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
243
244
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.
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
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.
247
248
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
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.
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.
250
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.
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252
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.
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