Biological Journal of the Linnean Society, 2011, 103, 458–474.

With 5 figures

Palaeoenvironmental and faunal inferences based on

the avian fossil record of Patagonia and Pampa: what
works and what does not
CLAUDIA P. TAMBUSSI*
CONICET, División Palaeontología Vertebrados, Museo de La Plata, Paseo del Bosque, 1900 La
Plata, Argentina

Received 28 February 2011; accepted for publication 28 February 2011 bij_1658 458..474

Analysing the effect of climatic/environmental changes on bird communities during the South American Cenozoic
is quite complicated. Taking into consideration the extremely complex evolution of such environmental conditions
and the incomplete and episodic fossil bird record in this part of the continent, any generalization should be
considered with caution. However, some aspects may be noted: (1) certain typically South American bird groups
evolved in total isolation, i.e. terrestrial or poorly flying birds, incapable of crossing important water barriers
(Rheiformes, Tinamiformes, Phorusrhacidae, Brontornithidae, Anhimidae); (2) other good flyers did not cross until
immediately before the definitive connection between both Americas (Teratornithidae, Passeriformes Suboscines);
(3) most of the families established important intercontinental relationships (Anhingidae, Pelecanidae, Ciconiidae,
Anatidae, Presbyornithidae, Rallidae, Falconidae and Accipitridae); (4) several taxa that are presently important
members of the rich South American bird fauna are unknown for certain geological time periods (Throchilidae); and
(5) there is a high prevalence of carnivorous birds over all other trophic habits, regardless of the association or age
analysed. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474.

ADDITIONAL KEYWORDS: Aves – biogeography – Cenozoic – faunal associations – palaeoenvironments.

Analizar el efecto que los cambios climáticos y ambientales tuvieron en las comunidades de aves durante el
Cenozoico sudamericano es complicado y cualquier generalización debe tomarse con cautela. Sin embargo pueden
señalarse algunos aspectos: (1) algunos grupos de aves típicamente sudamericanas evolucionaron en total
aislamiento (Rheiformes, Tinamiformes, Phorusrhacidae, Brontornithidae, Anhimidae); (2) otros grupos buenos
voladores no cruzaron a América del Norte hasta establecido el puente Panameño entre las dos Américas
(Teratornithidae, Passeriformes Suboscines); (3) la mayoría de las familias establecieron importantes relaciones
intercontinentales (Anhingidae, Pelecanidae, Ciconiidae, Anatidae, Presbyornithidae, Rallidae, Falconidae, Accipi-
tridae); (4) importantes miembros de la avifauna sudamericana actual son desconocidos en el registro fósil
(Throchilidae); (5) hay una prevalencia de aves carnívoras en todas las asociaciones cualquiera fuere su antigüedad.

PALABRAS CLAVE: Aves – biogeografía – Cenozoico – Asociaciones faunísticas – palaeoambientes.

INTRODUCTION stunted and dwarf plants; in the valleys the same thorn-
bearing bushes grow. Everywhere we see the same birds and
The country remains the same: terribly uninteresting. The insects []. The curse of sterility is on the land.
great similarity in productions is a very striking feature in all Charles Darwin’s Diary of the Voyage of H.M.S. Beagle,
Patagonia. The level plains of arid shingle support the same April 22nd, 1834

Patagonia is currently an arid (yearly precipitations

*E-mail: [email protected] under 300 mm), cold (mean annual temperature,

458 © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
 PATAGONIAN AND PAMPEAN FOSSIL BIRDS 459

12 °C) and windy territory with low-growing, steppe- sentatives of the earliest Neogene. These exposures
adapted shrub vegetation. Only towards the west, in are emblematic as they have been known since the
the vicinity of the Andean mountain range, do pre- late 19th century thanks to the work of the brothers
cipitations become more abundant and allow the Carlos and Florentino Ameghino, the former recog-
development of forests with dense undergrowth nized by his essential field work and the latter by his
(Barreda & Palazzesi, 2007) and more abundant and vast written production (Ameghino, 1891, 1895, 1906,
varied vegetation. According to phytogeographical 1908).
and ecological criteria, Argentina can be divided into Accordingly, the oldest Neornithes (or modern)
18 eco-regions, seven of which are represented in birds from Patagonia (and the whole of Argentina)
Patagonia and in the marine coastal sectors and come from the San Jorge Gulf Basin (Chubut and
neighbouring islands (Fig. 1). Each of these eco- Santa Cruz Provinces), corresponding fundamentally
regions is also characterized by the presence of to the Riochican (middle to late Palaeocene), Casa-
certain bird species, both residents and visitors. mayoran (late Eocene) and Deseadan (late Oligocene)
Taken as a whole, Patagonia currently hosts about 40 periods (Fig. 1A), whereas the Neogene records origi-
nonpasseriform and 14 passeriform bird families nate from the Santacrucian (late early Miocene) of the
(Narovsky & Babarskas, 2000) with heterogeneous Austral Basin, up to the most modern Huayquerian
concentration in each eco-region, with the prevalence (late Miocene), located on the northern San Jorge
of marine birds near the Atlantic Ocean coast, low Gulf Basin, in Chubut Province (Dozo et al., 2010), as
bird diversity on the steppe and highest diversity in well as north of Patagonia, in La Pampa Province
the mountain forests. (Montalvo et al., 2008; M. N. Cenizo, C. P. Tambussi &
Modern-day Patagonia has been moulded by a C. Montalvo, unpubl. data). The vertebrate fossil
transformation process resulting from the interaction record in Patagonia is very scarce from the end of the
of global tectonics, with palaeogeographical and Miocene to the present. Information for this period
palaeoclimatic changes. Two events are key and may thus needs to be complemented with extra-
be considered as iconic for this transformation: first, Patagonian (Pampasian) records to better understand
the opening of the Drake Passage at some point the macro-events in the Patagonian region (Tonni &
during the Eocene–Oligocene transition (c. 28 Ma ago; Carlini, 2008).
Madden et al., 2010) isolated Antarctica from Patago- At least two marine transgressions affected the
nia, bringing about a trend towards cooler conditions entire South American continent along different cor-
(Reguero & Marenssi, 2010), and, second, the rise of ridors during the Miocene. The first transgression
the Andes mountain range during the Miocene, which took place between 15 and 13 Ma and was dominated
blocked the passage of the humid East winds, induced by eustatic and tectonic factors. The second transgres-
a process of desertification. The trend towards global sion took place between 10 and 5 Ma and resulted
cooling, which was already evident after the Mid- mainly from tectonic events (Hernandez et al., 2005).
Eocene Climate Optimum, became more marked Thus, the continental sediments of the Palaeogene
towards the Miocene, although with some inter- and Neogene are interdigitated with highly fossilifer-
spersed warmer periods, such as the event known as ous sediments of marine origin containing both ver-
the Middle Miocene Climate Optimum (Zachos et al., tebrates and invertebrates. Among birds, penguins
2001; Madden et al., 2010, and references cited are the most frequent (Acosta Hospitaleche et al.,
therein). According to Pascual & Odreman Rivas 2007, 2008). Marine sediments are represented in the
(1971), two areas can be recognized in Patagonia: to San Julián, Monte León and Gaiman Formations
the north, the San Jorge Gulf Basin extends between (late Eocene to early Miocene) and the more modern
the Somun Curá massif and the Deseado massif Puerto Madryn Formation (middle to late Miocene)
(Fig. 1B), and the Austral Basin to the south of the outcropping in the Valdés Peninsula (Dozo et al.,
Deseado massif (Tonni & Carlini, 2008). This scheme 2010). The greatest of the Miocene transgressions is
provides a useful basis for the arrangement of the known as the Paranaensean; it began in the latest
bird records discussed here. Oligocene and ended during the early middle Miocene
Although the San Jorge Gulf Basin is dominated (15–13 Ma). The final phases of this transgression
by Palaeogene sediments (Palaeocene–Oligocene, are coincident with the Neogene Climatic Optimum.
approximately between 65 and 23 Ma, which com- The Paranaensean covered much of the Chaco-
prises the first part of the Cenozoic era, according to Paranaensean lowlands and eastern Patagonia,
Gradstein, Ogg & Smith, 2004), the Austral Basin affecting the current territories of Argentina,
comprises mostly sediments from the initial part of Uruguay, parts of Brazil and Paraguay, southern
the Neogene (Tonni & Carlini, 2008). The wonderful Bolivia, Venezuela, Colombia and Ecuador.
exposures of the Santa Cruz Formation (late early Although the fossil record of birds in the Cenozoic is
Miocene) have undoubtedly provided the best repre- incomplete, scarce and consisting mostly of isolated

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460 C. P. TAMBUSSI

Figure 1. A, Palaeocene to Recent time scale following Gradstein et al. (2004) including time scale for Cenozoic
mammalian faunas of South America (SALMA). B, Map of Patagonia showing the present-day major ecosystems (1, High
Andes; 2, Araucanian Forest; 3, Steppe; 4, Shrub) and four important fossil localities used in the text (SAN, Santa Cruz
Formation at Santa Cruz Province; PEN, Puerto Madryn Formation at Penínusula Valdés; CA, Cerro Azul Formation at
La Pampa Province; CHA, Chapadmalal at Buenos Aires Province). C–E, Vegetation-type physiognomies under increas-
ingly drier and/or more markedly seasonal climates (modified from Barreda & Palazzesi, 2007): C, late Oligocene–early
Miocene; D, middle late Miocene; E, late Miocene.

bones (very exceptionally complete skeletons have with extant species were already represented at about
been exhumed; Tambussi & Noriega, 1996; Acosta 20 Ma (Miocene). The bird fossil record in Patagonia
Hospitaleche et al., 2007), it is sufficient to suggest provides the basis for support of this scenario of first
that many of the orders of living birds were present appearances of the major groups of Neornithes birds
around 33 Ma (Oligocene) and that nearly all families (the clade that includes all modern birds), the analy-

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
 PATAGONIAN AND PAMPEAN FOSSIL BIRDS 461

sis of which is the focus of this work. Many of the than at present (for a synthesis of the fossil record of
records, especially those from the Palaeogene, penguins in Patagonia, see Acosta Hospitaleche &
however, correspond to collections made during the Tambussi, 2008).
late 19th and early 20th centuries, and they lack The record of Phorusrhacidae also begins during the
reliable provenance information. Inferences made Eocene. In particular, Psilopterines from the late
from these materials thus have a restricted value. Eocene (Mustersan) were reported by Tambussi &
In spite of their small sample sizes and incomplete Acosta Hospitaleche (2005). The Phorusrhacidae are
preservation, fossil birds may be a source of valuable an extinct group of predatory terrestrial birds
palaeoecological and palaeoenvironmental informa- (Andrews, 1899; Sinclair & Farr, 1932; Tonni, 1980;
tion, especially when they form assemblages and are Tambussi & Noriega, 1996; Alvarenga & Höfling, 2003;
analysed in a general context. The focus of this study Bertelli, Chiappe & Tambussi, 2007; Degrange et al.,
is to interpret the palaeoenvironmental, palaeoeco- 2010). Traditional studies of phorusrhacids classify
logical and faunal conditions of the Cenozoic using them into five subgroups; in order of increasing size
the four best bird assemblages recovered from and body mass, these are the psilopterines, mesembri-
Neogene sediments of Patagonia and Pampa. ornithines, patagornithines, phorusrhacines and bron-
tornithines. This subject is described in detail below in
the section on birds found in the Patagonian Miocene.
According to Agnolin (2006b), the diurnal birds of
THE PALAEOGENE BIRDS OF PATAGONIA
prey (Accipitridae, buzzards and hawks) also have
Very little is known about the Palaeogene birds of their first records in the Patagonian Eocene. (See
Patagonia: in all cases they are poorly represented. Appendix for other Eocene records.)
With isolated fossil material, their phylogenetic affini- The Oligocene bird record is fragmentary and little
ties are uncertain or their relationships are in need of progress has been made since Ameghino’s original
exhaustive revision (Tambussi & Noriega, 1996; Mayr, assignations. As far as I know, there have been no
2009). Indeed, for the Palaeocene, the only known new records of this age. Agnolin (2004) attempted a
records are feather imprints collected from fluvial review, but the existing record is highly fragmentary
levels of the Salamanca Formation (the Danian– and of dubious affinities (see Appendix for a synthe-
Selandian boundary, c. 61 Ma, early Palaeocene; sis). Only the appearance of new and more complete
Degrange et al., 2006). This is the oldest evidence of records would allow appropriate reconstruction of the
Neornithes birds for Patagonia and also for all of South continental scenarios of the Oligocene in Patagonia.
America. However, an isolated phalanx from the foot of Psilopterus affinis (Ameghino), a species that seems
a member of Rheiformes, the taxon that includes the to be the smallest among phorusrhacid species, is
living rheas or South American ostrich-like birds safely recorded in the late Oligocene of the San Jorge
(Tambussi, 1995), was collected in the Río Chico For- Gulf Basin. The first records of the middle-sized pho-
mation (middle Palaeocene). Both records come from rusrhacids, Patagornithinae (Andrewsornis abbotti
the San Jorge Gulf Basin (see Appendix). Patterson) and the giant Brontornithinae (Physornis
From the same area, but exhumed from Eocene fortis Ameghino) also occur in the Oligocene.
sediments, a few birds from the Casamayor Forma- With regard to the marine sediments from the San
tion and probably middle Eocene (Casamayoran) in Julián area, in Santa Cruz Province (late Eocene–
age have been described (Tonni, 1980; Tambussi & early Oligocene), the remains of two mid- to large-
Noriega, 1996; Mayr, 2009; see also Agnolin, 2004, sized penguins have been exhumed: Paraptenodytes
2006a, b, 2007a, b) as well as some from the robustus (Ameghino) and Arthrodytes andrewsi
Sarmiento Formation (late Eocene, Mustersan; Tam- (Ameghino) (Acosta Hospitaleche & Tambussi, 2008)
bussi & Acosta Hospitaleche, 2005). (see Appendix).
The Eocene of Patagonia holds the first record of the
Presbyornithidae, a very particular family of aquatic
or semi-aquatic Anseriformes birds with a duck-like
THE NEOGENE OF PATAGONIA: THE
head and a body reminiscent of that of flamingos
SANTACRUCIAN BIRDS
(Tambussi & Noriega, 1999). The affinity of these birds
with the Anseriformes has been recognized since the It is not possible to refer to the Patagonian or South
1970s (Olson & Feduccia, 1980), and currently they are American Neogene without mentioning the birds from
considered as a sister group of the Anatidae (swans the late early Miocene of the Santa Cruz Formation
and ducks) (Ericson, 1997; Livezey, 1997). (Santacrucian). Known since the times of Ameghino,
Penguins constitute a second group with first Pat- the Santa Cruz Formation occurs in southern Patago-
agonian records during the Eocene (see Appendix). At nia and is the most widespread and most richly
that time, their species’ diversity was much higher fossiliferous of all Patagonian nonmarine Tertiary

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462 C. P. TAMBUSSI

Figure 2. Location maps of the fossil localities used in the text. A, Late early Miocene localities of Santa Cruz Formation
at Santa Cruz Province. B, Late Pliocene Chapadmalal locality at Buenos Aires Province. C, Late Miocene localities of
Puerto Madryn Formation at Peninsula Valdés, Chubut Province. D, Late Miocene localities of Cerro Azul Formation of
La Pampa Province.

formations of southern Argentina (Tauber, 1994, recognized two biozones within the Santa Cruz For-
1997a, b; Tejedor et al., 2006; Vizcaíno et al., 2006, and mation, stratigraphically represented by the lower
references cited therein). This formation outcropping Estancia La Costa Member and the upper Estancia La
at the Atlantic Ocean coast represents the continental Angelina Member (see also Croft, 2001).
facies after the regression of the Patagonian Sea (Tonni By the late Oligocene–early Miocene, forest ele-
& Carlini, 2008). The richest fossil localities lie along ments are gradually replaced by other shrubby and
the Atlantic Ocean coast of southern Patagonia from herbaceous forms (Fig. 1B–D), signalling the begin-
Monte León to Río Gallegos and along the valleys of the ning of the expansion of xerophytic environments
Chico, Coyle, Chalía, Santa Cruz and Gallegos streams determined by a cooling and drying trend manifested
(Fig. 2A). Its lower levels are dated as 19.33 and in areas to the east of the Andes (Barreda & Palazz-
20.18 Ma, whereas the upper levels are in the 16.18– esi, 2007), and Patagonia begins to acquire its modern
20.61 Ma range (Tejedor et al., 2006). Tauber (1997b) appearance. Some evidence supports the existence of

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
 PATAGONIAN AND PAMPEAN FOSSIL BIRDS 463

Figure 3. Top: climatic scenario during the late early Miocene (Santacrucian stage) of Patagonia. Birds and some
landmark mammals are included. Bottom: explanation of the figure: 1, Tinamidae; 2, Cariama santacrucensis;
3, Psilopterus lemoinei; 4, Anhingidae indet.; 5, Psilopterus bachmanni; 6, Phorusrhacos longissimus; 7, Thegornis debilis;
8, Protibis cnemialis; 9, Eutelornis patagonicus; 10, Kelenken guillermoi; 11, Astrapotherium magnum (Astrapotheriidae);
12, Arctodictis muñizi (Borhyaenidae); 13, Brontornis burmeisteri; 14, Badiostes patagonicus; 15, Cracidae indet.;
16, Opistodactylus patagonicus; 17, Thegornis musculosus; 18, Steiromys sp. (Erethizontidae); 19, Patagornis marshi;
20, Soriacebus sp. (Atelidae); 21, Ankonetta larriestrai. Drawings by Marcos Cenizo.

both open and closed environments (see Vizcaíno described in the Reports of the Princeton University
et al., 2010, and references cited therein). Birds also Expeditions to Patagonia (Sinclair & Farr, 1932).
reflect this process. However, most specimens are fragmentary. Some
The Aves remains of the Santa Cruz Formation are have recently been reanalysed (Alvarenga & Höfling,
spectacular because of their diversity and morphologi- 2003; Agnolin, 2004, 2006a, b, 2007b, 2009), but a
cal disparity (Tonni, 1980; Olson, 1985; Tambussi & thorough review is still pending (Noriega et al., 2009).
Noriega, 1996; Alvarenga & Höfling, 2003; Agnolin, The Appendix summarizes all known records to date,
2004, 2006a, b, 2007a, b, 2009; Noriega, Vizcaíno & which, although from different localities within the
Bargo, 2009; Cenizo & Agnolin, 2010). These materi- Santa Cruz Formation, are considered jointly in the
als were beautifully illustrated and reasonably well present paper (Fig. 3).

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
464 C. P. TAMBUSSI

Rheiformes are herbivorous/omnivorous ground estimation according to Jones, 2010). All phorusrha-
birds, inhabitants of open grasslands. They are mag- cids, whose larger forms are popularly known as ‘terror
nificent runners and their long necks and excellent birds’, are characterized by a very large and com-
vision provide them with an effective monitoring pressed skull, raptor-like beak and elongate leg bones.
system. Opisthodactylus patagonicus Ameghino, the The wings seem to be greatly reduced in all species,
only Santacrucian record for this family, does not ranging from marked reductions in the largest forms
differ greatly from that of the living Rheidae that would not have allowed them to fly, to more
(Tambussi, 1995). moderate reduction in which flight might have been
Records of Santacrucian tinamous (Tinamidae) possible (e.g. Psilopterinae; Tonni & Tambussi, 1988;
from the fossil localities of Monte Observación, Monte Tambussi & Acosta Hospitaleche, 2005; for an alterna-
León and Cañadón de las Vacas have been reported tive view, see Alvarenga & Höfling, 2003). With or
(Chiappe, 1991; Bertelli & Chiappe, 2002). At present, without flight capability, all phorusrhacids were large
the tinamous are a Neotropical group of palaeog- terrestrial, swift cursorial birds, and it is generally
naths. They are basically ground birds and poor fliers. assumed that they were among the top predators of the
Modern Neotropical faunas are highly diverse in Patagonian Cenozoic scenarios. A carnivorous lifestyle
anseriforms, particularly anatids, but fossil ducks are was recently tested from a biomechanical point of view
poorly known in Patagonia and South America. Their (Degrange et al., 2010) using the skull of the medium-
fossil record is characterized by isolated and poorly sized Patagornithinae Andalgalornis steulleti Kra-
informative specimens, with most species known glievich from the Pliocene of northwestern Argentina.
from their original diagnosis. The list of Patagonian Because terror birds have no close analogues among
anseriforms includes Eoneornis australis Ameghino modern-day birds, this study was essential to under-
(Anhimidae sensu Cenizo & Agnolin, 2010), Eutelornis stand the biology of this extinguished group. In a broad
patagonicus Ameghino (a basal anseriform) and sense, we may assume that the skulls of other large
Ankoneta larriestrai Cenizo & Agnolin, a mid-sized phorusrhacids, characterized by high and compressed
anatid considered to have superficial similarities with beaks, would respond similarly to that of Andalgalor-
the whistling ducks (i.e. Dendrocygna) (see Appendix). nis (body mass, 40 kg). Phorusrhacids chased and
The idea originally suggested by Moreno & Mer- killed their prey using their huge beaks as a hatchet
cerat (1891) and recently revived by Agnolin (2007b), (Degrange et al., 2010). They probably could have
that Brontornis burmeisteri is a basal anseriform, is swallowed small prey whole or, at least in some cases,
quite reasonable. Brontornis was traditionally placed dismembered large prey with the help of their strong
within phorusrhacids, as a representative of the neck muscles and the claw on digit II of the foot. For
‘graviportal’ habit in the family. Systematic issues most of the existence of phorusrhacids, South America
aside, Brontornis was undoubtedly the giant beast of was isolated (like a large island?) and inhabited by a
the Patagonian Miocene, reaching a standing height peculiar fauna: the nonavian predators were repre-
of over 2 m and 380 kg (body mass estimation accord- sented by marsupials and reptiles, whereas herbivores
ing to Jones, 2010). Alvarenga & Höfling (2003) (as in the rest of the world) were placental mammals.
hypothesized that brontornithines may have been The marsupials tended to be heavily limbed forms (e.g.
scavengers/kleptoparasites, but Agnolin (2007b) con- borhyaenids) and the predaceous reptiles were terres-
sidered them to be herbivorous. Further studies are trial crocodiles: both probably inhabited forests,
needed, especially focused on the mandible, to cor- whereas the large phorusrhacids tended to prefer more
roborate any of these hypotheses. open plains. However, there is no contrary evidence to
During the late early Miocene, phorusrhacids expe- support that the smaller psilopterines foraged in the
rienced the greatest diversification ever recorded (at woods. The Americas were connected at about 3 Ma,
least four species) and were common inhabitants of the Panamanian bridge was completed, and this
coastal localities. They include Phorusrhacos longis- resulted in the interchange of faunas to and from
simus Ameghino, Patagornis marshi Moreno & Mer- South America. The phorusrhacids are one example of
cerat, Psilopterus bachmanni (Moreno & Mercerat) south to north movement. An array of modern placen-
and Psilopterus lemoinei (Moreno & Mercerat). The tals, such as modern carnivores, arrived to the south.
seriema-like Cariama santacrucensis Noriega, Viz- It is not easy to understand whether the phorusrhacids
caíno & Bargo was also present. Cariama is the oldest were the losers in the direct competition with these
South American genus with living representatives. carnivores and/or whether their diversity was already
Phorusrhacids show considerable variation in body in decline because their disappearance had already
size, including the small psilopterines, three or four begun. The truth is that the last record of phorusrha-
times larger than the living seriemas of 1.5 kg (body cids in South America dates from the early Pleistocene
mass from Dunning, 2008), and the giant Phorusrha- (Tambussi, Ubilla & Perea, 1999; see also Alvarenga,
cos longissimus at approximately 160 kg (body mass Jones & Rinderknecht, 2010, and later in this article).

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
 PATAGONIAN AND PAMPEAN FOSSIL BIRDS 465

The latest South American species recorded in These forms can only be associated with open habi-
Uruguay was among the largest phorusrhacids ever, tats. This does not exactly agree with the presence of
suggesting that it coexisted with the large carnivorous cebine monkeys (Tejedor et al., 2006) or anhingids
immigrants from North America with which it could (Cenizo & Agnolin, 2010). If anything, these latter
have competed. records suggest that the area was more humid and
Other carnivorous birds from the Santa Cruz For- less open, perhaps agreeing with Croft (2001).
mation include Accipitridae (eagles, hawks) and Fal- Diurnal raptors and psilopterines are compatible with
conidae (caracaras, falcons). Both families have a the presence of environments with alternating grass-
sharply hooked beak, strong legs, feet with raptorial lands and woodlands, as suggested from the floras
claws and diurnal habits. Again, Ameghino described (Barreda & Palazzesi, 2007). Considering only
three species: Badiostes patagonicus Ameghino, the unquestionable records, medium- to large-sized
Thegornis debilis Ameghino and T. musculosus carnivorous species markedly predominate within
Ameghino. Apparently, they all have falconid affini- the Santacrucian avifauna: phorusrhacids, diurnal
ties; furthermore, new findings would have estab- raptors, ibises and anhingids (Fig. 3).
lished that Thegornis is neither a Falconinae nor a
Polyborinae, but has similarities with the living forest
falcons and laughing falcon (Noriega et al., 2008).
BIRDS FROM THE LATE MIOCENE:
Darters (Pelecaniformes Anhingidae) are flying,
TWO HUAYQUERIAN ASSOCIATIONS
foot-propelled diving and fish-eating waterbirds. They
AS EXAMPLES
are mostly tropical and common in freshwater envi-
ronments, but also occur in marshes and marine The Cerro Azul Formation of La Pampa Province
coasts. There are four living species, all placed in the (central Argentina) has yielded numerous and varied
genus Anhinga. Their fossil record is rather abundant bird remains recovered from the Salinas Grandes de
(Noriega, 1992, 1995; Alvarenga & Guilherme, 2003; Hidalgo classical area, and several new localities that
Areta, Noriega & Agnolin, 2007; Cenizo & Agnolin, correspond to the most complete South American avian
2010) and began during the early Miocene. Fossil association of Huayquerian Age (late Miocene)
anhingids in South America include species similar (Fig. 2D) (M. N. Cenizo, C. P. Tambussi & C. Montalvo,
to those living today, as well as some very large unpubl. data). The Formation seems to have been
forms with a marked tendency to becoming flightless. deposited under a semi-arid, probably warm and sea-
The Santacrucian anhingids include Macranhinga sonal climate (Montalvo et al., 2008, and references
Noriega, recovered far from the ocean coast in Santa cited therein). This avian assemblage includes nine
Cruz Province, and Liptornis hesternus Ameghino, taxa representing at least six families, two of which do
which may also be a darter (Cenizo & Agnolin, 2010). not have living representatives (Fig. 4): teratorns, at
Another species recorded at the Santa Cruz For- least two genera of tinamids with modern representa-
mation is Protibis cnemialis Ameghino, a species tives (Eudromia and Nothura), phorushacids, the fal-
belonging to the Threskiornithidae (Brodkorb, 1963; conid Milvago and an undetermined Passeriformes
Pelecaniformes sensu Hackett et al., 2008). Living Tyraniidae (M. N. Cenizo, C. P. Tambussi & C. Mon-
members of the family are large terrestrial birds that talvo, unpubl. data). Other than passerines, all fami-
prey on invertebrates and small vertebrates; they lies had been recorded in previous times.
occur near slow-flowing freshwater or brackish water Undoubtedly, the most impressive is Argentavis
areas. magnificens Campbell & Tonni (Teratornithidae), the
Finally, Anisolornis excavatus Ameghino world’s largest flying bird of all times (80 kg; wing-
(?Cracidae), another species recognized by Ameghino span, 6.5–7.0 m). Teratorns are well known for com-
from the Santa Cruz Formation, is of uncertain affini- plete late Pleistocene skeletons preserved in the
ties (see Brodkorb, 1964, Cracraft, 1973 and Olson, Rancho La Brea tar pits (California) and were related
1985 for different proposals). to storks (Ciconiidae) or New World vultures (Vultu-
This overview of the Santacrucian avifauna allows ridae). Their wing bones are pneumatic, elongated
some inferences. Various lines of evidence suggest the and robust. The ulna exhibits large and widely spaced
presence of open vegetation and dry conditions for the nodes for the attachment of the feathers and, alto-
Santacrucian environments, although others are con- gether, the wing bones resemble those of large soaring
tradictory. At present, seriemas occupy semi-open and birds. Argentavis probably used a thermal soaring
dry landscapes, but Cariama cristata, at least, does mode over the open Pampas, a nonflapping style of
not appear to be a climatic indicator (Tambussi et al., flight energetically inexpensive (Chatterjee, Templin
2005). There is a clear predominance of ground and & Campbell, 2007). Nowadays, it is thought to have
pedestrian birds in the records known so far for this been a diurnal predator (see also Palmqvist & Viz-
age: rheids, giant anseriforms or large phorusrhacids. caíno, 2003) dependent on thermals for flying.

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
466 C. P. TAMBUSSI

Figure 4. Top: climatic scenario during the late Miocene (Huayquerian stage) of La Pampa Province according to the
Cerro Azul Formation record. Birds, some landmark mammals and reptiles are included. Bottom: explanation of the
figure: 1, Cyonasua sp. (Procyonidae); 2, Viperidae; 3, Cricetidae; 4, Nothura sp.; 5, Procariama simplex; 6, Macrauche-
niidae; 7, Tyrannidae; 8, Pterocnemia sp.; 9, Thylacosmilidae; 10, Tetrastylus sp. (Dinomyidae); 11, Glyptodontidae; 12,
Milvago sp.; 13, Argentavis magnificens; 14, Proeuphractus sp. (Dasypodidae); 15, Phorusrhacidae indet.; 16, Pseudoty-
potherium subinsigne (Mesotheriidae); 17, Eudromia sp. Drawings by Marcos Cenizo.

Within Psilopterinae, Procariama simplex Rovereto As pointed out in the Introduction, the climate
has been recorded from the Cerro Azul Formation. deteriorated markedly at the end of the Miocene in
This extinct monotypic genus was flightless, of northern Patagonia and open grassland areas, devel-
medium to large size with a body mass of approxi- oped during dry periods, characterized the environ-
mately 10 kg (Alvarenga & Höfling, 2003). It was ment. Such environmental changes allowed the
originally described from the Miocene–Pliocene of expansion of some faunal elements from northern
northwestern Argentina (Huayquerian, Catamarca localities to the southwest. This is the case for the
Province), representing the phorusrhacid with the Cerro Azul avian remains. These few but interesting
widest geographical range. remains are representatives of a typically Pampean
Remains attributable to Passeriformes Tyrannidae bird fauna at the end of the late Miocene in central-
are also present in the Cerro Azul Formation. south Argentina.

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
 PATAGONIAN AND PAMPEAN FOSSIL BIRDS 467

Another bird assemblage of Huayquerian age comes discussion is based on information provided by the
from the uppermost part of the late Miocene Puerto 1950s collections that had no rigorous provenance
Madryn Formation at Península Valdés. These information. Chapadmalalan (late Pliocene, c. 4.0–
marine deposits crop out along the southwestern 3.0 Ma) was chosen to contrast with the previous
coast of the peninsula, near Punta Delgada (Fig. 2C). scenarios in northern Patagonia. The Chapadmala-
The climate during the accumulation of the late lan age follows the Montehermosan and precedes
Miocene deposits in this region is inferred to have the Uquian age. Records of this age belong to the
been warmer and seasonally drier than that of today. families Rheidae, Tinamidae, Phorusrhacidae (Psi-
From terrestrial palynomorphic evidence, the Penín- lopterinae and Mesembriornithinae), Cathartidae,
sula Valdés Basin was characterized by the presence Charadriidae, Scolopacidae and Furnariidae (Fig. 5,
of low xerophytic open forests and several shrubs. In see Appendix).
addition, the record of sedges and bur-reed would Remains of the steppe tinamous Nothura and
indicate permanently saturated soils with freshwater Eudromia are frequent. Nothura parvula Tambussi is
environments with algae and aquatic ferns (Dozo the most common tinamous from the Chapadmalalan
et al., 2010). Eagles (Accipitridae), whistling ducks (Tambussi & Noriega, 1996). Another taxon recorded
(Dendrocygninae) and phorusrhacid Psilopterinae are there is Darwin’s Nothura (Nothura darwini), an
recorded there. Birds are only a part of the exhumed extant species that currently inhabits arid steppes.
vertebrate fauna that includes fishes (Loricariidae Eudromia elegans inhabits Andean steppes and
and Percomorpha) and abundant and varied mountainsides from Patagonia to Buenos Aires Prov-
mammals (Xenarthra, Rodentia and Litopterna) ince and from central to northwestern Argentina.
(Dozo et al., 2010). Vertebrates come from two close Echarri, Tambussi & Acosta Hospitaleche (2008)
sites of similar age and sedimentology: La Pastosa made a detailed bioclimatic analysis of the distri-
and Rincón Chico. In addition, a large species of bution of the elegant crested tinamou E. elegans,
marabou stork (Leptotilus patagonicus Noriega & allowing a refinement of palaeoenvironmental
Cladera) was recovered at the locality of Punta reconstructions. Low precipitation areas (mean of
Buenos Aires (Noriega & Cladera, 2008). They repre- 311.45 mm) were identified as suitable for Eudromia
sent the first assemblage of late Miocene continental elegans. Hinasuri nehuensis Tambussi, 1995 is the
vertebrates recorded in southern Río Negro Province. sturdiest Rheidae ever recorded.
Within birds, the most exquisite fossil remains is a Two species of Phorusrhacidae have been described:
fragment of the cranial roof for a large-sized adult Mesembriornis australis (Moreno & Mercerat) and M.
eagle (Picasso, Tambussi & Dozo, 2009). Whistling rapax (Patterson & Kraglievich). These large ground
ducks (Dendrocygninae) were also present. Some birds, plus Hinasuri nehuensis, provide evidence of
materials show considerable similarities with the open environments.
living Dendrocygna (Dozo et al., 2010). Dendrocygni- Currently, condors (Vulturidae) do not live in the
nae include herbivorous aquatic birds inhabiting Pampas region, but their fossil record is abundant
lentic environments with dense surface vegetation. (Tonni & Noriega, 1998; Tambussi & Noriega, 1999),
Within this continental assemblage, Psilopterinae are although there is no fossil record in Patagonia. This
also included. As mentioned, psilopterinae are the fact is used by several authors to support the North
smallest and most gracile phorusrhacids; some may American origin of condors with a posterior shift to
even have retained their flying capability. Unlike the South America once the Panamanian bridge was estab-
Cerro Azul Formation and other older sites, no large lished. It is well known that condors are soaring
ground birds, such as rheids or Phorusrhacinae, have scavengers with high orographic affinity. For the
yet been registered in the Puerto Madryn Formation. Montehermosan/Chapadmalalan stages, arid or semi-
The only carnivorous elements of these environments arid conditions compatible with grasslands and forest
were psilopterinae and eagles. Dendrocygninae patches have been inferred. Such conditions favour the
records are consistent with the presence of freshwater presence of updrafts, such as those used by condors for
environments. Large birds, such as Psilopterinae or soaring. Chapadmalalan condors were varied and
Accipitridae, are in agreement with the occurrence of exhibited different sizes. Their diversity appears to be
open forest and shrubs. concomitant with the decrease in phorusrhacids.
The waterbirds Charadriiformes Calidris and
Charadrius indicate the presence of freshwater envi-
BIRDS FROM THE PLIOCENE
ronments. Finally, the passerines are represented by
The main Pliocene localities are situated on the the ovenbirds (Furnariidae). They are small- to
Atlantic coast of Buenos Aires Province (Fig. 2B). medium-sized insectivores; most are forest birds, but
The most recent sampling expeditions have not suc- some occur in more open habitats, such as savannah or
ceeded in collecting new data. Much of the following grassland.

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
468 C. P. TAMBUSSI

Figure 5. Top: climatic scenario during the late Pliocene (Chapadmalalan stage) of Buenos Aires Province. Birds and
some landmark mammals are included. Bottom: explanation of the figure: 1, Actenomys priscus (Ctenomyidae); 2,
Thylophorops chapalmalensis (Didelphidae); 3, Tinamidae; 4, Calidris sp.; 5, Charadrius sp.; 6, Scelidodon sp. (Mylodon-
tidae); 7, Vulturidae indet.; 8, Phorusrhacidae indet.; 9, Chapadmalania altaefrontis (Procyonidae); 10, Mesembriornis
rapax; 11, Thylacosmilus atrox; 12, Ringueletia simpsoni (Dasypodidae); 13, Furnaridae indet. Drawings by Marcos
Cenizo.

The Chapadmalalan bird association (Fig. 5) shares modern bird community began to be formed during
the predominance of carnivores/scavengers with asso- the Oligocene–Miocene transition, South American
ciations of the early Miocene. However, there are no bird fossil records are too fragmentary to allow the
Patagonian fossil condors and the phorusrhacids are reconstruction of such a scenario.
smaller in the Chapadmalalan deposits. The Cenozoic avian fossil record from Patagonia is
still sparse – both geographically and temporally –
and taxonomically poor, relative to the diverse con-
WHAT WORKS AND WHAT DOES NOT
temporary avian fauna of South America. Palaeogene
Although it is generally recognized that the Neorni- fossils do not provide a firm foundation to begin the
thes diversified explosively within 10 Ma following reconstruction of the evolutionary and biogeographi-
the Cretaceous–Palaeogene boundary, and that the cal history of the taxa of higher levels. It is not within

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
 PATAGONIAN AND PAMPEAN FOSSIL BIRDS 469

the scope of the present contribution to cover in detail Teratornithidae have a South American origin and
the biogeographical patterns of the major lineages of did not reach North America until the late Neogene
Neornithes. I direct the reader to Cracraft (2001) and (Campbell & Tonni, 1981). They have a Palaeogene
Mayr (2009) and the various references reviewed by record from Brazil and, after that, during the late
them. Miocene. As phorusrhacids, they move in the Ameri-
Although our knowledge on Patagonian (and South cas in a south–north direction.
American) Neogene birds is characterized by many Two additional derived taxa, Psittaciformes and
uncertainties, some biogeographical inferences are Passeriformes, deserve consideration. Psittaciformes
possible. Rheas belong to a family deeply rooted in (parrots, cockatoos, macaws) represent a very particu-
South America. The fossil record for this family lar group of living birds with a probably Australasian
begins during the Palaeocene and is restricted to origin (Wright et al., 2008), with a long evolutionary
South America (Tambussi, 1995). There is evidence history predating their current panaustral distribu-
that Darwin’s rhea, Pterocnemia pennata, a typically tion. Approximately 43% of Psittaciformes living
Patagonian arid–semi-arid element that nowadays species (148 species within the tribe Arini) are in the
inhabits areas with annual precipitations up to Neotropics. In Argentina, they are recognized from
300 mm (Tambussi & Acosta Hospitaleche, 2002), the late Pliocene and all records are limited to the
extended its distribution to the pampas during the Pampean Region. All but one (Nandayus vorohuensis
Pleistocene (northeast of its current distribution) in Tonni & Noriega) are species of the genus Cyanoliseus
accordance with environmental changes. Similarly, Bonaparte. The burrowing parrot, Cyanoliseus pat-
there is evidence indicating that a second extant agonus, belongs to the most derived clade of parrots.
species, the Greater Rhea, extended further south Tambussi, Acosta Hospitaleche & Horlent (2007)
during the late Pleistocene, and this was facilitated characterize the modern area of distribution as
by the more humid conditions prevailing at that time subhumid–dry to semiarid, with rainfall up to
(Tambussi & Tonni, 1985). 600 mm and, exceptionally, 800 mm, and tempera-
The South American screamers (Anhimidae) consti- tures not below 8–9 °C. Taking into account the
tute a basal lineage of Anseriformes with deep roots current conditions of the Pampean Region, the Cyano-
in South America. Their fossil record may go back into liseus record suggests more arid conditions that at
the late Oligocene–early Miocene of Brazil and, if the present during the late Pleistocene–Holocene at the
assignation of Loxornis clivus to the Anhimidae is southeastern portion of the Pampean Region.
correct, they would have a record in the late Oli- The passerine radiation represents one of the great-
gocene of Patagonia. est avian success stories of all time. They are strong
Gruiformes exhibit a clear Gondwanan distribu- fliers of small size with exceptionally high metabolic
tion, but they also have a wide Laurasian Palaeogene rates. About 58% (5700 species) of the living birds
record and the relationships of these taxa with those (close to 9700 species) are passerines and they
of South America are unclear. Presently, Gruiformes became highly diverse in the Neogene. However, the
are not considered as a natural group (Mayr & fossil record of Passeriformes in Argentina is
Clarke, 2003; Hackett et al., 2008). extremely poor. The oldest record corresponds to an
Phorusrhacids (Cariamiformes) had their early and undetermined Passeriformes (probably belonging to
near complete history in South America. A member of Tyranni) for the early–middle Miocene (Noriega &
this group, Titanis, participates in the second episode Chiappe, 1993) of Pinturas. Fossil Tyranni began to
of the exchange between the Americas that took place be abundant in Argentina starting in the Pliocene
at about 5.0–4.7 Ma (see Woodburne, 2010, and ref- (Tonni & Noriega, 2001). Four species, Ciclodes
erences cited therein). The latest records in South major, Pseudoseisuropsis nehuen, Junco robustus and
America are Pliocene–early Pleistocene in age (Tam- an indeterminate species of Sicalis, are known from
bussi et al., 1999). Recently, Alvarenga et al. (2010) the middle Pleistocene of the Pampean Region. Pos-
described a distal portion of a tarsometatarsus from sibly, the very scarce record of passerines is the result
the late Pleistocene of Uruguay that they assigned to of the sampling mode and does not reflect real
a Phorusrhacidae. At least judging from the published absence.
figure, however, the material is very poor, and addi- The focus in the present article was on four
tional skeletal elements are needed to confirm Neogene associations of birds: one from the late early
identification. Miocene, two from the late Miocene and one from the
Falconidae, which today have a worldwide distri- late Pliocene. This choice is not random. First, the
bution, seem to have a South American origin. Palaeogene record of birds is virtually absent and
Falcons occur in the late early Miocene Santacrucian controversial. Second, the Miocene is a landmark
localities, but the record is extremely poor when com- period of transformation in the Patagonian – and
pared with its high modern diversity. South American – physiognomy. This transformation

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
470 C. P. TAMBUSSI

was the result of several processes: the decrease in nal habitats and biotic structures. Certain groups of
marine temperatures, the decrease in humid condi- birds persisted in their original distribution area,
tions and the increase in xeric conditions, resulting in whereas others succumbed or changed their distribu-
the expansion of the steppe across extra-Andean Pa- tions. Rheids that are strictly terrestrial did not
tagonia and the confinement of the forest to the extend their distribution to North America, even
western areas. The avian fossil record partially though the land connection existed; nor did the Tina-
reflects these processes but, in several cases, is not mids, although they are capable of flight. The Tera-
especially informative. tornithidae, which would have been excellent flyers
However, birds can usefully complement environ- and for which a sea barrier would not have been
mental information when they are members of a an impediment, have no records in North America
representative vertebrate assemblage. It is necessary until the Pliocene. Other senior groups, such as
to incorporate a reflection over the context in which Anhingidae, Pelecanidae, Ciconiidae, Anatidae,
we use the term ‘association’ here. Most of the Anhimidae, Presbyornithidae, Rallidae, Falconidae
Neogene material recognized for Patagonia and the and Accipitridae, established important interconti-
Pampas belongs to old collections. The provenance of nental relationships. Other taxa that are now impor-
some of these materials can be doubtful. However, tant members of the rich South American avian fauna
specimens – and taxa – are considered to belong to (e.g. hummingbirds) are unknown in certain periods
the same association when they come from the same of geological time. Lineages that were strictly terres-
sedimentary formation, although not necessarily from trial, flightless and with giant species persisted for a
the same locality (but nearby; e.g. Santacrucian long time in South America (phorusrhacids, the giant
birds). It is very difficult to extrapolate the environ- Brontornis and rheids). Only the latter survive among
mental requirements of the extinguished taxa from the Neogene mammalian predators to date.
the living fauna (that is, actualism). Although sub- Nowadays, the Patagonian avifauna includes
stantial advances have recently been made in the approximately 156 terrestrial, 52 aquatic and 11
recognition of which environmental variables best oceanic breeding species. Bearing in mind the wide
explain the distribution of some species (tinamids, extension of Patagonia, exceeding one million square
psittaciforms, cariamids, some anseriforms), substan- kilometres, it is clear that the Patagonian avifauna is
tial information is still absent. Environmental recon- presently impoverished.
struction from fossil remains will be reliable when
those remains correspond to extant species. Major
uncertainties arise when the species are not the ACKNOWLEDGEMENTS
same.
Thanks are due to F. Degrange and M. Cenizo for
Eduardo Tonni and Kenneth Campbell, in the
inspiring discussions on Cenozoic South American
1980s, stated that the South American Cenozoic bird
birds over the past few years. Thanks are also due to
fauna was dominated by carnivorous birds, with the
my colleagues C. Mosto and M. Picasso, as well as my
giant Phorusrhacidae and Teratornithidae being the
students, whose frequent questions demanded con-
most noticeable. We can now add the Falconidae,
stant updating. C. Morgan and C. Mosto improved
Accipitridae and Vulturidae (the latter only in the
substantially the English text. F. Degrange assisted
Pampean Region). Other nocturnal raptors (i.e.
with Figure 2. This contribution is based on a presen-
Strigiformes, owls) have not been recorded during the
tation at the Symposium on ‘Palaeogeography and
South American Palaeogene or early Neogene, but
Palaeoclimatology of Patagonia: Effects on Biodiver-
they are very abundant from the Palaeogene of the
sity’, held at the La Plata Museum in Argentina in
Northern Hemisphere. They argued that this domi-
May 2009 and organized by Jorge Rabassa, Eduardo
nance was correlated with a low diversity of large
Tonni, Alfredo Carlini and Daniel Ruzzante. I further
mammals (placental) carnivores. It is very interesting
thank the editors of this special issue for inviting my
to note that carnivore vertebrate communities in Pat-
participation. This study was supported by FONCYT
agonia were represented predominantly by terrestrial
32617 and 0365 grants.
birds and crocodiles (Sebecidae), whereas, in other
continents, this role was occupied by the placental
carnivores. The high prevalence of carnivorous birds
over all other trophic habits continues today, although REFERENCES
represented by different taxa (Tambussi, Noriega & Acosta Hospitaleche C, Castro L, Tambussi CP, Scasso
Tonni, 1993). R. 2008. Palaeospheniscus patagonicus (Aves, Sphenis-
After the connection between both Americas, there cidae): new discoveries from the Early Miocene of Argentina
was intrusion of potential competitors and invasive and its paleoenvironmental significance. Journal of Paleon-
species that undoubtedly had an impact on the origi- tology 82: 565–575.

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
 PATAGONIAN AND PAMPEAN FOSSIL BIRDS 471

Acosta Hospitaleche C, Tambussi CP. 2008. South Ameri- Neues Jahrbuch fur Geologie und Palaontologie Abhandlun-
can fossil penguins: a systematic update. Oryctos 7: 109– gen 243: 343–350.
127. Barreda V, Palazzesi L. 2007. Patagonian vegetation turn-
Acosta Hospitaleche C, Tambussi CP, Donato M, overs during the Paleogene–Early Neogene: origin of arid-
Cozzuol M. 2007. A new Miocene penguin from Patagonia adapted floras. The Botanical Review 73: 31–50.
and a phylogenetic analysis of living and fossil species. Acta Bertelli S, Chiappe L. 2002. Earliest tinamous (Aves:
Paleontologica Polonica 52: 299–314. Palaeognathae) from the Miocene of Argentina and their
Agnolin FL. 2004. Revisión sistemática de algunas aves phylogenetic position. Contributions in Science 502: 1–20.
deseadenses (Oligoceno Medio) descriptas por Ameghino en Bertelli S, Chiappe L, Tambussi CP. 2007. A new phorus-
1899. Revista del Museo Argentino de Ciencias Naturales rhacid (Aves: Cariamae) from the middle Miocene of Pat-
Nueva Serie 6: 239–244. agonia, Argentina. Journal of Vertebrate Paleontology 27:
Agnolin FL. 2006a. Notas sobre el registro de Accipitridae 409–419.
(Aves, Accipitriformes) fósiles argentinos. Studia Geológica Brodkorb P. 1963. Catalogue of fossil birds. Part 1: Archae-
Salmanticensia 42: 67–80. opterygiformes through Ardeiformes. Bulletin of the Florida
Agnolin FL. 2006b. Posición sistemática de algunas aves State Museum (Biological Sciences) 7: 179–293.
fororracoideas (Ralliformes; Cariamae) Argentinas. Revista Brodkorb P. 1964. Catalogue of fossil birds. Part II (Anseri-
del Museo Argentino de Ciencias Naturales Nueva Serie 8: formes through Galliformes). Bulletin of the Florida State
27–33. Museum (Biological Sciences) 8: 195–335.
Agnolin FL. 2007a. Argyrodyptes microtarsus Ameghino, Campbell KE Jr, Tonni EP. 1981. Preliminary observations
1905: un petrel (Procellariiformes) del Eoceno-Oligoceno de on the paleobiology and evolution of teratorns (Aves, Tera-
Argentina. Studia Geológica Salmanticensia 43: 207–213. tornithidae). Journal of Vertebrate Paleontology 1: 265–272.
Agnolin FL. 2007b. Brontornis burmeisteri Moreno & Mer- Cenizo MN, Agnolin FL. 2010. The southernmost records of
cerat, un Anseriformes (Aves) gigante del Mioceno Medio de Anhingidae and a new basal species of Anatidae (Aves) from
Patagonia, Argentina. Revista del Museo Argentino de Cien- the lower–middle Miocene of Patagonia, Argentina. Alcher-
cias Naturales Nueva Serie 9: 15–25. inga: an Australasian Journal of Palaeontology 34: 1–22.
Agnolin FL. 2009. Sistemática y filogenia de las aves Chatterjee S, Templin J, Campbell JK. 2007. The aerody-
fororracoideas (Gruiformes: Cariamae). Buenos Aires: namics of Argentavis, the world’s largest flying bird from
Monografías Fundación Azara/Adrián Giacchino. the Miocene of Argentina. Proceedings of the National
Alvarenga HM, Guilherme E. 2003. The Anhingas (Aves: Academy of Sciences 104: 12 398–12 403.
Anhingidae) from the upper Tertiary (Miocene–Pliocene) of Chiappe L. 1991. Fossil birds from the Miocene Pinturas
southwestern Amazonia. Journal of Vertebrate Paleontology Formation of southern Argentina. Journal of Vertebrate
23: 614–621. Palaeontology 11 (Supplement 3): 21A–22A.
Alvarenga HM, Höfling E. 2003. Systematic revision of the Cracraft J. 1973. Systematics and evolution of the Grui-
Phorusrhacidae (Aves: Ralliformes). Papéis Avulsos de Zoo- formes (Class Aves) 3. Phylogeny of the suborder Gruess.
logia 43: 51–91. American Museum Novitates 151: 1–127.
Alvarenga HM, Jones WW, Rinderknecht A. 2010. The Cracraft J. 2001. Avian evolution, Gondwana biogeography,
youngest record of phorusrhacid birds (Aves, Phorusrhaci- and the Cretaceous–Tertiary mass extinction event.
dae) from the late Pleistocene of Uruguay. Neues Jahrbuch Proceedings of the Royal Society of London 268B: 459–
für Geologie und Palaontologie-Abhandlungen 256: 229– 469.
234. Croft D. 2001. Cenozoic environmental change in South
Ameghino F. 1891. Enumeración de las aves fósiles de la America as indicated by mammalian body size distributions
República Argentina. Buenos Aires: Revista Argentina de (cenograms). Diversity and Distributions 7: 271–287.
Historia Natural, 1: 441–453. Degrange F, Tambussi C, Iglesias A, Zamuner A,
Ameghino F. 1895. Sur les oiseaux fossiles de Patagonie. Wilf P. 2006. Primer registro de Aves para el Daniano.
Boletín del Instituto Geográfico de Argentina 15: 501– Ameghiniana 43 Suplemento: 13R.
602. Degrange FJ, Tambussi CP, Moreno K, Witmer L, Wroe
Ameghino F. 1906. Les formations sédimentaires du Crétacé S. 2010. Mechanical analysis of feeding behavior in the
supérieur et du Tertiaire de Patagonie. Anales del Museo extinct ‘terror bird’ Andalgalornis steulleti (Gruiformes:
Nacional de Buenos Aires 15: v. Phorusrhacidae). PLoS ONE 5: e11856.
Ameghino F. 1908. Notes sur les poissons du Patagonien. Dozo T, Bouza P, Monti A, Palazzesi L, Barreda V,
Anales del Museo Nacional de Historia Natural de Buenos Massaferro G, Scasso R, Tambussi CP. 2010. Late
Aires XVI: (3a serie): 477–497. Miocene continental biota in northeastern Patagonia
Andrews CW. 1899. On the extinct birds of Patagonia. I. The (Península Valdés, Chubut, Argentina). Palaeogeography,
skull and skeleton of Phororhacos inflatus Ameghino. Trans- Palaeoclimatology, Palaeoecology 297: 100–109.
actions of the Zoological Society of London 15: 55–86. Dunning JB. 2008. CRC handbook of avian body masses.
Areta JI, Noriega JI, Agnolin FL. 2007. A giant darter Boca Raton, FL: CRC Press.
(Pelecaniformes: Anhingidae) from the Upper Miocene of Echarri F, Tambussi CP, Acosta Hospitaleche C. 2008.
Argentina and weight calculation of fossil Anhingidae. Predicting the distribution of the crested-tinamous,

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
472 C. P. TAMBUSSI

Eudromia spp. (Aves, Tinamiformes). Journal of Ornithol- tacrucian (Early–Middle Miocene) beds of Patagonia.
ogy 150: 75–84. Ameghiniana 45 (Suppl. 4): 30R.
Ericson PG. 1997. Systematic relationships of the Palaeo- Noriega JI, Chiappe L. 1993. An Early Miocene Passeriform
gene family Presbyornithidae (Aves: Anseriformes). Zoologi- from Argentina. The Auk 110: 936–938.
cal Journal of the Linnean Society 121: 429–483. Noriega JI, Cladera G. 2008. First record of an extinct
Gradstein F, Ogg J, Smith A. 2004. A geologic time scale. marabou stork in the Neogene of South America. Acta
Cambridge: Cambridge University Press. Palaeontologica Polonica 53: 593–600.
Hackett SJ, Kimball RT, Reddy S, Bowie RCK, Braun Noriega JI, Vizcaíno SF, Bargo MS. 2009. First record and
EL, Braun MJ, Chojnowski JL, Cox WA, Han K-L, a new species of Seriema (Aves: Ralliformes: Cariamidae)
Harshman J, Huddleton CJ, Marks BD, Miglia KJ, from Santacrucian (Early–Middle Miocene) beds of Patago-
Moore WA, Sheldon FH, Steadman DW, Witt CC, Yuri nia. Journal of Vertebrate Paleontology 29: 620–626.
T. 2008. A phylogenomic study of birds reveals their evolu- Olson S. 1985. The fossil record of birds. In: Farner DS, King
tionary history. Science 320: 1763–1768. JR, Parkes KC, eds. Avian biology. New York: Academic
Hernandez R, Jordan T, Farjat AD, Echavarria L, Press, 79–238.
Idleman B, Reynolds J. 2005. Age, distribution, tectonics Olson S, Feduccia A. 1980. Presbyornis and the origin of the
and eustatic controls of the Paranense and Caribbean Anseriformes (Aves: Charadriomorphae). Smithsonian Con-
marine transgressions in southern Bolivia and Argentina. tributions to Zoology 323: 1–24.
Journal of South American Earth Sciences 19: 495–512. Palmqvist P, Vizcaíno SF. 2003. Ecological and reproduc-
Jones WW. 2010. Nuevos aportes sobre la paleobiología de los tive constraints of body size in the gigantic Argentavis
fororrácidos (Aves: Ralliformes) basados en el análisis de magnificens (Aves, Theratornithidae) from the Miocene of
estructuras biológicas. PhD Thesis. PEDECIBA – Facultad Argentina. Ameghiniana 40: 379–385.
de Ciencias, Montevideo, Uruguay. Pascual R, Odreman Rivas O. 1971. Evolución de las
Livezey BC. 1997. A phylogenetic analysis of basal Anseri- comunidades de vertebrados del Terciario Argentino. Los
formes, the fossil Presbyornis, and the interordinal relation- aspectos paleozoogeográficos y paleoclimáticos relacionados.
ship of waterfowl. Zoological Journal of the Linnean Society Ameghiniana 8: 372–412.
121: 361–428. Picasso M, Tambussi CP, Dozo T. 2009. Neurocranial
Madden RH, Kay RF, Vucetich MG, Carlini AA. 2010. and brain anatomy of a late Miocene eagle (Aves, Accipi-
Gran Barranca: a 23 million-year record of middle Cenozoic tridae) from Patagonia. Journal of Vertebrate Paleontology
faunal evolution in Patagonia. In: Madden R, Carlini A, 29: 1–6.
Vucetuch G, Kay R, eds. The paleontology of Gran Barranca: Reguero MA, Marenssi S. 2010. Paleogene climatic and
evolution and environmental change through the middle biotic events in the terrestrial record of the Antarctic pen-
Cenozoic of Patagonia. London: Cambridge University insula: an overview. In: Madden RCAA, Vucetich G, Kay R,
Press, 458. eds. The paleontology of Gran Barranca: evolution and envi-
Mayr G. 2009. Paleogene fossil birds. Berlin: Springer-Verlag. ronmental change through the middle Cenozoic of Patago-
Mayr G, Clarke J. 2003. The deep divergences of neornithine nia. London: Cambridge University Press, 383–397.
birds: a phylogenetic analysis of morphological characters. Sinclair WJ, Farr MS. 1932. Aves of the Santa Cruz beds.
Cladistics 19: 527–553. Part II. Reports of the Princeton University Expedition to
Montalvo C, Melchor RN, Visconti G, Cerdeño E. 2008. Patagonia 7: 157–191.
Vertebrate taphonomy in loess–paleosol deposits: a case Tambussi CP. 1995. The fossil Rheiformes from Argentina.
study from the Late Miocene of central Argentina. Geobios Courier Forchungsinstitut Senckenberg 181: 121–129.
41: 133–143. Tambussi CP, Acosta Hospitaleche C. 2002. Reidos (Aves)
Moreno F, Mercerat A. 1891. Catálogo de los pájaros fósiles cuaternarios de Argentina: inferencias paleoambientales.
de la República Argentina conservados en el Museo de La Ameghiniana 39: 95–102.
Plata: La Plata. Tambussi CP, Acosta Hospitaleche C. 2005. Fororracos
Narovsky T, Babarskas M. 2000. Aves de la Patagonia – Psilopterinos (Aves) en la Formación Sarmiento de la local-
guía para su reconocimiento. Buenos Aires: Vázquez idad de Gran Hondonada (Eoceno tardío), Patagonia, Argen-
Manzini Editores. tina. Revista Española de Paleontología 20: 127–132.
Noriega JI. 1992. Un nuevo genero de Anhingidae (Aves: Tambussi CP, Acosta Hospitaleche C, Horlent N. 2007.
Pelecaniformes) de la Formacion Ituzaingo (Mioceno supe- La avifauna del cuaternario de Argentina: inferencias pale-
rior) de Argentina. Notas del Museo de La Plata 109: 217– oambientales a partir del registro de los Psittacidae. In:
223. Pons G, Vicens D, eds. Geomorfologi Litoral i Quaternari.
Noriega JI. 1995. The avifauna from the ‘Mesopotamian’ Balears: Monografía Societat d’Historie Natural de Balears,
(Ituzaingó Formation; Upper Miocene) of Entre Rıos Prov- 69–80.
ince, Argentina. Courier Forchungsinstitut Senckenberg Tambussi CP, Noriega JI. 1996. Summary of the avian
181: 141–148. fossil record from southern South America. In: Arratia G,
Noriega JI, Areta JI, Vizcaíno SF, Bargo MS. 2008. Reas- ed. Contributions of southern South America to vertebrate
sessment of Thegornis musculosus Ameghino 1894 (Aves: paleontology. Munich: Müncher Geowissenschaftliche
Falconidae) based on new material recovered from San- Abhandlungen, 245–264. 247 figs., 241 app.

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
 PATAGONIAN AND PAMPEAN FOSSIL BIRDS 473

Tambussi CP, Noriega JI. 1999. The fossil record of condors Tonni EP, Carlini AA. 2008. Neogene vertebrates from
(Aves, Vulturidae) of Argentina. Smithsonian Contribution Argentine Patagonia: their relationship with the most sig-
in Paleobiology 89: 171–184. nificant climatic changes. Developments in Quaternary Sci-
Tambussi CP, Noriega JI, Tonni EP. 1993. Fossil birds of ences 11: 269–283.
Buenos Aires province (Argentina): an attempt to quantitate Tonni EP, Noriega JI. 1998. Los cóndores (Ciconiiformes,
faunal changes. Palaeogeography, Palaeoclimatology, Vulturidae) de la región pampeana de la Argentina durante
Palaeoecology 3: 117–129. el Cenozoico tardío: distribución interacciones y extinciones.
Tambussi CP, Tonni E. 1985. Aves del sitio arqueológio Los Ameghiniana 35: 141–150.
Toldos, Cañadón de las Cuevas, Santa Cruz. Ameghiniana Tonni EP, Noriega JI. 2001. Una especie extinta de Pseu-
22: 69–74. doseisura Reichenbach 1853 (Passeriformes: Furnariidae)
Tambussi CP, Ubilla M, Acosta Hospitaleche C, Perea D. del Pleistoceno de la Argentina: comentarios filogenéticos.
2005. Paleoenvironmental implications of the fossil avi- Ornitología Neotropical 12: 29–44.
fauna from the Sopas Formation (Late Pleistocene), Tonni E, Tambussi CP. 1988. Un nuevo Psilopterinae (Aves:
Uruguay. Neues Jahrbuch für Geologie und Palaeontologie Ralliformes) del Mioceno tardío de la provincia de Buenos
5: 257–268. Aires, República Argentina. Ameghiniana 25: 155–160.
Tambussi CP, Ubilla M, Perea D. 1999. The youngest large Vizcaíno SF, Bargo MS, Kay RF, Milne N. 2006. The
carnassial bird (Phorusrhacidae, Phorusrhacinae) from armadillos (Mammalia, Xenarthra, Dasypodidae) of the
South America (Pliocene–early Pleistocene of Uruguay). Santa Cruz Formation (early–middle Miocene): an approach
Journal of Vertebrate Paleontology 19: 406–408. to their paleobiology. Palaeogeography, Palaeoclimatology,
Tauber AL. 1994. Estratigrafía y vertebrados fósiles de la Palaeoecology 237: 255–269.
Formación Santa Cruz (Mioceno inferior) en la costa atlán- Vizcaíno SF, Bargo MS, Kay RF, Fariña RA, Giacomo
tica entre las rías del Coyle y Río Gallegos, Provincia de MD, Perry JM, Prevosti FJ, Toledo N, Cassini GH,
Santa Cruz, República Argentina. PhD Thesis, Universidad Fernicola JC. 2010. A baseline paleoecological study for
Nacional de Córdoba. the Santa Cruz Formation (late–early Miocene) at the
Tauber AL. 1997a. Bioestratigrafía de la Formación Santa Atlantic coast of Patagonia, Argentina. Palaeogeography,
Cruz (Mioceno inferior) en el extremo sudeste de la Patago- Palaeoclimatology, Palaeoecology 292: 507–519.
nia. Ameghiniana 34: 413–426. Woodburne M. 2010. The great American biotic interchange:
Tauber AL. 1997b. Paleoecología de la Formación Santa Cruz dispersals, tectonics, climate, sea level and holding pens.
(Mioceno inferior) en el extremo sudeste de la Patagonia. Journal of Mammalian Evolution 17: 245–264.
Ameghiniana 34: 517–529. Wright T, Schirtzinger E, Matsumoto T, Eberhard J,
Tejedor MF, Tauber AA, Rosenberger AL, Swisher CC Graves G, Sanchez J, Capelli S, Müller H, Scharpegge
III, Palacios ME. 2006. New Primate genus from the J, Chambers GK, Fleischer R. 2008. A multilocus
Miocene of Argentina. Proceedings of the National Academy molecular phylogeny of the parrots (Psittaciformes): support
of Sciences of the United States of America 103: 5437– for a Gondwanan origin during the Cretaceous. Molecular
5441. Biology and Evolution 25: 2141–2156.
Tonni EP. 1980. The present state of knowledge of the Zachos J, Shackleton NJ, Revenaugh JS, Pälike H,
Cenozoic birds of Argentina. Contributions in Science, Flower BP. 2001. Climate response to orbital forcing
Natural History Museum, Los Angeles 330: 105–114. across the Oligocene–Miocene. Science 292: 274–278.

APPENDIX
Fossil birds from selected Patagonian and Pampean localities (see text for current status). Ac, Accipitriformes;
An, Anseriformes; Ca, Cariamiformes; Ch, Charadriiformes; Fa, Falconiformes; Pe, Pelecaniformes; Rh,
Rheiformes; Sp, Sphenisciformes; Ti, Tinamiformes.

Taxon Stratigraphy and age

Aves Neornithes Salamanca Formation, early Palaeocene

Rh Rheiformes Río Chico Formation, mid-Palaeocene
An Presbyornis pervetus Casamayor Formation, mid-Eocene
An Telmabates antiqus Casamayor Formation, mid-Eocene
An Presbyornithidae Vaca Mahuida Formation, early Eocene
Ac Accipitridae indet. Chubut province, lower–mid-Eocene
Ca Psilopterinae indet. Sarmiento Formation, late Eocene
Sp Arthrodytes andrewsi San Julián Formation, late Eocene–early Oligocene
Sp Paraptenodytes robustus San Julián Formation, late Eocene–early Oligocene
Sp Argyrodytes microtarsus San Julián Formation, late Eocene–early Oligocene
Ca Physornis fortis Deseado Formation, late Oligocene
Ca Andrewsornis abbotti Deseado Formation, late Oligocene

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474
474 C. P. TAMBUSSI

APPENDIX Continued
Taxon Stratigraphy and age

Ca Psilopterus affinis Sarmiento Formation, late Oligocene

? Pseudolarus guaraniticus Deseado Formation, late Oligocene
Ca Ciconiopsis antarctica Deseado Formation, late Oligocene
An Loxornis clivus Deseado Formation, late Oligocene
? Cruschedula revola Deseado Formation, late Oligocene
? Climacarthus incompletus Deseado Formation, late Oligocene
? Aminornis excavatus Deseado Formation, late Oligocene
An Teleornis impressus Deseado Formation, late Oligocene
? Riacama caliginea Deseado Formation, late Oligocene
Sp Eretiscus tonni Gaiman Formation, early Miocene
Sp Palaeospheniscus bergi Gaiman Formation, early Miocene
Sp Palaeospheniscus patagonicus Gaiman Formation, early Miocene
Sp Palaeospheniscus biloculata Gaiman Formation, early Miocene
Sp Paraptenodytes antarctica Monte León Formation, early Miocene; Puerto Madryn
Formation, early late Miocene
Sp Madrynornis mirandus Puerto Madryn Formation, early late Miocene
Rh Opisthodactylus patagonicus Santa Cruz Formation, late early Miocene
Ti Tinamidae Santa Cruz Formation, late early Miocene
? Anisolornis excavatus Santa Cruz Formation, late early Miocene
An Eoneornis australis Santa Cruz Formation, late early Miocene
An Eutelornis patagonicus Santa Cruz Formation, late early Miocene
An Ankoneta larriestrai Santa Cruz Formation, late early Miocene
An Brontornis burmeisteri Santa Cruz Formation, late early Miocene
Ca Phorusrhacos longissimus Santa Cruz Formation, late early Miocene
Ca Patagornis marshi Santa Cruz Formation, late early Miocene
Ca Psilopterus lemoinei Santa Cruz Formation, late early Miocene
Ca Psilopterus bachmanni Santa Cruz Formation, late early Miocene
Ca Cariama santacrucensis Santa Cruz Formation, late early Miocene
Fa Badiostes patagonicus Santa Cruz Formation, late early Miocene
Fa Thegornis debilis Santa Cruz Formation, late early Miocene
Fa T. musculosus Santa Cruz Formation, late early Miocene
Pe Macranhinga Santa Cruz Formation, late early Miocene
Pe Liptornis hesternus Santa Cruz Formation, late early Miocene
Pe Protibis cnemialis Santa Cruz Formation, late early Miocene
Rh Pterocnemia Cerro Azul Formation, late Miocene
Ti Nothura Cerro Azul Formation, late Miocene
Ti Eudromia sp Cerro Azul Formation, late Miocene
Ca Procariama simplex Cerro Azul Formation, late Miocene
Ci Argentavis magnificens Cerro Azul Formation, late Miocene
Fa Milvago Cerro Azul Formation, late Miocene
Pa Tyraniidae Cerro Azul Formation, late Miocene
Ci Leptoptilus patagonicus Puerto Madryn Formation, early late Miocene
Ac Accipitridae indet. Puerto Madryn Formation, early late Miocene
Ca Psilopterinae indet. Puerto Madryn Formation, early late Miocene
An Dendrocygninae Puerto Madryn Formation, early late Miocene
Rh Hinasuri nehuensis Chapadmalal Formation, late Pliocene
Rh Rheidae indet. Chapadmalal Formation, late Pliocene
Ti Nothura parvula Chapadmalal Formation, late Pliocene
Ti Eudromia Chapadmalal Formation, late Pliocene
Ca Mesembriornis australis Chapadmalal Formation, late Pliocene
Ca M. rapax Chapadmalal Formation, late Pliocene
Ch Calidris Chapadmalal Formation, late Pliocene
Ch Charadrius Chapadmalal Formation, late Pliocene

© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474