ments and suggestions. We are grateful to CONICET for per-
manent support. This work was supported by PICT 2014-2330
(to CPT), and PUE 2016-CONICET-CICTERRA.
DATA ARCHIVING STATEMENT
Data for this study are available in MorphoBank: https://doi.org/10.
7934/P3896
Editor. Philip Mannion
 DEGRANGE ET AL.: MORPHOLOGY AND PHYLOGENY OF DRYORNIS
REFERENCES
A G N O L
IN , F. L. 2016. A brief history of South American
birds. Contribuciones del MACN, 6, 157–172.
- B R I S S ON
E G L I , F., S O I B E L Z O N , E., R O D R I -
G U E Z , S. G., S O I B E L Z O N , L. H., I A C O N A , F. and
P I A Z Z A , D. 2017. A new large Cathartidae from the quater-
nary of Argentina, with a review of the fossil record of con-
dors in South America. Contribuciones del MACN, 7, 1–16.
A L V A R E N G A , H. M. F. and O L S O N , S. L. 2004. A new
genus of tiny condor from the Pleistocene of Brazil (Aves:
Vulturidae). Proceedings of the Biological Society of Washington,
117, 1–9.
A L V A R E N G A , H., B R I T O , G. R. R., M I G O T T O , R., H U -
B B E , A. and H OF€ L I N G , E. 2008. Pleistovultur nevesi gen. et
sp. nov. (Aves: Vulturidae) and the diversity of condors and
vultures in the South American Pleistocene. Ameghiniana, 45,
613–618.
B A U M E L , J. J. and W I T M E R , L. M. 1993. Osteologia. 45–
132. In B A U M E L , J. J., K I N G , A. S., B R E A Z I L E , J. E.,
E V A N S , H. E. and V A N D E N B E R G E , J. C. (eds) Hand-
book of avian anatomy: Nomina anatomica avium. Publications
of the Nuttall Ornithological Club, 23, 779 pp.
B E C K , R. and T A G L I O R E T T I , M. L. 2019. A nearly com-
plete juvenile skull of the marsupial Sparassocynus derivatus
from the Pliocene of Argentina, the affinities of “sparasso-
cynids”, and the diversification of opossums (Marsupialia;
Didelphimorphia; Didelphidae). Journal of Mammalian Evolu-
tion, 27, 385–417.
B R I T O , G. R. R. 2008. An
alise filogen
etica de Cathartidae
(Aves) com base em caracteres osteol
ogicos. Unpublished PhD
thesis, Universidad de S~ao Paulo, Brazil.
B R O D K O R B , P. 1964. Catalogue of fossil birds: part 2 (Anser-
iformes). Bulletin of Florida State Museum, 8, 195–335.
- 1967. Catalogue of fossil birds, part III (Ralliformes,
Ichthyornithiformes, Charadriiformes). Bulletin of Florida State
Museum, 2, 99–220.
C A M P B E L L , K. E. Jr 1979. The non-passerine Pleistocene avi-
fauna of the Talara Tar Seeps, northwestern Peru. Life Science
Contribution, Royal Ontario Museum, 118, 1–203.
-1995. Additional specimens of the Giant Teratorn, Argen-
tavis magnificens, from Argentina (Aves: Teratornithidae).
Courier Forschungsinstitut Senckenberg, 181, 199–201.
-and T O N N I , E. P. 1980. A new genus of teratorn from
the Huayquerian of Argentina (Aves: Teratornithidae). Natural
History Museum of Los Angeles County, Contributions in
Science, 330, 59–68.
- - 1981. Preliminary observations on the paleobiology
and evolution of teratorns (Aves: Teratornithidae). Journal of
Vertebrate Paleontology, 1, 265–272.
- -1983. Size and locomotion in teratorns (Aves, Tera-
tornithidae). The Auk, 100, 390–403.
C E N I Z O , M., T A M B U S S I , C. P. and M O N T A L V O , C. I.
2012. Late Miocene continental birds from the Cerro Azul for-
mation in the Pampean region (central-southern Argentina).
Alcheringa, 36, 47–68.
-N O R I E G A , J. I., T O M A S S I N I , R. and T A S S A R A , D.
2016. First records of teratorns (Aves, Teratornithidae) in the
13
Pleistocene of South America. 7. In Proceedings of the 9th
International Meeting of the Society of Avian Paleontology and
Evolution.
C H A T T E R J E E , S., T E M P L I N , R. J. and C A M P B E L L , K.
E. Jr 2007. The aerodynamics of Argentavis, the world’s largest
flying bird from the Miocene of Argentina. Proceedings of the
National Academy of Sciences, 104, 12398–12403.
C I O N E , A. L. and T O N N I , E. P. 1995a. Chronostratigraphy
and “Land-Mammal Ages” in the Cenozoic of southern South
America: principles, practices, and the “Uquian” problem.
Journal of Paleontology, 69, 135–159.
- -1995b. Los estratotipos de los pisos Montehermosense y
Chapadmalalense (Plioceno) del esquema cronol
ogico sudameri-
cano. Ameghiniana, 32, 369–374.
- -2005. Bioestratigraf
ıa basada en mam
ıferos del Ceno-
zoico superior de la provincia de Buenos Aires, Argentina.
183–200. In D E B A R R I O , R. E., E T C H E V E R R Y , R. O.,

M. F. and L L A M B
IA S , E. (eds) Geolog
ıa y
C A B A L L E,
recursos minerales de la Provincia de Buenos Aires. Relatorio
del XVI Congreso Geol
ogico Argentino, 11, 480 pp.
- G A S P A R I N I , G. M., S O I B E L Z O N , E., S O I B E L -
Z O N , L. H. and T O N N I , E. P. 2015. The Great American bio-
tic interchange: A South American perspective. Springer, 97 pp.
C O L L A R , J. 1996. Family Otididae (Bustards). 240–273. In
D E L H O Y O , J., E L L I O T , A. and S A R G A T A L , J. (eds)
Handbook of birds of the world. Vol. 3: Hoatzin to auks. Lynx,
Barcelona, 821 pp.
C O U E S , E. 1894. Key to North American birds. Estes & Lauriat,
Boston, 907 pp.
C R U Z , L. E., F E R N Í C O L A , J. C., T A G L I O R E T T I , M. L.
and T O L E D O , N. 2016. A reassessment of the taxonomic
status of Paraglyptodon (Mammalia, Cingulata, Glyptodonta).
Journal of South American Earth Sciences, 66, 32–40.
D E M E N D O Z A , R. S. and P I C A S S O , M. B. J. 2019. A tera-
torn (Aves: Teratornithidae) from the early late Miocene of
Buenos Aires, Argentina. Historical Biology, published
online 26 June. https://doi.org/10.1080/08912963.2019.1634
706
D E G R A N G E , F. J., T A M B U S S I , C. P., T A G L I O R E T T I ,
M. L., D O N D A S , A. and S C A G L I A , F. 2015. A new Mesem-
briornithinae (Aves, Phorusrhacidae) provides new insights into
the phylogeny and sensory capabilities of terror birds. Journal of
Vertebrate Paleontology, 35, e912656.
--- and S C A G L I A F. 2021. Data from: Phylo-
genetic affinities and morphology of the Pliocene cathartiform
Dryornis pampeanus Moreno & Mercerat. MorphoBank, P3896.
https://doi.org/10.7934/p3896
D E S C H A M P S , C. M., V U C E T I C H , M. G., V E R Z I , D. H.
and O L I V A R E S , A. I. 2012. Biostratigraphy and correlation
of the Monte Hermoso Formation (early Pliocene, Argentina):
the evidence from caviomorph rodents. Journal of South Amer-
ican Earth Sciences, 35, 1–9.
D O D G E , S., B O H R E R , G., B I L D S T E I N , K., D A V I D -
S O N , S. C., W E I N Z I E R L , R., B E C H A R D , M. J., B A R -
B E R , D., K A Y S , R., B R A N D E S , D., H A N , J. and
W I K E L S K I , M. 2014. Environmental drivers of variability in
the movement ecology of turkey vultures (Cathartes aura) in
North and South America. Philosophical Transactions of the
Royal Society B, 369, 20130195.
 14 PAPERS IN PALAEONTOLOGY
D U N N I N G , J. B. 2008. CRC handbook of avian body masses.
Taylor & Francis, 672 pp.
E L Z A N O W S K I , A., B I E NK
O W S K A - W A S I L U K , M.,
C H O D Y N,
R. and B O G D A N O W I C Z , W. 2012. Anatomy
of the coracoid and diversity of the Procellariiformes (Aves)
in the Oligocene of Europe. Palaeontology, 55, 1199–1221.
E M S L I E , S. D. 1988. The fossil history and phylogenetic rela-
tionships of condors (Ciconiiformes: Vulturidae) in the New
World. Journal of Vertebrate Paleontology, 8, 212–228.
F A R R I S , J. S. 1969. A successive approximation approach to
character weighting. Systematic Zoology, 18, 374–385.
- 1989. The retention index and the rescaled consistency
index. Cladistics, 5, 417–419.
F I E L D , D. J., L Y N N E R , C., B R O W N , C. and D A R R O C H ,
S. A. 2013. Skeletal correlates for body mass estimation in
modern and fossil flying birds. PLoS One, 8, e82000.
F I S H E R , H. I. 1945. Locomotion in the fossil vulture Terator-
nis. The American Midland Naturalist, 33, 725–742.
G O L O B O F F , P. A. 1993. Estimating character weights during
tree search. Cladistics, 9, 83–92.
-and C A T A L A N O , S. A. 2016. TNT version 1.5, including
a full implementation of phylogenetic morphometrics. Cladis-
tics, 32, 221–238.

O V A S , C., M O L E ON

R E Z - C AN
G U T I ER
, M., M A T E O -
T O M AS
, P., O L E A , P. P., S E B A S T I AN

- G O N Z AL
EZ,
E. and S AN
C H E Z - Z A P A T A , J. A. 2020. Large home range
scavengers support higher rates of carcass removal. Functional
Ecology, 34, 1929–1932.
H A C K E T T , S. J., K I M B A L L , R. T., R E D D Y , S., B O W I E ,
R. C., B R A U N , E. L., B R A U N , M. J., C H O J N O W S K I , J.
L., C O X , W. A., H A N , K. L., H A R S H M A N , J., H U D -
D L E S T O N , C. J., M A R K S , B. D., M I G L I A , K. J.,
M O O R E , W. S., S H E L D O N , F. H., S T E A D M A N , D.
W., W I T T , C. C. and Y U R I , T. 2008. A phylogenomic
study of birds reveals their evolutionary history. Science, 320,
1763–1768.
H O L L A N D , A. E., B Y R N E , M. E., B R Y A N , A. L., D E V -
A U L T , T. L., R H O D E S , O. E. and B E A S L E Y , J. C. 2017.
Fine-scale assessment of home ranges and activity patterns for
resident black vultures (Coragyps atratus) and turkey vultures
(Cathartes aura). PLoS One, 12, e0179819.
H O U S T O N , C. S., M c L O U G H L I N , P. D., M A N D E L , J.
T., B E C H A R D , M. J., S T O F F E L , M. J., B A R B E R , D. R.
and B I L D S T E I N , K. L. 2011. Breeding home ranges of
migratory turkey vultures near their northern limit. The Wil-
son Journal of Ornithology, 123, 472–478.
H O W A R D , H. 1962. A comparison of prehistoric avian assem-
blages from individual pits at Rancho La Brea, California.
Natural History Museum of Los Angeles County, Contributions
in Science, 58, 1–24.
J A R V I S , E. D., M I R A R A B , S., A B E R E R , A. J., L I , B.,
H O U D E , P., L I , C., H O , S. Y. W., F A I R C L O T H , B. C.,
N A B H O L Z , B., H O W A R D , J. T., S U H , A., W E B E R , C.
C., D A F O N S E C A , R. R., L I , J., Z H A N G , F., L I , H.,
Z H O U , L., N A R U L A , N., L I U , L., G A N A P A T H Y , G.,
B O U S S A U , B., B A Y Z I D , S., Z A V I D O V Y C H , V.,
S U B R A M A N I A N , S., G A B A L D ON
, T., C A P E L L A -

R E Z , S., H U E R T A - C E P A S , J., R E K E P A L L I ,
G U T I ER
B., M U N C H , K., S C H I E R U P , M., L I N D O W , B., W A R -
R E N , W. C., R A Y , D., G R E E N , R. E., B R U F O R D , M.
W., Z H A N , X., D I X O N , A., L I , S., L I , N., H U A N G , Y.,
D E R R Y B E R R Y , E. P., B E R T E L S E N , M. F., S H E L D O N ,
F. H., B R U M F I E L D , R. T., M E L L O , C. V., L O V E L L , P.
V., W I R T H L I N , M., S C H N E I D E R , M. P. C., P R O S -
D O C I M I , F., S A M A N I E G O , J. A., V A R G A S V E L A Z -
Q U E Z , A. M., A L F A R O - N U
NE
~ Z , A., C A M P O S , P. F.,
P E T E R S E N , B., S I C H E R I T Z - P O N T E N , T., P A S , A.,
B A I L E Y , T., S C O F I E L D , P., B U N C E , M., L A M B E R T ,
D. M., Z H O U , Q., P E R E L M A N , P., D R I S K E L L , A. C.,
S H A P I R O , B., X I O N G , Z., Z E N G , Y., L I U , S., L I , Z.,
L I U , B., W U , K., X I A O , J., Y I N Q I , X., Z H E N G , Q.,
Z H A N G , Y., Y A N G , H., W A N G , J., S M E D S , L., R H E -
I N D T , F. E., B R A U N , M., F J E L D S A , J., O R L A N D O ,
L., B A R K E R , F. K., J Ø N S S O N , K. A., J O H N S O N , W.,
K O E P F L I , K., O ’ B R I E N , S., H A U S S L E R , D., R Y D E R ,
O. A., R A H B E K , C., W I L L E R S L E V , E., G R A V E S , G. R.,
G L E N N , T. C., M c C O R M A C K , J., B U R T , D., E L L E -
G R E N , H., A L S T R OM € , P., E D W A R D S , S. V., S T A -
M A T A K I S , A., M I N D E L L , D. P., C R A C R A F T , J.,
B R A U N , E. L., W A R N O W , T., J U N , W., G I L B E R T , M.
T. P. and Z H A N G , G. 2014. Whole-genome analyses resolve
early branches in the tree of life of modern birds. Science, 346,
1320–1331.
J O H N S O N , J. A., B R O W N , J. W., F U C H S , J. and M I N -
D E L L , D. P. 2016. Multi-locus phylogenetic inference among
New World Vultures (Aves: Cathartidae). Molecular Phylo-
genetics & Evolution, 105, 193–199.
K L U G E , A. 1997. Testability and the refutation and corrobora-
tion of cladistics hypotheses. Cladistics, 13, 81–96.
K R A G L I E V I C H , J. L. 1952. El perfil geol
ogico de Chapad-
malal y Miramar, Provincia de Buenos Aires. Revista Museo
Municipal Ciencias Naturales y Tradicional Mar del Plata, 1,
8–37.
L A F R E S N A Y E , N. F. 1839. Nouvelle classification des oiseaux
de proie ou rapaces. Revue Zoologique par la Soci
et
e Cuvi
eri-
enne, 193–196.
L I N N A E U S , C. V. 1758. Systema naturae per regna tria natu-
rae. Tenth edition. Laurentii Salvii, Stockholm, 824 pp.
M A N D E L , J. T., B I L D S T E I N , K. L., B O H R E R , G. and
W I N K L E R , D. W. 2008. Movement ecology of migration in
turkey vultures. Proceedings of the National Academy of
Sciences, 105, 19102–19107.
M A Y R , G. 2009. Paleogene fossil birds. Springer, 262 pp.
-and R U B I L A R - R O G E R S , D. 2010. Osteology of a new
giant bony-toothed bird from the Miocene of Chile, with a
revision of the taxonomy of Neogene Pelagornithidae. Journal
of Vertebrate Paleontology, 30, 1313–1330.
M I L L E R , L. 1925. The birds of Rancho La Brea. Carnegie Insti-
tute of Washington Publication, 349, 63–106.
M O R E N O , F. P. and M E R C E R A T , A. 1891. Cat
alogo de los
p
ajaros f
osiles de la Rep
ublica Argentina conservados en el
Museo de La Plata. Anales del Museo de La Plata, 1, 7–71.
N O R I E G A , J. I. and T O N N I , E. P. 2007. Geronogyps reliquus
Campbell (Ciconiiformes: Vulturidae) en el Pleistoceno tard
ıo
de la provincia de Entre R
ıos y su significado paleoambiental.
Ameghiniana, 44, 245–250.
 DEGRANGE ET AL.: MORPHOLOGY AND PHYLOGENY OF DRYORNIS
O L S O N , S. L. 1985. The fossil record of birds. 79–252. In
F A R N E R , D., K I N G , J. and P A R K E S , K. (eds) Avian biol-
ogy. Academic Press, 256 pp.
-and A L V A R E N G A , H. M. F. 2002. A new genus of small
teratorn from the Middle Tertiary of the Taubate Basin, Brazil
(Aves: Teratornithidae). Proceedings of the Biological Society of
Washington, 115, 701–705.
P A L M Q V I S T , P. and V I Z C A
IN O , S. F. 2003. Ecological
and reproductive constraints of body size in the gigantic
Argentavis magnificens (Aves, Teratornithidae) from the Mio-
cene of Argentina. Ameghiniana, 40, 379–385.
P A S C U A L , R. and O D R E M A N R I V A S , O. E. 1973. Las
unidades estratigr
aficas del Terciario portadoras de mam
ıferos.
Su distribuci
on y sus relaciones con los acontecimientos
distr
oficos. Anales V Congreso Geol
ogico Argentino, 3, 293–338.
- O R T E G A H I N O J O S A , E. J., G O N D A R , D. and
T O N N I , E. P. 1965. Las edades del Cenozoico mamal
ıfero
de la Argentina, con especial atenci
on a aquellas del territorio
bonaerense. Anales Comisi
on Investigaciones Cient
ıficas Buenos
Aires, 6, 165–193.
P A T T E R S O N , B. and K R A G L I E V I C H , L. 1960. Sistem
atica
y nomenclatura de las aves fororracoideas del Plioceno Argen-
tino. Publicaci
on del Museo Municipal Ciencias Naturales y
Tradicionales de Mar del Plata, 1, 1–51.
P R E V O S T I , F. J., F O R A S I E P I , A. and Z I M I C Z , N. 2011.
The evolution of the Cenozoic terrestrial mammalian predator
guild in South America: competition or replacement? Journal
of Mammalian Evolution, 20, 3–21.
P R U M , R. O., B E R V , J. S., D O R N B U R G , A., F I E L D , D.
J., T O W N S E N D , J. P., L E M M O N , E. M. and L E M -
M O N , A. R. 2015. A comprehensive phylogeny of birds
(Aves) using targeted next-generation DNA sequencing. Nat-
ure, 526, 569–573.
P Y C R A F T , W. P. 1900. The morphology and phylogeny of
the Palaeognathae (Ratitae and Crypturi) and the Neog-
nathae (Carinatae). Transactions of the Zoological Society, 15,
149–290.
R O D R
IG U E Z B R I Z U E L A , R. 2004. Registro de Vulturidae
(Aves, Ciconiiformes) en el Ne
ogeno de la provincia de la
Rioja, Argentina. Revista del Museo Argentino de Ciencias Nat-
urales, Nueva Serie, 6, 307–311.
S C H U L T Z , P. H., Z AR
A T E , M. A., H A M E S , W., C A M -

, C. and K I N G , J. 1998. A 3.3-Ma impact in Argen-
I L I ON
tina and possible consequences. Science, 282, 2061–2063.
S T U C C H I , M. and E M S L I E , S. M. 2005. A new condor
(Ciconiiformes, Vulturidae) from the late Miocene/early Plio-
cene Pisco Formation, Peru. Condor, 107, 107–113.
- - V A R A S - M A L C A , R. M. and U R B I N A -
S C H M I T T , M. 2015. A new late Miocene condor (Aves,
Cathartidae) from Peru and the origin of South American
condors. Journal of Vertebrate Paleontology, 35, e972507.
S U Á R E Z , W. and O L S O N , S. L. 2009. A new genus for the
Cuban teratorn. Proceedings of the Biological Society of Wash-
ington, 122, 103–116.
T A G L I O R E T T I , M. L., M I NO ~ - B O I L I N I , A. R., S C A -
G L I A , F. and D O N D A S , A. P. 2014. Presencia de Prosceli-
dodon patrius (Xenarthra, Scelidotheriinae) en la Formaci
on
Chapadmalal (Plioceno superior), Mar del Plata, Buenos Aires,
15
Argentina: implicancias bioestratigr
aficas. Ameghiniana, 51,
420–427.
T A M B U S S I , C. P. 2011. Paleoenvironmental and faunal infer-
ences based upon the avian fossil record of Patagonia and
Pampa: what works and what does not. Biological Journal of
the Linnean Society, 103, 458–474.
- and D E G R A N G E , F. J. 2013. South American and
Antarctic continental Cenozoic birds: Paleobiogeographic affini-
ties and disparities. Springer, 113 pp.
- and N O R I E G A , J. I. 1996. Summary of the Avian fossil
record from southern South America. 245–264. In A R R A T I A ,
G. (eds) Contributions of southern South America to vertebrate
paleontology. M€ unchner Geowissenschaftliche Abhandlungen
30. Friedrich Pfeil, 340 pp.
-- 1999. The fossil record of condors (Ciconiiformes:
Vulturidae) in Argentina. Smithsonian Contributions to Paleon-
tology, 89, 177–184.
T O M A S S I N I , R. L. and M O N T A L V O , C. I. 2013. Tapho-
nomic modes on fluvial deposits of the Monte Hermoso Forma-
tion (early Pliocene), Buenos Aires province, Argentina.
Palaeogeography, Palaeoclimatology, Palaeoecology, 369, 282–294.
-- D E S C H A M P S , C. M. and M A N E R A , T. 2013.
Biostratigraphy and biochronology of the Monte Hermoso For-
mation (early Pliocene) at its type locality, Buenos Aires Pro-
vince, Argentina. Journal of South American Earth Sciences, 48,
31–42.
T O N N I , E. P. 1980. The present state of knowledge of the
Cenozoic birds of Argentina. Natural History Museum of
Los Angeles County, Contributions in Science, 330, 104–
114.
-and N O R I E G A , J. I. 1998. Los c
ondores (Ciconiiformes,
Vulturidae) de la regi
on pampeana de la Argentina durante el
Cenozoico tard
ıo: distribuci
on, interacciones y extinciones.
Ameghiniana, 35, 141–150.
-P R A D O , J., F I D A L G O , F. and L A Z A , J. 1992. El Piso/
Edad Montehermosense (Plioceno) y sus mam
ıferos. Jornadas
Geol
ogicas Bonaerenses, 3, 113–118.
T Y R B E R G , T. 2008. The late Pleistocene continental avian
extinction: an evaluation of the fossil evidence. Oryctos, 7,
249–269.
V I Z C A
IN O , S. F. and F A R I N A , R. A. 1999. On the flight
capabilities and distribution of the giant Miocene bird
Argenvtavis magnificens (Teratornithidae). Lethaia, 32, 271–
278.
-F A R I NA ~ , R. A., Z AR
A T E , M. A., B A R G O , M. S. and
S C H U L T Z , P. 2004. Palaeoecological implications of the
mid-Pliocene faunal turnover in the Pampean Region (Argen-
tina). Palaeogeography, Palaeoclimatology, Palaeoecology, 213,
101–113.
W E T M O R E , A. 1927. Fossil birds from the Oligocene of Col-
orado. Proceedings of the Colorado Museum of Natural History,
7, 1–14.
Z A M O R A N O , M., T A G L I O R E T T I , M., Z U R I T A , A. E.,
S C I L L A T O - Y A N É , G. J. and S C A G L I A , F. 2014. El reg-
istro m
as antiguo de Panochthus (Xenarthra, Glyptodontidae).
Estudios Geol
ogicos, 70, 5–11.

A T E , M. 2005. El Cenozoico tard
ıo continental de la
Z AR
provincia de Buenos Aires. 139–158. In D E B A R R I O , R. E.,
 16 PAPERS IN PALAEONTOLOGY
E T C H E V E R R Y , R. O., C A B A L L É , M. F. and L L A M -
B Í A S , E. (eds) Geolog
ıa y recursos minerales de la Provincia de
Buenos Aires. XVI Congreso Geol
ogico Argentino: Relatorio.
Asociaci
on Geol
ogica Argentina, Buenos Aires.
-and F A S A N O , J. L. 1989. The Plio-Pleistocene record of
Central Eastern Pampas, Buenos Aires Province, Argentina:
the Chapadmalal case study. Palaeogeography, Palaeoclimatol-
ogy, Palaeoecology, 72, 27–52.
Z U R I T A , A. E., T A G L I O R E T T I , M., D E L O S R E Y E S , M.,
O L I V A , C. and S C A G L I A , F. 2014. First Neogene skulls of
Doedicurinae (Xenarthra, Glyptodontidae): morphology and
phylogenetic implications. Historical Biology, 28, 423–432.
- - - - -2016a. First Neogene skulls of Doedi-
curinae (Xenarthra Glyptodontidae): morphology and phylo-
genetic implications. Historical Biology, 28, 423–432.
--- C U A D R E L L I , F. and P O I R E , D. 2016b.
Regarding the real diversity of Glyptodontidae (Mammalia,
Xenarthra) in the late Pliocene (Chapadmalalan age/stage) of
Argentina. Anais da Academia Brasileira de Ci^encias, 88,
809–827.