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Journal of South American Earth Sciences 37 (2012) 242e255

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Journal of South American Earth Sciences


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An overview of the dinosaur fossil record from Chile


David Rubilar-Rogers a, *, Rodrigo A. Otero a, Roberto E. Yury-Yez b, Alexander O. Vargas c,
Carolina S. Gutstein d, e
a

rea de Paleontologa, Museo Nacional de Historia Natural, Casilla 787, Santiago, Chile
Laboratorio de Zoologa de Vertebrados, Departamento de Ciencias Ecolgicas, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, uoa, Santiago, Chile
c
Laboratorio de Ontogenia y Filogenia, Departamento de Biologa, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, uoa, Santiago, Chile
d
Laboratorio de Ecosiologa, Departamento de Ciencias Ecolgicas, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, uoa, Santiago, Chile
e
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA
b

a r t i c l e i n f o

a b s t r a c t

Article history:
Received 19 January 2012
Accepted 23 March 2012

In Chile, the record of dinosaurs in Jurassic and Cretaceous sediments is often restricted to footprints,
with few skeletal remains. Tetanuran theropods are known in the Upper Jurassic, and bones of titanosaur
sauropods in the Late Cretaceous, including partial skeletons (e.g. Atacamatitan chilensis Kellner et al.).
Also from the late Cretaceous, an ornithopod vertebra, a pair of theropod teeth and one tarsometatarsus
of a gaviiform bird (Neogaeornis wetzeli Lambrecht) have been reported. The Cenozoic fossil record
comprises abundant and well-preserved marine birds from Eocene and Miocene units, with a specially
abundant record of Sphenisciformes and less frequently, Procellariiformes. There is an excellent Miocene
ePliocene record of other birds such as Odontopterygiformes, including the most complete skeleton ever
found of a pelagornithid, Pelagornis chilensis Mayr and Rubilar-Rogers. Fossil birds are also known from
Pliocene and Pleistocene strata. A remarkable collection of birds was discovered in lacustrine sediments
of late Pleistocene age associated to human activity. The perspectives in the study of dinosaurs in Chile
are promising because plenty of material stored in institutional collections is not described yet. The
record of Chilean dinosaurs is relevant for understanding the dynamics and evolution of this group of
terrestrial animals in the western edge of Gondwana, while Cenozoic birds from the Region may
contribute to the understanding of current biogeography for instance, the effect of the emergence and
establishment of the Humboldt Current.
2012 Elsevier Ltd. All rights reserved.

Keywords:
Dinosaurs
Birds
Mesozoic
Cenozoic
Chile

1. Introduction
Continental vertebrates of the Mesozoic terrestrial fauna of
Chile are mainly represented by dinosaurs, with the exception of an
aetosaurid from the Triassic of the Antofagasta Region, Chilenosuchus forttae Casamiquela (Casamiquela, 1980; Desojo, 2003) and
another yet undetermined non-dinosaurian Ornithodira; isolated
bones of pterosaurs from the Cretaceous of the Atacama Region
(Bell and Surez, 1989; Martill et al., 2000, 2006); terrestrial crocodiles from the Aysn Region (Lio et al., 2011); and isolated fragments of turtle plates from the Cretaceous of Coquimbo Region
(Casamiquela et al., 1969). All other continental vertebrates are
represented by dinosaurs.
In the last two decades, knowledge of the record of dinosaurs
has received much attention, not only because of the information
provided about geological problems, for instance, narrowing the
* Corresponding author. Tel.: 56 26804651; fax: 56 28958513.
E-mail address: drubilar@mnhn.cl (D. Rubilar-Rogers).
0895-9811/$ e see front matter 2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jsames.2012.03.003

temporal range of continental deposits (e.g. Blanco et al., 2000), but


also because of their contribution to the knowledge of the evolution
of this clade in this Region of the South American continent, once
part of the occidental margin of Gondwana.
Thus, for example, the rich documentation of dinosaur prints in
layers of the Jurassic and lower Cretaceous (Moreno et al., 2004;
Rubilar-Rogers et al., 2008; among others) allows to appreciate the
wide diversity of kind and size of dinosaurs, and the faunistic
successions undergone between these periods. Further, the remains
of titanosaurs found in different points throughout northern Chile
can indicate how dinosaurs were affected by geographic restrictions
(transgression forming epicontinental seas and expansion of arid
zones) from land emerging during the latter part of the mesozoic,
and if these phenomena generated isolation such that endemism
evolved with regard to other dinosaurs of South America.
During the cenozoic, the dinosaur record is much better represented temporally and in number of discoveries. In the Paleogene
and Neogene, avian remains proceed from the rich marine deposits
of the Atacama Region (e.g. Baha Inglesa Formation) and Patagonia

D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

(Loreto, Ro Turbio, and Ro Baguales formations). Quaternary


records of birds came mainly from lacustrine deposits of the
OHiggins Region (San Vicente de Taguatagua).
The record of cenozoic fossil birds has received great attention,
with several contributions dedicated to documenting these materials (Chvez, 2007a,b; Sallaberry et al., 2008, 2010a). So far, implications of these studies include, for example, the effects of the
establishment of the Humboldt Cold Marine Current System and the
succession of avian faunas throughout the cenozoic in the South
Pacic.
The current work presents an overview of the knowledge
available about dinosaurs in Chile in recent years, updating
previous work that implicitly or explicitly addressed this fauna (e.g.
Chong and Gasparini, 1976; Gasparini, 1979; Salinas et al., 1991a;
Rubilar-Rogers, 2003). Records are ordered by periods and within
each period, the order is by antiquity. A list of the geological units
where dinosaur remains have been found is also delivered below.
Two general maps of Chile show geographical position of records of
the Mesozoic and Cenozoic, respectively.
1.1. Institutional abbreviations
SGO.PV.: Collection of Museo Nacional de Historia Natural,
Santiago. SNGM: Servicio Nacional de Geologa y Minera, Santiago.
2. Geological setting
Several records of Mesozoic dinosaurs from Chile are known
only by their mention in the literature since the materials were not
gured or no hosting institution was properly given in the original
publications (e.g. Biese, 1961; Salinas et al., 1991a, 1991b). As
a consequence, the present review only considered the materials
(Fig. 1) that are available with adequate institutional repository.

243

as a distinctive belt extended from Sierra Moreno to Huatacondo


locality. This latter unit was dated based on ammonoids at several
main local ravines (e.g. Quebrada Arca and Quebrada Huatacondo,
among others, Vicente, 2006), indicating an Oxfordian age for the
marine hosting levels that underlies the typical evaporitic beds,
assigned to the early Kimmeridgian. In consequence, the age of the
continental levels of the Quinchamale Formation is constrained to
a maximum mid Kimmeridgian age. In Sierra Moreno, the roof of
the unit is conformably overlaid by the volcanic succession Cuesta
de Montecristo Volcanites (Charrier et al., 2007), that is partially
equivalent to the Arca Formation (Maksaev, 1978) in the Quebrada
Arca sector. The presence of dinosaur footprints in Quebrada Arca
occurred on multiple layers of sandstones, associated to ripple
marks and mud cracks (Rubilar-Rogers and Otero, 2008). Such
structures and ichnites are consistent with those described in the
upper levels of the Quinchamale Formation, and have not been
observed in the volcanic facies of the overlying Cuesta de Montecristo Volcanites (Ladino et al., 1999). This is the reason why the
outcrops that host the dinosaur footprints are here assigned to the
upper levels of the Quinchamale Formation, and in consequence,
constrained to a Kimmeridgian age.
2.1.3. El Toqui Formation (Surez and De la Cruz, 1994; De la Cruz
and Surez, 2006)
Deposits of a marine transgression over subaerial rocks, conformed mainly by calcareous and volcanoclastic sediments deposited on high-energy shorelines (Charrier et al., 2007). The
volcanoclastic levels are characterized by alternating sandstones
and sedimentary breccias with some levels of tuffs and ignimbrites,
which contains fragmentary bone remains of dinosaurs, as well as
fossil trunks and traces. This formation was assigned to the Upper
Jurassic (Tithonian) based on stratigraphic correlations and radioisotopic dates (Salgado et al., 2008). It is conformably overlaid by
the marine, anoxic sediments of the Katterfeld Formation.

2.1. Mesozoic units


Besides the unconrmed mention (without gures or repository
information) of remains referred to Megalosauridae from early
Callovian marine beds at Cerritos Bayos, northern Chile (Biese,
1961), the oldest record of dinosaurs in the country is restricted
to the upper Jurassic (Salgado et al., 2008). Regrettably, most of the
units of interest lack detailed striatigraphic/biostratigraphic
correlations to constrain the age of the hosting levels of dinosaur
remains. Due to this, most ndings have been referred to broad
chronostratigraphic intervals. The units including veriable records
of dinosaurs in Chile are the following:
2.1.1. Chacarilla Formation (Galli and Dingman, 1962)
Clastic succession conformed by lower marine levels that reach
a thickness of 1100 m. They were assigned by Galli and Dingman
(1962) to the Oxfordian based on fossil invertebrates. The upper
levels are comprised by gray-to-red sandstones that host dinosaur
footprints and can be constrained to a minimal Kimmeridgiane
lower Cretaceous age based on stratigraphic correlations (Charrier
et al., 2007).
2.1.2. Quinchamale Formation (sensu lato, Maksaev, 1978;
Skarmeta and Marinovic, 1981)
Backarc deposits with the lower beds comprising mainly limestones, and shales of Sinemurian to Oxfordian age based on fossil
invertebrates (Maksaev, 1978; Vicente, 2006). The upper levels are
characterized by evaporites (Vicente, 2006), while its uppermost
levels are comprised by quarcites and red, continental shales
(Skarmeta and Marinovic, 1981). The Quinchamale Formation is in
part equivalent to the Aquiuno Formation dened by Garcia (1967)

2.1.4. San Salvador Formation (Lira, 1989)


This unit is comprised by paralic and ne-grained continental
deposits that overlie through a conformable contact to the Cerritos
Bayos Formation (Biese, 1961; Baeza, 1979), and conformably
underlie the Cuesta de Montecristo Volcanites. Due to its stratigraphic relations, it can be directly correlated with the continental
levels of the Quinchamale Formation exposed in the northern and
eastern part of Sierra Moreno, while the beds of the San Salvador
Formation represent equivalent levels exposed in the southern part
of Sierra Moreno (Charrier et al., 2007). In this unit, Moreno et al.
(2004) reported the presence of dinosaur footprints, assigned by
these authors to the Kimmeridgianelower Cretaceous. The beds
with footprints appear to be consistent in facies and stratigraphic
position with the upper levels of the Quinchamale Formation that
host dinosaur trackways. In consequence, we propose a temptative
Kimmeridgian age for the footprints of the San Salvador Formation
(Moreno et al., 2004).
2.1.5. Quebrada Monardes Formation (Muzzio, 1980; Mercado,
1982)
Succession of clastic rocks, mostly reddish in color, exposed in
the Precordillera of Copiap, in the Atacama Region. The basal
member is comprised by mid-to-coarse grained sandstones and
siltstones with cross-bedding (Surez and Bell, 1986). It was interpreted by these authors (Surez and Bell, 1986) as a system of
braided rivers and eolic dunes with proximity to an inner lake, with
several localities hosting dinosaur trackways. It overlies in
conformable contact to the marine limestones of the Lautaro
Formation assigned to the lowereearly middle Jurassic
(Segerstrom, 1968), while its roof is uncertain. Due to this, it was

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D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

Fig. 1. Geographic map showing the distribution of localities in Chile that have yielded remains of Mesozoic Dinosauria (marked with a circle): 1) Quebrada Chacarilla. 2) Huatacondo. 3) El Abra. 4) Quebrada Arca. 5) San Salvador. 6) Pajonales. 7) Quebrada Codocedo. 8) Cerro la Isla. 9) Cerro Algarrobito. 10) Pichasca. 11) Termas del Flaco. 12) Southern part
of Lago General Carrera.

assigned by Surez and Bell (1986) to the Kimmeridgianelower


Cretaceous, while the basal member hosting dinosaur footprints
(Bell and Surez, 1989) could represent Kimmeridgian beds that are
consistent with the age and facies of the backarc units of the second
substage (Charrier et al., 2007) with similar ichnologic records.

volcanic unit Icanche Formation, assigned to the Eocene based on


radioisotopic dating (Charrier et al., 2007). The Tolar Formation is
assigned to the undifferentiated Late Cretaceous based on stratigraphic correlations. This unit has yielded the type material of
Atacamatitan chilensis.

2.1.6. Tolar Formation (Maksaev, 1978)


This unit is located on the west side of the Domeyko Range,
Region de Antofagasta. It is comprised of well-stratied, red
succession of breccias, conglomerates and sandstones. Its lower
levels unconformably contact with the Early Cretaceous levels of
the Arca Formation and probably with the rhyolites of the Pea
Morada Formation, while its roof conformably contacts with the

2.1.7. Pajonales Formation (Harrington, 1961)


It comprises continental red sandstones and conglomerates best
exposed in the homonymous ravine, on Sierra de Almeyda, Antofagasta Region. It overlies the Pular Formation (Brggen, 1942)
through an angular, erosive discordant contact, and is covered by
the same kind of contact with the Guanaqueros Formation (Pino
and Fuenzalida, 1988). The dinosaur remains recovered in the

D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

Pajonales Formation have been assigned to Sauropoda indet


(Salinas et al., 1991b), and were found in lower levels very close to
the discordant contact with the underlying unit. The age of the
fossiliferous beds was assigned to the Maastrichtian based on
stratigraphic correlation (Salinas et al., 1991b).
2.1.8. Hornitos Formation (Segerstrom, 1959)
Breccias and conglomerate levels in coarse sandstone matrix,
including lenses of sandstones, calcareous mudstones and limestones, with discrete volcanic levels to the roof. It is assigned to the
CampanianeMaastrichtian based on stratigraphic correlations
(Seplveda and Naranjo, 1982; Charrier et al., 2007). This unit
contained titanosaur bone remains (Chong, 1985).
2.1.9. Viita Formation (Aguirre and Egert, 1965)
It is comprised mainly by andesites and volcano-sedimentary
rocks with pyroclastic intercalations. It includes fossiliferous beds
with dinosaur bones and continental turtles (Casamiquela et al.,1969),
characterized by a reddish matrix with volcanic, well-selected clasts.
Its radioisotopic dates in levels of underlying and overlying units
allowed constraining the age of the Viita Formation to the
SantonianeMaastrichtian interval (Pineda and Emparn, 2006).
2.1.10. Baos del Flaco Formation (Klohn, 1960)
This unit is characterized by basal marine beds with fossil
invertebrates (ammonoids) that allow assignation to a Tithonian
age. Upper levels are comprised by near-shore, continental beds
that include dinosaur trackways (Casamiquela and Fasola, 1968),
presenting volcanic levels to the roof. These latter authors assigned
an early Cretaceous age to the dinosaur trackways based on their
relative stratigraphic position respect to lower Tithonian beds.
2.1.11. Quiriquina Formation (Bir-Bagczky, 1982)
This unit was formalized by this author, with its type locality on
the homonymous island. It is comprised by a basal conglomerate,
cross-bedded yellow sandstones with conglomerate lenses, coquinaceous horizons and green sandstones at the top that include
concretionary nodules. It was originally assigned to the
CampanianeMaastrichtian and later constrained to the late Maastrichtian based on biostratigraphic correlations (Stinnesbeck, 1996;
Salazar et al., 2010).
2.2. Tertiary units
The PaleogeneeNeogene units are the most recently studied
and produced interesting and abundant avian records (Fig. 2). Like
marine Eocene birds of Loreto Formation and the diverse fauna of
marine birds of Bahia Inglesa Formation.
2.2.1. Estratos de Algarrobo (Gana et al., 1996)
This succession is conformed by sandstones of variable grain
size and hardness, very fossiliferous, with abundant concretionary
nodules in different levels. It reaches about 150 m along the coast
(95 m of thickness) and overlies the strata of Quebrada Municipalidad through an erosive discordance. The roof of the unit is
constrained by a granitic basement through an inferred fault. This
unit was assigned to the middle-to-late Eocene based on fossil
invertebrates with good chronostratigraphic resolution (Brggen,
1915; Tavera, 1980).
2.2.2. Ro Turbio Formation (Feruglio, 1938; sensu lato.; emend.
Hnicken, 1955; sensu Malumin and Carams, 1997)
This unit, originally dened in Argentina, crops out in part of
southernmost Chile. This succession is comprised by sandstones,
conglomerates and intercalated coquinaceous levels. Its age is

245

currently accepted to be middle-to-late Eocene, based on stratigraphic correlations and microfossils (Malumin and Carams,
1997). This unit has yielded fossils of spheniscid and tentative
ardeid birds (Sallaberry et al., 2010b).
2.2.3. Ro Baguales Formation (Cecioni, 1956)
Fossiliferous outcrops out in the northern part of Magallanes. Its
stratigraphic relationships are not completely clear, nevertheless, it
is possible to recognize a thick section of marine sediments that are
consistent with the original denition of the Ro Baguales Formation (Cecioni, 1956), reason why these are referred to this unit. The
age of the beds that host fossil birds is based on the abundant and
typical Eocene associated cartilaginous shes (Sallaberry et al.,
2010b).
2.2.4. Loreto Formation (Hoffstetter et al., 1957)
It is comprised mostly by marine sediments that include
abundant palynomporphs, wood remains, leaf prints, coal beds, as
well as vertebrates, represented by spheniscid penguins and
abundant cartilaginous shes. The age of this unit was recently
constrained to the late Eocene (Priabonian) based on vertebrates
with good chronostratigraphic value, paleobotany and radiometric
date (U/Pb Shrimp) (Otero et al., 2012).
2.2.5. La Portada Formation (Ferraris and Di Biase, 1978)
This unit crops out in Mejillones Peninsula, Antofagasta, and is
comprised by sandstones, occasional diatomites and ne
conglomerates that host a rich vertebrate fauna, including bird
remains. The age of this formation was assigned to the Pliocene
based on microfossils and fossil invertebrates (Tsuchi et al., 1988;
DeVries and Vermeij, 1997), as well as radiometric dating
(Marquardt et al., 2005; Corts et al., 2007).
2.2.6. The Baha Inglesa Formation (Rojo, 1985)
Located on the coast of Atacama Region (Rojo, 1985). It
comprises phosphatic sandstones and conglomerates that overlie
the Oligoceneeearly Miocene Gravas de Angostura unit and is
discordantly covered by the Pleistocene unit of Estratos de Caldera
(Marquardt et al., 2000). Also the unit is in lateral contact with
uvial deposits of the Gravas de Copiap. Most of the fossil vertebrates come from the phosphatic bonebed (sensu Walsh and
Naish, 2002), including an unusual abundance and diversity of
pinnipeds (Walsh and Naish, 2002), cetaceans (Gutstein et al.,
2009) and cartilaginous shes (Long, 1993; Surez et al., 2004).
The age of this formation is constrained to the middle
Mioceneeearly Pliocene based on radioisotopic dates on K/Ar
(Marquardt et al., 2000), and middle Mioceneelate Pliocene on Sr
(Achurra et al., 2009) that are consistent with the chronostratigraphic distribution of several fossil vertebrates hosted in the unit
(Surez and Marquardt, 2003). In the Baha Inglesa Formation at
least 4 localities are distinct, in order of antiquity, Las Arenas,
Mina Fosforita, El Morro, and Los Negros, and Las Arenas,
with bird remains mentioned in this work from the four units
mentioned before.
2.2.7. Coquimbo Formation (Moscoso et al., 1982)
Marine terraces conformed by sandstones and conglomerates,
in part phosphatic, with abundant fossil invertebrates and vertebrates (the latter including birds), exposed on the coast of northcentral Chile. The age of this unit was originally assigned to the
Pliocene and later constrained to the middle Mioceneeearly Pliocene based on fossil invertebrates (Covacevich and Frassinetti,
1990). Additional radioisotopic dates on Sr indicate 14.6 Ma for
beds near the base and 2 Ma for their uppermost levels (Le Roux
et al., 2005).

246

D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

Fig. 2. Geographic map showing the distribution of localities in Chile that have yielded remains of Cenozoic Dinosauria (marked with a circle): 1) La Portada Formation, Antofagasta.
2) Baha Inglesa Formation. 3) Coquimbo Formation. 4). Horcon Formation. 5) Estratos de Algarrobo. 6) Taguatagua. 7) Curamalln Formation. 8) Mocha Island. 9) Ro Baguales
Formation. 10) Ro Turbio Formation. 11) Loreto Formation.

2.2.8. Horcn Formation (Thomas, 1958)


It is comprised almost entirely by sandstones and less frequent
ne conglomerates, exposed in the coast of central Chile. Its age was
assigned to the Pliocene based on fossil invertebrates (Tavera, 1960).
2.2.9. Curamalln Formation (Gonzlez and Vergara, 1962)
Unit that crops out in the cordillieran part of south-central Chile.
Lacustrine beds conformed by ne limolites, claystones and ne
sandstones, assigned to the middleelate Miocene (Niemeyer and
Muoz, 1983). This unit has yielded bird remains in beds with
radioisotopic date (KeAr), indicating 17.5  0.6 and 13.0  1.6 Ma
(Surez and Emparan, 1995).

2.3. Pleistocene units


The following units are not ordered stratigraphically in formally
described formations, so the term locality is used to refer to the
deposits (Fig. 2).
2.3.1. Taguatagua
Systematic excavations were carried out since 1967. An absolute
dating of 11.380  320 years was obtained with Carbon14 from the
level with human artifacts (Montan, 1968).
Fossils of birds were found along with remains of gomphoteres,
horses and human tools of archaeological interest. Although the

D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

larger fauna has been previously recognized (e.g. Casamiquela,1970;


Casamiquela, 1976), besides minor mention in previous work, the
smaller fauna has been studied only recently, comprising micromammals, amphibians (Jimnez-Huidobro et al., 2009) and birds.
2.3.2. Mocha Island
Located in the Lebu province, Biobio Region, the rst paleontological and geological studies of this island were carried out by
Tavera and Veyl (1958) who recognized Miocene levels and assigned
them to the Ranquil Formation (Garca, 1968) from the nearby
pennsula of Arauco, while an underlying unit was assigned to
Navidad Formation. Toward the south of the island Pliocene levels
were also recognized. Later studies of the invertebrate fauna were
carried out by Nielsen and Frassinetti (2007) who describe that, with
minor differences, the fauna of the island resembles Navidad and
Ranquil formations. Finger et al. (2007) consider that Ranquil and
Navidad formations are equivalent in age, sedimentology, fauna and
history of depositation. The Pleistocene levels are the only in which
remains of birds have been reported. These are recognized as
sandstones that are found in a terrace of marine abrasion, these
levels also contain much fractured remains of invertebrate shells.
2.3.3. Lower Pilauco
This paleoindian site is found inward to the los Notros
neighborhood in the city of Osorno. Absolute dating delivers an age
of 14,649  382 years. Casually discovered in 1986, since 2007
systematic excavations have been carried out at this site. Initially,
remains of mastodons were recovered and more recently, other
large and small-sized mammals (horses, chingues, rodents) as well
as insects, coprolites and plant remains (Gonzlez-Guarda et al.,
2008; Montero et al., 2008).
3. Jurassic
The oldest bone remains of dinosaurs in Chile correspond to
materials of two theropods (Theropoda and Tetanurae indet) from
the Toqui Formation (late Jurassic, Tithonian) in the Aysn Region
(south Chile). These materials correspond to an appendicular
skeleton composed of a right Ilium (SNGM-1889), proximal end of
left tibia (SNGM-1885), partial left pes, astragalus and calcaneum,
distal tarsal IV, metatarsal IIeIV and articulated phalanges (SNGM1888) and distal end of right tibia (SNGM-1901) assigned to Theropoda; and dorsal vertebrae (SNGM-1894, 1898, 1900, 1903) and
a partial left manus (SNGM-1887) referred to Tetanurae (Salgado
et al., 2008).
Interesting assemblages of sauropod and theropod icnites are
known from the Tithonian of central Chile (Moreno and Pino, 2002;
Moreno and Benton, 2005; Rubilar-Rogers, 2006) found in a coastal
layer belonging to the Baos del Flaco Formation (late Jurassic,
Tithonian). Other prints have been interpreted as belonging to
ornithopods including an ichnospecies Camptosaurichnus fasolae
(Casamiquela and Fasola,1968), however, given the preservation it is
difcult to evaluate this proposal and these prints have been alternatively referred as Ornithopoda indet (Rubilar-Rogers, 2003). In
nearby layers, the presence of dinosaur bones has been mentioned
(Salinas et al., 1991b), however, the destiny of this material is
unknown and thus this information cannot be conrmed.
An assemblage of theropod prints was reported by Moreno et al.
(2004) at the locality of San Salvador in the Antofagasta Region
consisting of sauropod and theropod prints. The sauropod prints
are narrow-gauged, a condition of sauropods with narrow hips (e.g.
Diplodocimorpha). Three kinds of theropod prints are mentioned.
Such association of narrow-hipped sauropods with theropods is
characteristic of the Jurassic period, which is in concordance with
the age proposed by Moreno et al. (2004) for the red sedimentites

247

from the Estacin member of the San Salvador Formation as


belonging to the Kimmeridgian.
At least four trackways of narrow-hipped sauropods (aff. Brontopodus) and an isolated theropod print were reported from Quebrada Arca (Quinchamale Formation) in the Antofagasta Region
(Rubilar-Rogers and Otero, 2008). These trackways were attributed
to this ichnogenus for its wide-gauge trackway, step angle greater
than 100 , and the absence of claw marks in the manus. The
theropod is distinguished by the separation of the digits with
acuminate distal ends. Several hundreds of prints were also identied from photographs taken by amateurs, in different crops of this
sedimentary sequence, data that still requires systematic recopilation. These sedimentary sequences extend for several kilometers
and it is possible that several dinosaur types remain yet to be found.
Biese (1961) mentions the presence of remains of a Megalosauridae in sandstones from the Tithonian of the Cerritos Bayos
Formation. This nding could not be conrmed since the ultimate
destiny of this material is unknown.
4. Cretaceous
In situ Cretaceous prints are documented from the Codocedo
ravine, the Tambera ravine, Los Pantanos, cerros bravos and cerro
La Isla (Bell and Surez, 1989). These tridactyl prints may correspond to theropods.
Fragmentary dinosaur bones were collected from cerro La Isla. A
caudal vertebra was mentioned by Bell and Surez (1989) (identied
by A.C. Milner) as belonging to an Iguanodontid. This would be the
only reference to skeletal remains of ornithischians in Chile, however,
this material is not available and its destiny is unknown, thus it is
impossible to conrm this specimens taxonomic assignation. Also
from cerro La Isla, a sauropod ulna or bula is mentioned but as in the
previous case the destiny of this important material is unknown.
In the upper Jurassicelower cretaceous range, great assemblages of dinosaur prints are known from the Chacarilla Formation
in northern Chile.
At site III of the Chacarilla ravine and unusual assemblage of
theropod dinosaurs is described (80% of ichnites referred to this
group, the remaining 20% belonging to ornithopods) where at least
two pedal morphotypes can be distinguished (Rubilar-Rogers et al.,
2008).
These generally are large predators which indicate these carnivores were established during the lower Cretaceous along the
occidental margin of Gondwana. In other outcrops of the same
ravine there are narrow and wide-gauged prints of trackways of
sauropods (related to the ichnogenera Parabrontopodus and Brontopodus) as well as some trackways possibly belonging to Thyreophora. Toward the base of the Chacarilla Formation, ornithopod
prints were reported that are consistent with a Cretaceous age for
this formation (Blanco et al., 2000; Rubilar-Rogers et al., 2000). The
prints of the Chacarilla Formation were made in a meandriform
river paleoenvironment.
Tridactyl prints were reported in Huatacondo ravine by Salinas
et al. (1991b), who suggested these could belong to theropods
and ornithopods.
The rst new species of non-avian dinosaur ever discovered in
Chile A. chilensis Kellner et al. (2011) was found in the Tolar
Formation. This titanosaur comprises a right femur, a proximal end
of a humerus, two dorsal vertebrae, two posterior caudal vertebrae,
dorsal ribs and a plate-shaped bone referred as part of a sternal
plate (Kellner et al., 2011, Fig. 3). It is worth noting that the remains
of A. chilensis reveal a gracile titanosaur with femur proportions
different to those of titanosaurs of similar body size.
In order to establish the phylogenetic position of A. chilensis,
a preliminary cladistic analysis using parsimony was performed for

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this review. The data matrix used here corresponds to the one
proposed by Wilson (2002), which was specically designed to test
sauropod phylogeny at the genus level. It was possible to score only
nineteen characters for A. chilensis, out of a total of 324 characters in
Wilsons matrix (presented in Appendix). The analysis of the matrix
was performed using PAUP* software version 4.0 (Swofford, 2003)
with ACCTRAN as character-state optimization and unordered
multistate characters. In this analysis A. chilensis is included as
a Lithostrotia (Fig. 4), a clade that includes the most recent common
ancestor of Saltasaurus and Malawisaurus and all its descendants
(Upchurch et al., 2004). A heuristic search produced 21 most
parsimonious trees (MPTs) with a length of 434 steps (CI 0.6;
RI 0.8). In the strict consensus tree A. chilensis appears inside
Lithostrotia group more closely related to Saltasaurus than to
Malawisaurus, in a node that includes a politomy with Saltasauridae, Nemegtosaurus, Rapetosaurus and Titanosaurus colberti.
Similar results were obtained with bootstrap and jackknife analysis.
In the 50% majority rule bootstrap analysis of the matrix (using
a search with 100 replicates) the tree topology shows a wellsupported node for Saltasauridae (clade formed by the most
recent common ancestor between Saltasaurus and Neuquensaurus),
Rapetosaurus, Nemegtosaurus, T. colberti and A. chilensis. In a posterior phylogenetic analysis Curry-Rogers (2005) proposed a more

inclusive denition for Saltasauridae, which includes genera such


as Malawisaurus, Rapetosaurus, Nemegtosaurus and Titanosaurus
considered non saltasaurids by Wilson (2002). However, the database of Curry-Rogers (2005) is of little help, because in the matrix
there are a lot of uninformative characters, resulting in wide polytomies and low support values for titanosaur ingroups. Additionally a revision of all titanosaur taxa is beyond the purpose of the
present work. A broader data set (not available at the moment)
with more characters may change the hypothesis of relationships of
this specimen specially in reference to Saltasauridae. Up to the
moment another specimen referred to the Lithostrotia clade is
reported for the Cretaceous of the Atacama Region (SNGM-1;
Rubilar-Rogers et al., in prep.), however, the scarce materials of
A. chilensis and the geographic distance of these records (approximately more than 700 km) makes it difcult to comment on the
relationship between these specimens.
An interesting fact is that in comparison with other titanosaurs
of the same size, A. chilensis has slender proportions. Under this
assumption we plotted measurements in different sections of the
femur of saltasaurids such as Titanosaurus falloti, Titanosaurus
robustus, Saltasaurus loricatus, Opisthocoelicaudia skarszinsky,
Rapetosaurus krauseni, Alamosaurus sanjuanensis, Magyarosaurus
dacus, Neuquensaurus australis and the specimens referred to the

Fig. 3. Bones of Atacamatitan chilensis, so far the only new species of dinosaur discovered in Chile. AeH, dorsal vertebrae in AeB, EeF, anterior view and CeD, GeH lateral view; IeL,
rib; MeN, humerus; and OeP, right femur. nc: neural channel, pl: pleurocoel, lb: lateral bulge. Scale bar equals 10 cm. Illustrations: Jocelyn Navarro.

D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

249

Fig. 4. Consensus tree of the 21 most parsimonious trees. IC: 0.66; IR: 0.8; TL: 434, indicating Phylogenetic position of A. chilensis based on a matrix preformed by Wilson (2002) for
sauropod dinosaurs.

last species MCS-5/28 and MCS-5/25 indicated as Neuquensaurus C


and Neuquensaurus D (Fig. 5). In the case of A. chilensis the
measurement of the distal condyles is only estimated, due to
diagenetic distortion. In the graph it is possible to see the gracile
limbs of the Chilean specimen.
The evolution of these differences may relate to geographic
isolation or special environmental conditions during the cretaceous
in northern Chile, restricted to a slender continental border surrounded by epicontinental seas that were only connected to the
mainland by its southern portion, a predominant condition during
the cretaceous in the north and central Chile. The evolution of these
slender-limbed titanosaurs could relate to this restricted area,
which could be compared with the island rule of ecological theory
(e.g. Foster, 1964). This hypothesis requires a comparison with more
taxa and a deeper study of geological conditions during the Upper
Cretaceous in the Atacama Desert, including the dynamics of
deserts in northern Chile. Slender limbs may reect a general
evolutionary trend of a linage including the Chilean specimens. To
discuss this possibility, a complete phylogenetic analysis, including
some taxa suspected of dwarsm (e.g. Magyarosaurus) is necessary
(Rubilar-Rogers, 2008).
Elements of another sauropod specimen referred to Titanosauriformes (Rubilar-Rogers, 2005) have been recovered from the
same Tolar Formation which indicates the potential for further
discovery of more specimens.

Large-sized bones (SGO.PV. 322) were found in the Pajonales


Formation (Upper Cretaceous). P. Taquet (in Salinas et al., 1991b)
identied these remains as titanosaurian sauropods. Since only the
diaphysis of these bones is preserved, it is impossible to identify
them at a less inclusive level than Sauropoda (Rubilar-Rogers,
2003).
Several fossil remains of titanosaurian sauropods have been
reported for the Viita Formation, including the incomplete proximal portion of a right humerus, part of a left scapulo-coracoid, rib
fragments, ischium fragment, the proximal portion of a metapodial
bone, and an incomplete centrum of a caudal vertebra. Based on
these materials Casamiquela et al. (1969) distinguish at least two
kind of titanosaurs. The proximal portion of the humerus was
compared and referred to the genus Antarctosaurus (cf. Antarctosaurus wichmannianus), while the scapulo-coracoid was reconsidered to be an undetermined titanosaur. Rubilar-Rogers (2003)
pointed out that the afnities of the humerus fragment to
A. wichmannianus, cannot be discussed since this element is not
used in the diagnosis of this taxon (Powell, 1986). More recently,
additional material has been found and described for this same site.
This includes the anterior half of a large dorsal vertebra, which
preserves the base of the spine and part of the neural arch (SGO.PV.
959, Vargas et al., 2000). Additionally two teeth assigned to titanosaurs were reported (Salinas and Marshall, 1991; see also
Rubilar-Rogers, 2003). The distal ends are preserved which

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D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

An interesting specimen of titanosaur was found in the Hornitos


Formation comprising several axial and appendicular elements. This
material is currently under study (Rubilar-Rogers et al., in prep.).
Previously Chong (1985) reported a fragment of a left humerus and
rib segments from the same geological unit that was identied and
assigned to the Titanosauridae by Bonaparte (in Chong, 1985).
The only material of Mesozoic birds known from Chile proceeds
from the Quiriquina Formation, an isolated metatarsal referred to
a species of diving bird Neogaeornis wetzeli Lambrecht (1929).
Another similar element was mentioned by Oliver-Schneider
(1940) found in San Vicente bay, however, it was not illustrated
nor adequately described.
Salinas et al. (1991a) reported remains of a large bone from the
strata of Quebrada Blanca that was attributed to a dinosaur,
possibly a sauropod.
Salinas et al. (1991a) mentioned the presence of dinosaur bones
and large bones in cerro Negro and cerro Mesa, both localities
belonging to the Las Chilcas Formation. However, the destiny of
these materials is unknown. Thus, these may even belong to marine
reptiles since they are found in predominantly marine levels.
5. Paleogene

Fig. 5. Charts showing femur width in different specimens of titanosaurs (n 1 for


each species, except Neuquensaurus C and D). A, ratio between the width at midshaft
and total length in natural logarithmic scale. B, width of femur across the shaft: 1)
width of condyles, 2) width at the distal quarter, 3) width at the midshaft, 4) width at
proximal quarter, 5) width at the lateral bulge level, 6) width at femoral head level.
Note that the Chilean specimen (A. chilensis) show low values of width/length ratio
(below) and greater relative length than width values (above).

resemble pencil tips and are 13.0  4.6 mm and 7.75  4.5 mm
respectively, with the larger one possessing, in transeversal section,
an ellipse of 3.0  4.0 mm. In the smaller tooth, it is possible to see
a weared surface toward lingual, while the larger tooth a wear
surface is hardly visible. This suggests a recently erupted tooth or
alternatively a juvenile individual.
The only known cranial remains of non-avian theropod dinosaurs proceed from the Monumento Natural Pichasca (Pichasca
Natural Monument) in the Viita Formation. These consist of two
isolated teeth reported by Salinas and Marshall (1991) and referred
by these authors to coleurosaurs. Rubilar-Rogers (2003) described
these teeth and assigned them as Theropoda indet.
The specimens are small lateral teeth, with no preserved root
(crown heights of 16.6 and 12.0 mm). The teeth are laterally
compressed: in the best, larger, specimen the fore-aft basal length
is the double of the basal width. The basal cross section is not
preserved but the tooth tends to be nearly symmetrical and
teardrop-shaped. It possesses 3 denticles/mm (per millimeter)
along the posterior carina and 4 denticles/mm along the anterior
one. The denticles are generally low. The denticles in anterior and
posterior borders are slightly pointed toward the apex of the crown.
On the posterior carina, denticles are almost as long as they are
wide. Blood grooves are generally shallow and poorly dened. They
are oriented slightly toward the axis of the tooth. This kind of
morphology is very similar to an undescribed non-avian theropod
closely related to dromaeosaurids found in the cretaceous of
Madagascar (Fanti and Therrien, 2007). Nevertheless possible
afnity to abelisaurid cannot be excluded.

The fossil record of Dinosauria in Chile after the K/T event is


restricted to the living order Neornithes of modern birds. The fossil
record of birds in Chile has been revised in several publications
(Chvez, 2007a; Sallaberry et al., 2008, 2010a) but the increasing
work on the group and different localities have made every review
rapidly obsolete.
The fossil record of birds in the Paleogene is represented by levels
strictly of middle to late Eocene age from central Chile (Fm. Algarrobo) and best represented in southernmost Chile (with three
localities) (Sallaberry et al., 2010b; Yury-Yez et al., in press),
contrary to the previous assumption that Paleogene marine
vertebrate-bearing strata were absent in the Chilean record (Clarke
et al., 2003; Chvez, 2007a).
Paleogene fossil birds are almost entirely restricted to the order
Sphenisciformes (penguins), marine diving birds exclusively from
the Southern Hemisphere, which lost their ability to y. Because of
their heavy bones, and their high colonial concentrations, is one of
best represented marine vertebrates during the Cenozoic (de la Cruz,
2007). Also the record of aquatic birds attributed to Ardeidae and
a record of a supposed marine bird is considered in the currently
understudied Chilean Paleogene (Sallaberry et al., 2010b; YuryYez et al., in press).
Paleogene fossil penguins are recognized from three localities in
Magallanes Region, southernmost Chile, described in Sallaberry
et al. (2010b): Ro de Las Minas (Loreto Formation Hoffstetter
et al., 1957), Sierra Dorotea (Ro Turbio Formation Feruglio, 1938;
sensu lato; emend. Hnicken, 1955; sensu Malumin and Carams,
1997), and Sierra Baguales (Ro Baguales Formation, Cecioni, 1956).
In these formations, fossil penguins are represented by two
fragments: an ungual phalanx, and a tarsometatarsus in Ro de las
Minas. Even though the tarsometatarsus is very diagnostic in
penguin taxonomy, the poor preservation of this fossil does not
allow a more accurate designation. However, the high level of fusion
of the metatarsals is considered a common feature of Paleogene
penguins, not shared by the crown group Spheniscidae (Ghlich,
2007) that comprises the extant species and dates back to the
middle Miocene. The ungual phalanx also is considered similar to
one described from the late Eocene of Seymour Island, Antarctica
(g. 19:iej; Jadwiszczak, 2006a). Sierra Baguales represents the
poorest record of fossil penguins: two heavily eroded fragments of
humeri which are recognized by the attened cross section that is
characteristic of penguins.

D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

In contrast, Sierra Dorotea yields the largest amount of fossil


penguins with seven remains, including one that can be assigned at
genus level as Palaeeudyptes. This genus represents one of the most
widespread fossil penguins from the Paleogene being registred in
Australia, New Zealand, and Antarctica (besides Chile). Jadwiszczak
(2011) proposed it to be congeneric with the endemic penguin
fossils from southern Peru, such as, Inkayacu or Icadyptes (Clarke
et al., 2007, 2010). More recently it has been proposed that both
genera of giant penguins, Anthropornis and Palaeeudyptes from the
Eocene of Antarctica should be considered monoespecic
(Jadwiszczak and Mrs, 2011) and not two separate species.
Highly abundant in the marine La Meseta Formation from Seymour Island are the fossils of penguins. Assignations of bones
different to the tarsometatarsus, the most diagnostic element of
Eocene penguins (Myrcha et al., 2002) were based on dubious
criteria, such as, size (Jadwiszczak, 2006b) highly criticized
(Tambussi et al., 2006). The nding of articulated skeletons of
Palaeeudyptes gunnari allows correlating bones for which taxonomical criteria were problematic such as the humerus (AcostaHospitaleche and Reguero, 2010). In the light of the new data it
can be conrmed that Chilean humerus is assignable to Palaeeudyptes, conrming the criteria of Sallaberry et al. (2010b).
Two size classes of fossil penguins are found in the Chilean
Eocene strata. Although only one big sized humerus could be
recognized at the genus level, the remaining bones show size
afnities with other medium and small sized genera from the
Eocene of Antarctica such as Marambiornis, Mesetaornis or Delphinornis (Sallaberry et al., 2010b).
Non spheniscid remains are only known from a record assigned
to Ardeidae also from the Sierra Dorotea locality, a proximal fragment of tibiotarsus lacking an extension from the crista cnemialis
cranialis in proximal direction, as in other aquatic birds assigned to
this family, suggesting also it was a non-exclusively marine environment (Sallaberry et al., 2010b). From strata of Algarrobo, central
Chile, an incomplete fragmentary femur is the only record of an
Eocene bird from this locality. This does not represent a fossil
penguin, and thus poses question about the environment and
taphonomical inuences during depositation of the sediments in
this locality. It also comprises a locality that is intermediate in
latitude between southernmost Chile, Antarctica and fossil penguin
localities in Southern Peru (Yury-Yez et al., in press). The fragmentary nature of the fossil does not allow more specic taxonomic
afnities, but in general anatomy it could be related to an aquatic
bird, most probably a procelariid.
6. Neogene
The Neogene fossil record of modern birds is the most studied and
abundant of the order Dinosauria. Regardless of the enormous
localities of Cenozoic vertebrates from continental sediments
studied during the last decades (Croft et al., 2008), almost no bird has
been recorded from continental levels, with the exception of Meganhinga chilensis. In this sense, as with the Paleogene record, most
birds are also from marine localities in the Neogene. We will introduce rst the marine taxa (given the amount of data available) and
conclude with the fewer continental records. Three formations
northern to 30 S are the most studied and yield the highest amounts
of fossil birds. These are, from North to South: La Portada Formation
(late MioceneePliocene, Antofagasta Region), Baha Inglesa Formation (late MioceneePliocene, Atacama Region) and the Coquimbo
Formation (middle Miocene-Pliocene, Coquimbo Region). In the
central part of the country, isolated remains have been recorded in
the Horcn Formation (Pliocene, Valparaso Region). The only
exclusively continental formation with bird remains is the lacustrine
Curamalln Formation (Miocene) in the Biobio Region.

251

Fossil penguins are again the most remarkable group (in


number and diversity), including several endemic taxa. All Neogene
records belong to the crown group Spheniscidae and particularly to
extant genera, with the exception of one record of a humerus cf.
Palaeospheniscus from the Coquimbo Formation (AcostaHospitaleche et al., 2006a). This extinct genus is considered in
phylogenetical analysis just outside the crown group and is recorded originally in the early Miocene of Argentinean Patagonia
(Acosta-Hospitaleche et al., 2007). One enigmatic tarsometatarsus
from the Baha Inglesa Formation is attributable to Spheniscidae cf.
Palaeospheniscus in the size range of P. biloculata, but preservation
does not allow assessing the presence of this genus with certainty
in the late Miocene of Chile (Soto-Acua et al., 2008). Records from
Baha Inglesa Formation assigned to Argentinean genera (such as
Paraptenodytes or Palaeospheniscus) were reassigned to extant taxa,
particularly cranial remains. The taxonomical problem of Miocene
Chilean penguins have been discussed elsewhere, and here we will
only be considering recent assignments accepted in previous
reviews, in which all cranial and appendicular remains (with the
exceptions listed above) are restricted to extant genera of the
crown group (Chvez, 2007a,b; Acosta-Hospitaleche and Canto,
2005; Acosta-Hospitaleche and Canto, 2007; Acosta-Hospitaleche
et al., 2005; Soto-Acua et al., 2008).
Three neogene endemic species have being described: Spheniscus chilensis Emslie and Guerra-Correa (2003), Pygoscelis grandis
Walsh and Surez (2006), and Pygoscelis calderensis AcostaHospitaleche et al. (2006b). The rst one from the La Portada
Formation is a specimen of the penguin genus Spheniscus that is
present in South American Atlantic and Pacic coasts comprising
the species S. humboldti and Spheniscus magellanicus and the Galpagos penguin S. mendiculus. S. chilensis is a medium small sized
penguin Pliocene in age.
P calderensis and P. grandis are both recovered from different
levels of the Baha Inglesa Formation. P. calderensis was recovered
from late Miocene levels of phosphatic deposits and comprises
a species in the size range of the extant members of the genera.
P. grandis is a large sized extinct species that was recovered from
Pliocene levels of the Bahia Inglesa Formation near the size of the
extant king penguin (Aptenodytes forsteri).
From the late Miocene levels of Baha Inglesa Formation there
are records belonging to the extant genus Spheniscus and characterized by their big size. Spheniscus urbinai and Spheniscus megaramphus are represented by several neurocrania, an isolated
rostrum and appendicular bones (Soto-Acua et al., 2008; Chvez,
2007c, 2006). Only S. urbinai is known from associated skeletons
from the levels of El Morro locality from the Baha Inglesa
Formation (Yury-Yez et al., 2009) and is also represented (by
cranial remains) in the Chaaral de Aceituno locality from the
Coquimbo Formation (Chvez, 2005).
Besides Spheniscidae, seven other families of Neornithes are
known from the Chilean Neogene fossil record: Diomedeidae, Procellariidae, Phalacrocoracidae, Sulidae, Anhingidae, Falconidae and
Pelagornithidae. The order Procellariiformes is the second best
represented in Baha Inglesa Formation (after penguins) and
comprises two families. Diomedeidae (mollymawks or albatrosses)
is recognized by cranial remains of the genus Thalassarche (according to Nunn et al., 1996) or Diomedea (Walsh and Hume, 2001).
Procellariidae (petrels and others) is represented by several genera
currently under study, recognizing at least the tribe Pufninni
(particularly the genus Calonectris, Yury-Yez et al., 2008), gady
petrels comparable to Pterodroma, petrels related to Pagodroma,
a procelariid of uncertain afnities (probably Pterodroma, YuryYez et al., 2010a) and the genus Pachiptyla (Sallaberry et al., 2007).
The diversity of sulids (also exclusively from Baha Inglesa
Formation) comprises big sized forms with afnities to known

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D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

Fig. 6. Pelagornis chilensis, the most complete pelagornithid worldwide. A, cranium in lateral view. B, mounted skeleton. C, cranium in palatal view. Fur, furrow; nar, narial opening;
nvf, neurovascular foramen; qud, quadrate; fos, fossae for reception of the mandibular pseudo-teeth. Image by S. Trnkner.

species in Pisco Formation (Peru), and two other size classes, all
recognized as belonging to the genus Sula. Three forms of Sulidae
are known (Walsh and Hume, 2001; Chvez and Stucchi, 2006;
Soto-Acua et al., 2009) Sula cf. variegata, Sula magna and Sula cf.
sulita. S. magna should be considered the best represented based in
cranial remains. Other family, Phalacrocoracidae, is based on
appendicular bones that were described by Walsh and Hume
(2001).
With the Neogene marine avian fauna, the Pelagornithidae is
the most outstanding bird family, typically known from Paleogene
and Neogene levels from every continent. Until now, in Chile, it is
only recognized from the late Miocene of Baha Inglesa Formation
based on several isolated remains and the complete articulated
skeleton of Pelagonis chilensis, which Mayr and Rubilar-Rogers
(2010) considered as the bird with the largest well established
wingspan, within both living and extinct birds (Fig. 6). This specimen is also important due to its exquisite preservation, allowing
cranialepostcranial correlations which were impossible until this
nding. Based on Mayr and Rubilar-Rogers (2010), it is highly
probable that all Baha Inglesa Formation material (several isolated
bones) belong to the genus Pelagornis, coinciding with earlier
identications (Chvez et al., 2007).
Undescribed fossil birds from the Pliocene of the Horcn
Formation have been mentioned (Carrillo-Briceo et al., 2011) as
Charadriiformes, Sphenisciformes and Phalacrocoracidae, but at
least one fossil of Spheniscus from this unit is recognized at the
genus level (Yury-Yez et al., 2010b).
Two records of Neogene birds are ascribed as the only nonmarine birds, the Falconidae Milvago sp. from La Portada Formation (Emslie and Guerra Correa, 2003) and M. chilensis described as
ightless and big sized snake bird (Anhingidae; Alvarenga, 1995).
7. Quaternary
A large number of unpublished materials (hundreds of isolated
specimens) have been recovered from the Pleistocene of San
Vicente de Taguatagua, mainly from plaster jackets of vertebrate
remains recovered by Lautaro Nez and Rodolfo Casamiquela in
the 1980s. These comprise well-preserved and diagnostic remains
that will allow determining temporal variation in the taxa
composition of the zone from the Pleistocene to present.
Bird remains have also been found at the Pleistocene locality of
Pilauco Bajo, in the city of Osorno which have been informally
mentioned and are currently under study (Montero et al., 2008). The
collection of the department of Geology of the University of Chile
houses undetermined fragments from the Pleistocene of Mocha
Island of penguins of similar size to the modern genus Spheniscus that
were collected by Tavera and Veyl in 1958, but remain unstudied.

8. Conclusions and perspectives


No dinosaur remains have been recorded from the Triassic
period in Chile. However, this trend may be more related to the lack
of systematic eld work rather than the absence of such remains.
The Triassic outcrops of the strata of El Bordo, in the Antofagasta
Region, are auspicious since they consist of a sequence of lacustrine
deposits where two forms of non-dinosaurian archosaurs have
been found. Projected prospection of such outcrops should be
a priority because of the great importance that these levels represent for understanding the evolutionary history of dinosaurs.
For now, Jurassic dinosaurs are scarcely represented. The
discovery of theropod skeletons in the Aysen Region and the
deposits of dinosaur footprints from the Antofagasta Region are
promising. Doubtless, more material of theropod dinosaurs can be
recovered from the strata of El Toqui and possibly new prints will
be documented in quebrada Arca.
Cretaceous deposits have delivered a very abundant record of
dinosaur prints that allow us to know about the fauna of the time
despite the absence of skeletal remains. Some formations of this
period overlie Jurassic deposits, such that it becomes possible to
understand changes in faunal composition across these periods.
Additionally, the titanosaurs represent the best-documented nonavian dinosaurs in Chile. Doubtless, new forms of these dinosaurs
will continue to be discovered in sites like La Higuera ravine in the
Atacama Region, where new and more complete specimens are
likely to be collected.
As mentioned above, teeth possibly belonging to a dromaeosaurid are the only cranial remains of non-avian dinosaurs
currently known from Chile. If this identication is correct, this
evidence would be the rst record of this kind of dinosaur from
Chile and one of the few currently known for Gondwana.
The only record of birds from the cretaceous in Chile is two
isolated tarsometatarsi from the Quiriquina Formation. This
material is important since it is one of the few occurrences of
Mesozoic birds known for Gondwana. New eld work in this
formation is necessary in order to nd new evidence of these birds.
Until now, there is no information from Chile about other
terrestrial taxa that must have co-existed with dinosaurs during the
Mesozoic, such as mammals and ophidians. This fact is dened
mainly by the lack of systematic search for other components of the
Mesozoic biota. Reports of lizards and amphibians must be
corroborated since current mentions are unclear and the destiny of
these materials is unknown.
Continental Mesozoic vertebrates are very important to narrow
down possible ages of continental levels (Blanco et al., 2000;
Rubilar-Rogers, 2003). Thus they will continue to be a good tool for
establishing biochrones of Mesozoic continental levels in Chile.

D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255

Among perspectives for research on dinosaurs and other


Mesozoic continental vertebrates, an interesting problem is the
degree of endemism that may have evolved in relation to the
separation produced by epicontinental seas.
Acknowledgments
We are very grateful of the editors of JSES specially of Timothy
Horscroft for his invitation to publish this paper. Many thanks to
Jocelyn Navarro who made the line drawings of Atacamasaurus
chilensis. Conicyt-Chile funded PhD degree of D. R.-R. and C.S.G. and
master degree of R. Y.-Y. We thank Jurassic Foundation for eld
support.
Appendix
List of characters codied for A. chilensis based in the matrix
from Wilson (2002).
10
A. chilensis

20

30

40

50

?????????? ?????????? ?????????? ?????????? ??????????


60

70

80

90

100

?????????? ?????????? ??????11?? ?????????? ??????????


110

120

????11???? ?????????? ??????????


160

170

130

140

10?1?????? ????????1?

180

190

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