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-Yáñez b, Alexander O. Vargas c, Carolina S. Gutstein d,e a Área de Paleontología, Museo Nacional de Historia Natural, Casilla 787, Santiago, Chile b Laboratorio de Zoología de Vertebrados, Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Ñuñoa, Santiago, Chile c Laboratorio de Ontogenia y Filogenia, Departamento de Biología, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Ñuñoa, Santiago, Chile d Laboratorio de Ecofisiología, Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Ñuñoa, Santiago, Chile e Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA article info abstract

Article history: In Chile, the record of dinosaurs in Jurassic and Cretaceous sediments is often restricted to footprints, Received 19 January 2012 with few skeletal remains. Tetanuran theropods are known in the Upper Jurassic, and bones of titanosaur Accepted 23 March 2012 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 Keywords: of a gaviiform bird (Neogaeornis wetzeli Lambrecht) have been reported. The Cenozoic fossil record Dinosaurs comprises abundant and well-preserved marine birds from Eocene and Miocene units, with a specially Birds abundant record of Sphenisciformes and less frequently, Procellariiformes. There is an excellent Miocene Mesozoic e Cenozoic Pliocene record of other birds such as Odontopterygiformes, including the most complete skeleton ever Chile 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. Ó 2012 Elsevier Ltd. All rights reserved.

1. Introduction temporal range of continental deposits (e.g. Blanco et al., 2000), but also because of their contribution to the knowledge of the evolution Continental vertebrates of the Mesozoic terrestrial fauna of of this clade in this Region of the South American continent, once Chile are mainly represented by dinosaurs, with the exception of an part of the occidental margin of Gondwana. aetosaurid from the Triassic of the Antofagasta Region, Chileno- Thus, for example, the rich documentation of dinosaur prints in suchus forttae Casamiquela (Casamiquela, 1980; Desojo, 2003) and layers of the Jurassic and lower Cretaceous (Moreno et al., 2004; another yet undetermined non-dinosaurian Ornithodira; isolated Rubilar-Rogers et al., 2008; among others) allows to appreciate the bones of pterosaurs from the Cretaceous of the Atacama Region wide diversity of kind and size of dinosaurs, and the faunistic (Bell and Suárez, 1989; Martill et al., 2000, 2006); terrestrial croc- successions undergone between these periods. Further, the remains odiles from the Aysén Region (Lio et al., 2011); and isolated frag- of titanosaurs found in different points throughout northern Chile ments of turtle plates from the Cretaceous of Coquimbo Region can indicate how dinosaurs were affected by geographic restrictions (Casamiquela et al., 1969). All other continental vertebrates are (transgression forming epicontinental seas and expansion of arid represented by dinosaurs. zones) from land emerging during the latter part of the mesozoic, In the last two decades, knowledge of the record of dinosaurs and if these phenomena generated isolation such that endemism has received much attention, not only because of the information evolved with regard to other dinosaurs of South America. provided about geological problems, for instance, narrowing the During the cenozoic, the dinosaur record is much better repre- sented temporally and in number of discoveries. In the Paleogene * Corresponding author. Tel.: þ56 26804651; fax: þ56 28958513. and Neogene, avian remains proceed from the rich marine deposits E-mail address: [email protected] (D. Rubilar-Rogers). of the Atacama Region (e.g. Bahía Inglesa Formation) and Patagonia

0895-9811/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved. doi:10.1016/j.jsames.2012.03.003 D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255 243

(Loreto, Río Turbio, and Río Baguales formations). Quaternary as a distinctive belt extended from Sierra Moreno to Huatacondo records of birds came mainly from lacustrine deposits of the locality. This latter unit was dated based on ammonoids at several O’Higgins Region (San Vicente de Taguatagua). main local ravines (e.g. Quebrada Arca and Quebrada Huatacondo, The record of cenozoic fossil birds has received great attention, among others, Vicente, 2006), indicating an Oxfordian age for the with several contributions dedicated to documenting these mate- marine hosting levels that underlies the typical evaporitic beds, rials (Chávez, 2007a,b; Sallaberry et al., 2008, 2010a). So far, impli- assigned to the early Kimmeridgian. In consequence, the age of the cations of these studies include, for example, the effects of the continental levels of the Quinchamale Formation is constrained to establishment of the Humboldt Cold Marine Current System and the a maximum mid Kimmeridgian age. In Sierra Moreno, the roof of succession of avian faunas throughout the cenozoic in the South the unit is conformably overlaid by the volcanic succession Cuesta Pacific. de Montecristo Volcanites (Charrier et al., 2007), that is partially The current work presents an overview of the knowledge equivalent to the Arca Formation (Maksaev, 1978) in the Quebrada available about dinosaurs in Chile in recent years, updating Arca sector. The presence of dinosaur footprints in Quebrada Arca previous work that implicitly or explicitly addressed this fauna (e.g. occurred on multiple layers of sandstones, associated to ripple Chong and Gasparini, 1976; Gasparini, 1979; Salinas et al., 1991a; marks and mud cracks (Rubilar-Rogers and Otero, 2008). Such Rubilar-Rogers, 2003). Records are ordered by periods and within structures and ichnites are consistent with those described in the each period, the order is by antiquity. A list of the geological units upper levels of the Quinchamale Formation, and have not been where dinosaur remains have been found is also delivered below. observed in the volcanic facies of the overlying Cuesta de Mon- Two general maps of Chile show geographical position of records of tecristo Volcanites (Ladino et al., 1999). This is the reason why the the Mesozoic and Cenozoic, respectively. outcrops that host the dinosaur footprints are here assigned to the upper levels of the Quinchamale Formation, and in consequence, 1.1. Institutional abbreviations constrained to a Kimmeridgian age.

SGO.PV.: Collection of Museo Nacional de Historia Natural, 2.1.3. El Toqui Formation (Suárez and De la Cruz, 1994; De la Cruz Santiago. SNGM: Servicio Nacional de Geología y Minería, Santiago. and Suárez, 2006) Deposits of a marine transgression over subaerial rocks, con- 2. Geological setting formed mainly by calcareous and volcanoclastic sediments depos- ited on high-energy shorelines (Charrier et al., 2007). The Several records of Mesozoic dinosaurs from Chile are known volcanoclastic levels are characterized by alternating sandstones only by their mention in the literature since the materials were not and sedimentary breccias with some levels of tuffs and ignimbrites, figured or no hosting institution was properly given in the original which contains fragmentary bone remains of dinosaurs, as well as publications (e.g. Biese, 1961; Salinas et al., 1991a, 1991b). As fossil trunks and traces. This formation was assigned to the Upper a consequence, the present review only considered the materials Jurassic (Tithonian) based on stratigraphic correlations and radio- (Fig. 1) that are available with adequate institutional repository. isotopic dates (Salgado et al., 2008). It is conformably overlaid by the marine, anoxic sediments of the Katterfeld Formation. 2.1. Mesozoic units 2.1.4. San Salvador Formation (Lira, 1989) Besides the unconfirmed mention (without figures or repository This unit is comprised by paralic and fine-grained continental information) of remains referred to Megalosauridae from early deposits that overlie through a conformable contact to the Cerritos Callovian marine beds at Cerritos Bayos, northern Chile (Biese, Bayos Formation (Biese, 1961; Baeza, 1979), and conformably 1961), the oldest record of dinosaurs in the country is restricted underlie the Cuesta de Montecristo Volcanites. Due to its strati- to the upper Jurassic (Salgado et al., 2008). Regrettably, most of the graphic relations, it can be directly correlated with the continental units of interest lack detailed striatigraphic/biostratigraphic levels of the Quinchamale Formation exposed in the northern and correlations to constrain the age of the hosting levels of dinosaur eastern part of Sierra Moreno, while the beds of the San Salvador remains. Due to this, most findings have been referred to broad Formation represent equivalent levels exposed in the southern part chronostratigraphic intervals. The units including verifiable records of Sierra Moreno (Charrier et al., 2007). In this unit, Moreno et al. of dinosaurs in Chile are the following: (2004) reported the presence of dinosaur footprints, assigned by these authors to the Kimmeridgianelower Cretaceous. The beds 2.1.1. Chacarilla Formation (Galli and Dingman, 1962) with footprints appear to be consistent in facies and stratigraphic Clastic succession conformed by lower marine levels that reach position with the upper levels of the Quinchamale Formation that a thickness of 1100 m. They were assigned by Galli and Dingman host dinosaur trackways. In consequence, we propose a temptative (1962) to the Oxfordian based on fossil invertebrates. The upper Kimmeridgian age for the footprints of the San Salvador Formation levels are comprised by gray-to-red sandstones that host dinosaur ( Moreno et al., 2004). footprints and can be constrained to a minimal Kimmeridgiane lower Cretaceous age based on stratigraphic correlations (Charrier 2.1.5. Quebrada Monardes Formation (Muzzio, 1980; Mercado, et al., 2007). 1982) Succession of clastic rocks, mostly reddish in color, exposed in 2.1.2. ‘Quinchamale Formation’ (sensu lato, Maksaev, 1978; the Precordillera of Copiapó, in the Atacama Region. The basal Skarmeta and Marinovic, 1981) member is comprised by mid-to-coarse grained sandstones and Backarc deposits with the lower beds comprising mainly lime- siltstones with cross-bedding (Suárez and Bell, 1986). It was inter- stones, and shales of Sinemurian to Oxfordian age based on fossil preted by these authors (Suárez and Bell, 1986) as a system of invertebrates (Maksaev, 1978; Vicente, 2006). The upper levels are braided rivers and eolic dunes with proximity to an inner lake, with characterized by evaporites (Vicente, 2006), while its uppermost several localities hosting dinosaur trackways. It overlies in levels are comprised by quarcites and red, continental shales conformable contact to the marine limestones of the Lautaro (Skarmeta and Marinovic, 1981). The Quinchamale Formation is in Formation assigned to the lowereearly middle Jurassic part equivalent to the Aquiuno Formation defined by Garcia (1967) (Segerstrom, 1968), while its roof is uncertain. Due to this, it was 244 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) Hua- tacondo. 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 Suárez and Bell (1986) to the Kimmeridgianelower volcanic unit Icanche Formation, assigned to the Eocene based on Cretaceous, while the basal member hosting dinosaur footprints radioisotopic dating (Charrier et al., 2007). The Tolar Formation is (Bell and Suárez, 1989) could represent Kimmeridgian beds that are assigned to the undifferentiated Late Cretaceous based on strati- consistent with the age and facies of the backarc units of the second graphic correlations. This unit has yielded the type material of substage (Charrier et al., 2007) with similar ichnologic records. Atacamatitan chilensis.

2.1.6. Tolar Formation (Maksaev, 1978) 2.1.7. Pajonales Formation (Harrington, 1961) This unit is located on the west side of the Domeyko Range, It comprises continental red sandstones and conglomerates best Region de Antofagasta. It is comprised of well-stratified, red exposed in the homonymous ravine, on Sierra de Almeyda, Anto- succession of breccias, conglomerates and sandstones. Its lower fagasta Region. It overlies the Pular Formation (Brüggen, 1942) levels unconformably contact with the Early Cretaceous levels of through an angular, erosive discordant contact, and is covered by the Arca Formation and probably with the rhyolites of the Peña the same kind of contact with the Guanaqueros Formation (Pino Morada Formation, while its roof conformably contacts with the and Fuenzalida, 1988). The dinosaur remains recovered in the D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255 245

Pajonales Formation have been assigned to Sauropoda indet currently accepted to be middle-to-late Eocene, based on strati- (Salinas et al., 1991b), and were found in lower levels very close to graphic correlations and microfossils (Malumián and Caramés, the discordant contact with the underlying unit. The age of the 1997). This unit has yielded fossils of spheniscid and tentative fossiliferous beds was assigned to the Maastrichtian based on ardeid birds (Sallaberry et al., 2010b). stratigraphic correlation (Salinas et al., 1991b). 2.2.3. Río Baguales Formation (Cecioni, 1956) 2.1.8. Hornitos Formation (Segerstrom, 1959) Fossiliferous outcrops out in the northern part of Magallanes. Its Breccias and conglomerate levels in coarse sandstone matrix, stratigraphic relationships are not completely clear, nevertheless, it including lenses of sandstones, calcareous mudstones and lime- is possible to recognize a thick section of marine sediments that are stones, with discrete volcanic levels to the roof. It is assigned to the consistent with the original definition of the Río Baguales Forma- CampanianeMaastrichtian based on stratigraphic correlations tion (Cecioni, 1956), reason why these are referred to this unit. The (Sepúlveda and Naranjo, 1982; Charrier et al., 2007). This unit age of the beds that host fossil birds is based on the abundant and contained titanosaur bone remains (Chong, 1985). typical Eocene associated cartilaginous fishes (Sallaberry et al., 2010b). 2.1.9. Viñita Formation (Aguirre and Egert, 1965) It is comprised mainly by andesites and volcano-sedimentary 2.2.4. Loreto Formation (Hoffstetter et al., 1957) rocks with pyroclastic intercalations. It includes fossiliferous beds It is comprised mostly by marine sediments that include with dinosaur bones and continental turtles (Casamiquela et al.,1969), abundant palynomporphs, wood remains, leaf prints, coal beds, as characterized by a reddish matrix with volcanic, well-selected clasts. well as vertebrates, represented by spheniscid penguins and Itsradioisotopicdatesinlevelsofunderlyingandoverlyingunits abundant cartilaginous fishes. The age of this unit was recently allowed constraining the age of the Viñita Formation to the constrained to the late Eocene (Priabonian) based on vertebrates SantonianeMaastrichtian interval (Pineda and Emparán, 2006). with good chronostratigraphic value, paleobotany and radiometric date (U/Pb Shrimp) (Otero et al., 2012). 2.1.10. Baños del Flaco Formation (Klohn, 1960) This unit is characterized by basal marine beds with fossil 2.2.5. La Portada Formation (Ferraris and Di Biase, 1978) invertebrates (ammonoids) that allow assignation to a Tithonian This unit crops out in Mejillones Peninsula, Antofagasta, and is age. Upper levels are comprised by near-shore, continental beds comprised by sandstones, occasional diatomites and fine that include dinosaur trackways (Casamiquela and Fasola, 1968), conglomerates that host a rich vertebrate fauna, including bird presenting volcanic levels to the roof. These latter authors assigned remains. The age of this formation was assigned to the Pliocene an early Cretaceous age to the dinosaur trackways based on their based on microfossils and fossil invertebrates (Tsuchi et al., 1988; relative stratigraphic position respect to lower Tithonian beds. DeVries and Vermeij, 1997), as well as radiometric dating (Marquardt et al., 2005; Cortés et al., 2007). 2.1.11. Quiriquina Formation (Biró-Bagóczky, 1982) This unit was formalized by this author, with its type locality on 2.2.6. The Bahía Inglesa Formation (Rojo, 1985) the homonymous island. It is comprised by a basal conglomerate, Located on the coast of Atacama Region (Rojo, 1985). It cross-bedded yellow sandstones with conglomerate lenses, coqui- comprises phosphatic sandstones and conglomerates that overlie naceous horizons and green sandstones at the top that include the Oligoceneeearly Miocene Gravas de Angostura unit and is concretionary nodules. It was originally assigned to the discordantly covered by the Pleistocene unit of Estratos de Caldera CampanianeMaastrichtian and later constrained to the late Maas- (Marquardt et al., 2000). Also the unit is in lateral contact with trichtian based on biostratigraphic correlations (Stinnesbeck, 1996; fluvial deposits of the Gravas de Copiapó. Most of the fossil verte- Salazar et al., 2010). brates come from the phosphatic “bonebed” (sensu Walsh and Naish, 2002), including an unusual abundance and diversity of 2.2. Tertiary units pinnipeds (Walsh and Naish, 2002), cetaceans (Gutstein et al., 2009) and cartilaginous fishes (Long, 1993; Suárez et al., 2004). The PaleogeneeNeogene units are the most recently studied The age of this formation is constrained to the middle and produced interesting and abundant avian records (Fig. 2). Like Miocene eearly Pliocene based on radioisotopic dates on K/Ar marine Eocene birds of Loreto Formation and the diverse fauna of (Marquardt et al., 2000), and middle Mioceneelate Pliocene on Sr marine birds of Bahia Inglesa Formation. (Achurra et al., 2009) that are consistent with the chronostrati- graphic distribution of several fossil vertebrates hosted in the unit 2.2.1. Estratos de Algarrobo (Gana et al., 1996) (Suárez and Marquardt, 2003). In the Bahía Inglesa Formation at This succession is conformed by sandstones of variable grain least 4 localities are distinct, in order of antiquity, “Las Arenas”, size and hardness, very fossiliferous, with abundant concretionary “Mina Fosforita”, “El Morro”, and “Los Negros”, and “Las Arenas”, nodules in different levels. It reaches about 150 m along the coast with bird remains mentioned in this work from the four units (95 m of thickness) and overlies the strata of Quebrada Munici- mentioned before. palidad through an erosive discordance. The roof of the unit is constrained by a granitic basement through an inferred fault. This 2.2.7. Coquimbo Formation (Moscoso et al., 1982) unit was assigned to the middle-to-late Eocene based on fossil Marine terraces conformed by sandstones and conglomerates, invertebrates with good chronostratigraphic resolution (Brüggen, in part phosphatic, with abundant fossil invertebrates and verte- 1915; Tavera, 1980). brates (the latter including birds), exposed on the coast of north- central Chile. The age of this unit was originally assigned to the 2.2.2. Río Turbio Formation (Feruglio, 1938; sensu lato.; emend. Pliocene and later constrained to the middle Mioceneeearly Plio- Hünicken, 1955; sensu Malumián and Caramés, 1997) cene based on fossil invertebrates (Covacevich and Frassinetti, This unit, originally defi ned in Argentina, crops out in part of 1990). Additional radioisotopic dates on Sr indicate 14.6 Ma for southernmost Chile. This succession is comprised by sandstones, beds near the base and 2 Ma for their uppermost levels (Le Roux conglomerates and intercalated coquinaceous levels. Its age is 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) Bahía Inglesa Formation. 3) Coquimbo Formation. 4). Horcon Formation. 5) Estratos de Algarrobo. 6) Taguatagua. 7) Curamallín Formation. 8) Mocha Island. 9) Río Baguales Formation. 10) Río Turbio Formation. 11) Loreto Formation.

2.2.8. Horcón Formation (Thomas, 1958) 2.3. Pleistocene units It is comprised almost entirely by sandstones and less frequent fine conglomerates, exposed in the coast of central Chile. Its age was The following units are not ordered stratigraphically in formally assigned to the Pliocene based on fossil invertebrates (Tavera,1960). described formations, so the term “locality” is used to refer to the deposits (Fig. 2). 2.2.9. Curamallín Formation (González and Vergara, 1962) Unit that crops out in the cordillieran part of south-central Chile. 2.3.1. Taguatagua Lacustrine beds conformed by fine limolites, claystones and fine Systematic excavations were carried out since 1967. An absolute sandstones, assigned to the middleelate Miocene (Niemeyer and dating of 11.380 320 years was obtained with Carbon14 from the Muñoz, 1983). This unit has yielded bird remains in beds with level with human artifacts (Montané, 1968). radioisotopic date (KeAr), indicating 17.5 0.6 and 13.0 1.6 Ma Fossils of birds were found along with remains of gomphoteres, (Suárez and Emparan, 1995). horses and human tools of archaeological interest. Although the D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255 247 larger fauna has been previously recognized (e.g. Casamiquela,1970; from the “Estación” member of the San Salvador Formation as Casamiquela, 1976), besides minor mention in previous work, the belonging to the Kimmeridgian. smaller fauna has been studied only recently, comprising micro- At least four trackways of narrow-hipped sauropods (aff. Bron- mammals, amphibians (Jiménez-Huidobro et al., 2009) and birds. topodus) and an isolated theropod print were reported from Que- brada Arca (Quinchamale Formation) in the Antofagasta Region 2.3.2. Mocha Island (Rubilar-Rogers and Otero, 2008). These trackways were attributed Located in the Lebu province, Biobio Region, the first paleonto- to this ichnogenus for its wide-gauge trackway, step angle greater logical and geological studies of this island were carried out by than 100, and the absence of claw marks in the manus. The Tavera and Veyl (1958) who recognized Miocene levels and assigned theropod is distinguished by the separation of the digits with them to the Ranquil Formation (García, 1968) from the nearby acuminate distal ends. Several hundreds of prints were also identi- península of Arauco, while an underlying unit was assigned to fied from photographs taken by amateurs, in different crops of this Navidad Formation. Toward the south of the island Pliocene levels sedimentary sequence, data that still requires systematic recopila- were also recognized. Later studies of the invertebrate fauna were tion. These sedimentary sequences extend for several kilometers carried out by Nielsen and Frassinetti (2007) who describe that, with and it is possible that several dinosaur types remain yet to be found. minor differences, the fauna of the island resembles Navidad and Biese (1961) mentions the presence of remains of a Mega- Ranquil formations. Finger et al. (2007) consider that Ranquil and losauridae in sandstones from the Tithonian of the Cerritos Bayos Navidad formations are equivalent in age, sedimentology, fauna and Formation. This finding could not be confirmed since the ultimate history of depositation. The Pleistocene levels are the only in which destiny of this material is unknown. remains of birds have been reported. These are recognized as sandstones that are found in a terrace of marine abrasion, these 4. Cretaceous levels also contain much fractured remains of invertebrate shells. In situ Cretaceous prints are documented from the Codocedo 2.3.3. Lower Pilauco ravine, the Tambería ravine, Los Pantanos, cerros bravos and cerro This paleoindian site is found inward to the “los Notros” La Isla (Bell and Suárez, 1989). These tridactyl prints may corre- neighborhood in the city of Osorno. Absolute dating delivers an age spond to theropods. of 14,649 382 years. Casually discovered in 1986, since 2007 Fragmentary dinosaur bones were collected from cerro La Isla. A systematic excavations have been carried out at this site. Initially, caudal vertebra was mentioned by Bell and Suárez (1989) (identified remains of mastodons were recovered and more recently, other by A.C. Milner) as belonging to an “Iguanodontid”. This would be the large and small-sized mammals (horses, chingues, rodents) as well only reference to skeletal remains of ornithischians in Chile, however, as insects, coprolites and plant remains (González-Guarda et al., this material is not available and its destiny is unknown, thus it is 2008; Montero et al., 2008). impossible to confirm this specimen’s taxonomic assignation. Also from cerro La Isla, a sauropod ulna or fibula is mentioned but as in the 3. Jurassic previous case the destiny of this important material is unknown. In the upper Jurassicelower cretaceous range, great assem- The oldest bone remains of dinosaurs in Chile correspond to blages of dinosaur prints are known from the Chacarilla Formation materials of two theropods (Theropoda and Tetanurae indet) from in northern Chile. the Toqui Formation (late Jurassic, Tithonian) in the Aysén Region At “site III” of the Chacarilla ravine and unusual assemblage of (south Chile). These materials correspond to an appendicular theropod dinosaurs is described (80% of ichnites referred to this skeleton composed of a right Ilium (SNGM-1889), proximal end of group, the remaining 20% belonging to ornithopods) where at least left tibia (SNGM-1885), partial left pes, astragalus and calcaneum, two pedal morphotypes can be distinguished (Rubilar-Rogers et al., distal tarsal IV, metatarsal IIeIV and articulated phalanges (SNGM- 2008). 1888) and distal end of right tibia (SNGM-1901) assigned to Ther- These generally are large predators which indicate these carni- opoda; and dorsal vertebrae (SNGM-1894, 1898, 1900, 1903) and vores were established during the lower Cretaceous along the a partial left manus (SNGM-1887) referred to Tetanurae (Salgado occidental margin of Gondwana. In other outcrops of the same et al., 2008). ravine there are narrow and wide-gauged prints of trackways of Interesting assemblages of sauropod and theropod icnites are sauropods (related to the ichnogenera Parabrontopodus and Bron- known from the Tithonian of central Chile (Moreno and Pino, 2002; topodus) as well as some trackways possibly belonging to Thyreo- Moreno and Benton, 2005; Rubilar-Rogers, 2006) found in a coastal phora. Toward the base of the Chacarilla Formation, ornithopod layer belonging to the Baños del Flaco Formation (late Jurassic, prints were reported that are consistent with a Cretaceous age for Tithonian). Other prints have been interpreted as belonging to this formation (Blanco et al., 2000; Rubilar-Rogers et al., 2000). The ornithopods including an ichnospecies Camptosaurichnus fasolae prints of the Chacarilla Formation were made in a meandriform (Casamiquela and Fasola,1968), however, given the preservation it is river paleoenvironment. difficult to evaluate this proposal and these prints have been alter- Tridactyl prints were reported in Huatacondo ravine by Salinas natively referred as Ornithopoda indet (Rubilar-Rogers, 2003). In et al. (1991b), who suggested these could belong to theropods nearby layers, the presence of dinosaur bones has been mentioned and ornithopods. (Salinas et al., 1991b), however, the destiny of this material is The first new species of non-avian dinosaur ever discovered in unknown and thus this information cannot be confirmed. Chile A. chilensis Kellner et al. (2011) was found in the Tolar An assemblage of theropod prints was reported by Moreno et al. Formation. This titanosaur comprises a right femur, a proximal end (2004) at the locality of San Salvador in the Antofagasta Region of a humerus, two dorsal vertebrae, two posterior caudal vertebrae, consisting of sauropod and theropod prints. The sauropod prints dorsal ribs and a plate-shaped bone referred as part of a sternal are narrow-gauged, a condition of sauropods with narrow hips (e.g. plate (Kellner et al., 2011, Fig. 3). It is worth noting that the remains Diplodocimorpha). Three kinds of theropod prints are mentioned. of A. chilensis reveal a gracile titanosaur with femur proportions Such association of narrow-hipped sauropods with theropods is different to those of titanosaurs of similar body size. characteristic of the Jurassic period, which is in concordance with In order to establish the phylogenetic position of A. chilensis, the age proposed by Moreno et al. (2004) for the red sedimentites a preliminary cladistic analysis using parsimony was performed for 248 D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255 this review. The data matrix used here corresponds to the one inclusive definition for Saltasauridae, which includes genera such proposed by Wilson (2002), which was specifically designed to test as Malawisaurus, Rapetosaurus, Nemegtosaurus and Titanosaurus sauropod phylogeny at the genus level. It was possible to score only considered non saltasaurids by Wilson (2002). However, the data- nineteen characters for A. chilensis, out of a total of 324 characters in base of Curry-Rogers (2005) is of little help, because in the matrix Wilson’s matrix (presented in Appendix). The analysis of the matrix there are a lot of uninformative characters, resulting in wide pol- was performed using PAUP* software version 4.0 (Swofford, 2003) ytomies and low support values for titanosaur ingroups. Addi- with ACCTRAN as character-state optimization and unordered tionally a revision of all titanosaur taxa is beyond the purpose of the multistate characters. In this analysis A. chilensis is included as present work. A broader data set (not available at the moment) a Lithostrotia (Fig. 4), a clade that includes the most recent common with more characters may change the hypothesis of relationships of ancestor of Saltasaurus and Malawisaurus and all its descendants this specimen specially in reference to Saltasauridae. Up to the (Upchurch et al., 2004). A heuristic search produced 21 most moment another specimen referred to the Lithostrotia clade is parsimonious trees (MPTs) with a length of 434 steps (CI ¼ 0.6; reported for the Cretaceous of the Atacama Region (SNGM-1; RI ¼ 0.8). In the strict consensus tree A. chilensis appears inside Rubilar-Rogers et al., in prep.), however, the scarce materials of Lithostrotia group more closely related to Saltasaurus than to A. chilensis and the geographic distance of these records (approxi- Malawisaurus, in a node that includes a politomy with Saltasaur- mately more than 700 km) makes it difficult to comment on the idae, Nemegtosaurus, Rapetosaurus and Titanosaurus colberti. relationship between these specimens. Similar results were obtained with bootstrap and jackknife analysis. An interesting fact is that in comparison with other titanosaurs In the 50% majority rule bootstrap analysis of the matrix (using of the same size, A. chilensis has slender proportions. Under this a search with 100 replicates) the tree topology shows a well- assumption we plotted measurements in different sections of the supported node for Saltasauridae (clade formed by the most femur of saltasaurids such as Titanosaurus falloti, Titanosaurus recent common ancestor between Saltasaurus and Neuquensaurus), robustus, Saltasaurus loricatus, Opisthocoelicaudia skarszinsky, Rapetosaurus, Nemegtosaurus, T. colberti and A. chilensis. In a poste- Rapetosaurus krauseni, Alamosaurus sanjuanensis, Magyarosaurus rior phylogenetic analysis Curry-Rogers (2005) proposed a more 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 Large-sized bones (SGO.PV. 322) were found in the Pajonales and Neuquensaurus D (Fig. 5). In the case of A. chilensis the Formation (Upper Cretaceous). P. Taquet (in Salinas et al., 1991b) measurement of the distal condyles is only estimated, due to identified these remains as titanosaurian sauropods. Since only the diagenetic distortion. In the graph it is possible to see the gracile diaphysis of these bones is preserved, it is impossible to identify limbs of the Chilean specimen. them at a less inclusive level than Sauropoda (Rubilar-Rogers, The evolution of these differences may relate to geographic 2003). isolation or special environmental conditions during the cretaceous Several fossil remains of titanosaurian sauropods have been in northern Chile, restricted to a slender continental border sur- reported for the Viñita Formation, including the incomplete prox- rounded by epicontinental seas that were only connected to the imal portion of a right humerus, part of a left scapulo-coracoid, rib mainland by its southern portion, a predominant condition during fragments, ischium fragment, the proximal portion of a metapodial the cretaceous in the north and central Chile. The evolution of these bone, and an incomplete centrum of a caudal vertebra. Based on slender-limbed titanosaurs could relate to this restricted area, these materials Casamiquela et al. (1969) distinguish at least two which could be compared with the island rule of ecological theory kind of titanosaurs. The proximal portion of the humerus was (e.g. Foster, 1964). This hypothesis requires a comparison with more compared and referred to the genus Antarctosaurus (cf. Antarcto- taxa and a deeper study of geological conditions during the Upper saurus wichmannianus), while the scapulo-coracoid was reconsid- Cretaceous in the Atacama Desert, including the dynamics of ered to be an undetermined titanosaur. Rubilar-Rogers (2003) deserts in northern Chile. Slender limbs may reflect a general pointed out that the affinities of the humerus fragment to evolutionary trend of a linage including the Chilean specimens. To A. wichmannianus, cannot be discussed since this element is not discuss this possibility, a complete phylogenetic analysis, including used in the diagnosis of this taxon (Powell, 1986). More recently, some taxa suspected of dwarfism (e.g. Magyarosaurus) is necessary additional material has been found and described for this same site. (Rubilar-Rogers, 2008). This includes the anterior half of a large dorsal vertebra, which Elements of another sauropod specimen referred to Titano- preserves the base of the spine and part of the neural arch (SGO.PV. sauriformes (Rubilar-Rogers, 2005) have been recovered from the 959, Vargas et al., 2000). Additionally two teeth assigned to tita- same Tolar Formation which indicates the potential for further nosaurs were reported (Salinas and Marshall, 1991; see also discovery of more specimens. Rubilar-Rogers, 2003). The distal ends are preserved which 250 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 identified 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

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 (Chávez, 2007a; Sallaberry et al., 2008, 2010a) but the increasing work on the group and different localities have made every review rapidly obsolete. Fig. 5. Charts showing femur width in different specimens of titanosaurs (n ¼ 1 for The fossil record of birds in the Paleogene is represented by levels each species, except Neuquensaurus C and D). A, ratio between the width at midshaft strictly of middle to late Eocene age from central Chile (Fm. Algar- and total length in natural logarithmic scale. B, width of femur across the shaft: 1) robo) and best represented in southernmost Chile (with three 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. localities) (Sallaberry et al., 2010b; Yury-Yáñez et al., in press), Note that the Chilean specimen (A. chilensis) show low values of width/length ratio contrary to the previous assumption that Paleogene marine (below) and greater relative length than width values (above). vertebrate-bearing strata were absent in the Chilean record (Clarke et al., 2003; Chávez, 2007a). Paleogene fossil birds are almost entirely restricted to the order resemble pencil tips and are 13.0 4.6 mm and 7.75 4.5 mm Sphenisciformes (penguins), marine diving birds exclusively from respectively, with the larger one possessing, in transeversal section, the Southern Hemisphere, which lost their ability to fly. Because of an ellipse of 3.0 4.0 mm. In the smaller tooth, it is possible to see their heavy bones, and their high colonial concentrations, is one of a weared surface toward lingual, while the larger tooth a wear best represented marine vertebrates during the Cenozoic (de la Cruz, surface is hardly visible. This suggests a recently erupted tooth or 2007). Also the record of aquatic birds attributed to Ardeidae and alternatively a juvenile individual. a record of a supposed marine bird is considered in the currently The only known cranial remains of non-avian theropod dino- understudied Chilean Paleogene (Sallaberry et al., 2010b; Yury- saurs proceed from the “Monumento Natural Pichasca (Pichasca Yáñez et al., in press). Natural Monument) in the Viñita Formation. These consist of two Paleogene fossil penguins are recognized from three localities in isolated teeth reported by Salinas and Marshall (1991) and referred Magallanes Region, southernmost Chile, described in Sallaberry by these authors to coleurosaurs. Rubilar-Rogers (2003) described et al. (2010b): Río de Las Minas (Loreto Formation Hoffstetter these teeth and assigned them as Theropoda indet. et al., 1957), Sierra Dorotea (Río Turbio Formation Feruglio, 1938; The specimens are small lateral teeth, with no preserved root sensu lato; emend. Hünicken, 1955; sensu Malumián and Caramés, (crown heights of 16.6 and 12.0 mm). The teeth are laterally 1997), and Sierra Baguales (Río Baguales Formation, Cecioni, 1956). compressed: in the best, larger, specimen the fore-aft basal length In these formations, fossil penguins are represented by two is the double of the basal width. The basal cross section is not fragments: an ungual phalanx, and a tarsometatarsus in Río de las preserved but the tooth tends to be nearly symmetrical and Minas. Even though the tarsometatarsus is very diagnostic in teardrop-shaped. It possesses 3 denticles/mm (per millimeter) penguin taxonomy, the poor preservation of this fossil does not along the posterior carina and 4 denticles/mm along the anterior allow a more accurate designation. However, the high level of fusion one. The denticles are generally low. The denticles in anterior and of the metatarsals is considered a common feature of Paleogene posterior borders are slightly pointed toward the apex of the crown. penguins, not shared by the crown group Spheniscidae (Göhlich, On the posterior carina, denticles are almost as long as they are 2007) that comprises the extant species and dates back to the wide. Blood grooves are generally shallow and poorly defined. They middle Miocene. The ungual phalanx also is considered similar to are oriented slightly toward the axis of the tooth. This kind of one described from the late Eocene of Seymour Island, Antarctica morphology is very similar to an undescribed non-avian theropod (fig. 19:iej; Jadwiszczak, 2006a). Sierra Baguales represents the closely related to dromaeosaurids found in the cretaceous of poorest record of fossil penguins: two heavily eroded fragments of Madagascar (Fanti and Therrien, 2007). Nevertheless possible humeri which are recognized by the flattened cross section that is affinity to abelisaurid cannot be excluded. characteristic of penguins. D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255 251

In contrast, Sierra Dorotea yields the largest amount of fossil Fossil penguins are again the most remarkable group (in penguins with seven remains, including one that can be assigned at number and diversity), including several endemic taxa. All Neogene genus level as Palaeeudyptes. This genus represents one of the most records belong to the crown group Spheniscidae and particularly to widespread fossil penguins from the Paleogene being registred in extant genera, with the exception of one record of a humerus cf. Australia, New Zealand, and Antarctica (besides Chile). Jadwiszczak Palaeospheniscus from the Coquimbo Formation (Acosta- (2011) proposed it to be congeneric with the endemic penguin Hospitaleche et al., 2006a). This extinct genus is considered in fossils from southern Peru, such as, Inkayacu or Icadyptes (Clarke phylogenetical analysis just outside the crown group and is recor- et al., 2007, 2010). More recently it has been proposed that both ded originally in the early Miocene of Argentinean Patagonia genera of giant penguins, Anthropornis and Palaeeudyptes from the (Acosta-Hospitaleche et al., 2007). One enigmatic tarsometatarsus Eocene of Antarctica should be considered monoespecific from the Bahía Inglesa Formation is attributable to Spheniscidae cf. (Jadwiszczak and Mörs, 2011) and not two separate species. Palaeospheniscus in the size range of P. biloculata, but preservation Highly abundant in the marine La Meseta Formation from Sey- does not allow assessing the presence of this genus with certainty mour Island are the fossils of penguins. Assignations of bones in the late Miocene of Chile (Soto-Acuña et al., 2008). Records from different to the tarsometatarsus, the most diagnostic element of Bahía Inglesa Formation assigned to Argentinean genera (such as Eocene penguins (Myrcha et al., 2002) were based on dubious Paraptenodytes or Palaeospheniscus) were reassigned to extant taxa, criteria, such as, size (Jadwiszczak, 2006b) highly criticized particularly cranial remains. The taxonomical problem of Miocene (Tambussi et al., 2006). The finding of articulated skeletons of Chilean penguins have been discussed elsewhere, and here we will Palaeeudyptes gunnari allows correlating bones for which tax- only be considering recent assignments accepted in previous onomical criteria were problematic such as the humerus (Acosta- reviews, in which all cranial and appendicular remains (with the Hospitaleche and Reguero, 2010). In the light of the new data it exceptions listed above) are restricted to extant genera of the can be confirmed that Chilean humerus is assignable to Palae- crown group (Chávez, 2007a,b; Acosta-Hospitaleche and Canto, eudyptes, confirming the criteria of Sallaberry et al. (2010b). 2005; Acosta-Hospitaleche and Canto, 2007; Acosta-Hospitaleche Two size classes of fossil penguins are found in the Chilean et al., 2005; Soto-Acuña et al., 2008). Eocene strata. Although only one big sized humerus could be Three neogene endemic species have being described: Sphe- recognized at the genus level, the remaining bones show size niscus chilensis Emslie and Guerra-Correa (2003), Pygoscelis grandis affinities with other medium and small sized genera from the Walsh and Suárez (2006), and Pygoscelis calderensis Acosta- Eocene of Antarctica such as Marambiornis, Mesetaornis or Delphi- Hospitaleche et al. (2006b). The first one from the La Portada nornis (Sallaberry et al., 2010b). Formation is a specimen of the penguin genus Spheniscus that is Non spheniscid remains are only known from a record assigned present in South American Atlantic and Pacific coasts comprising to Ardeidae also from the Sierra Dorotea locality, a proximal frag- the species S. humboldti and Spheniscus magellanicus and the Gal- ment of tibiotarsus lacking an extension from the crista cnemialis ápagos penguin S. mendiculus. S. chilensis is a medium small sized cranialis in proximal direction, as in other aquatic birds assigned to penguin Pliocene in age. this family, suggesting also it was a non-exclusively marine envi- P calderensis and P. grandis are both recovered from different ronment (Sallaberry et al., 2010b). From strata of Algarrobo, central levels of the Bahía Inglesa Formation. P. calderensis was recovered Chile, an incomplete fragmentary femur is the only record of an from late Miocene levels of phosphatic deposits and comprises Eocene bird from this locality. This does not represent a fossil a species in the size range of the extant members of the genera. penguin, and thus poses question about the environment and P. grandis is a large sized extinct species that was recovered from taphonomical influences during depositation of the sediments in Pliocene levels of the Bahia Inglesa Formation near the size of the this locality. It also comprises a locality that is intermediate in extant king penguin (Aptenodytes forsteri). latitude between southernmost Chile, Antarctica and fossil penguin From the late Miocene levels of Bahía Inglesa Formation there localities in Southern Peru (Yury-Yáñez et al., in press). The frag- are records belonging to the extant genus Spheniscus and charac- mentary nature of the fossil does not allow more specific taxonomic terized by their big size. Spheniscus urbinai and Spheniscus mega- affinities, but in general anatomy it could be related to an aquatic ramphus are represented by several neurocrania, an isolated bird, most probably a procelariid. rostrum and appendicular bones (Soto-Acuña et al., 2008; Chávez, 2007c, 2006). Only S. urbinai is known from associated skeletons 6. Neogene from the levels of “El Morro” locality from the Bahía Inglesa Formation (Yury-Yáñez et al., 2009) and is also represented (by The Neogene fossil record of modern birds is the most studied and cranial remains) in the Chañaral de Aceituno locality from the abundant of the order Dinosauria. Regardless of the enormous Coquimbo Formation (Chávez, 2005). localities of Cenozoic vertebrates from continental sediments Besides Spheniscidae, seven other families of Neornithes are studied during the last decades (Croft et al., 2008), almost no bird has known from the Chilean Neogene fossil record: Diomedeidae, Pro- been recorded from continental levels, with the exception of Mega- cellariidae, Phalacrocoracidae, Sulidae, Anhingidae, Falconidae and nhinga chilensis. In this sense, as with the Paleogene record, most Pelagornithidae. The order Procellariiformes is the second best birds are also from marine localities in the Neogene. We will intro- represented in Bahía Inglesa Formation (after penguins) and duce first the marine taxa (given the amount of data available) and comprises two families. Diomedeidae (mollymawks or albatrosses) conclude with the fewer continental records. Three formations is recognized by cranial remains of the genus Thalassarche (accord- northern to 30S are the most studied and yield the highest amounts ing to Nunn et al., 1996)orDiomedea (Walsh and Hume, 2001). of fossil birds. These are, from North to South: La Portada Formation Procellariidae (petrels and others) is represented by several genera (late MioceneePliocene, Antofagasta Region), Bahía Inglesa Forma- currently under study, recognizing at least the tribe Puffininni tion (late MioceneePliocene, Atacama Region) and the Coquimbo (particularly the genus Calonectris, Yury-Yáñez et al., 2008), gadfly Formation (middle Miocene-Pliocene, Coquimbo Region). In the petrels comparable to Pterodroma, petrels related to Pagodroma, central part of the country, isolated remains have been recorded in a procelariid of uncertain affinities (probably Pterodroma, Yury- the Horcón Formation (Pliocene, Valparaíso Region). The only Yáñez et al., 2010a) and the genus Pachiptyla (Sallaberry et al., 2007). exclusively continental formation with bird remains is the lacustrine The diversity of sulids (also exclusively from Bahía Inglesa Curamallín Formation (Miocene) in the Biobio Region. Formation) comprises big sized forms with affinities to known 252 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. Tränkner.

species in Pisco Formation (Peru), and two other size classes, all 8. Conclusions and perspectives recognized as belonging to the genus Sula. Three forms of Sulidae are known (Walsh and Hume, 2001; Chávez and Stucchi, 2006; No dinosaur remains have been recorded from the Triassic Soto-Acuña et al., 2009) Sula cf. variegata, Sula magna and Sula cf. period in Chile. However, this trend may be more related to the lack sulita. S. magna should be considered the best represented based in of systematic field work rather than the absence of such remains. cranial remains. Other family, Phalacrocoracidae, is based on The Triassic outcrops of the strata of El Bordo, in the Antofagasta appendicular bones that were described by Walsh and Hume Region, are auspicious since they consist of a sequence of lacustrine (2001). deposits where two forms of non-dinosaurian archosaurs have With the Neogene marine avian fauna, the Pelagornithidae is been found. Projected prospection of such outcrops should be the most outstanding bird family, typically known from Paleogene a priority because of the great importance that these levels repre- and Neogene levels from every continent. Until now, in Chile, it is sent for understanding the evolutionary history of dinosaurs. only recognized from the late Miocene of Bahía Inglesa Formation For now, Jurassic dinosaurs are scarcely represented. The based on several isolated remains and the complete articulated discovery of theropod skeletons in the Aysen Region and the skeleton of Pelagonis chilensis, which Mayr and Rubilar-Rogers deposits of dinosaur footprints from the Antofagasta Region are (2010) considered as the bird with the largest well established promising. Doubtless, more material of theropod dinosaurs can be wingspan, within both living and extinct birds (Fig. 6). This spec- recovered from the strata of “El Toqui” and possibly new prints will imen is also important due to its exquisite preservation, allowing be documented in quebrada Arca. cranialepostcranial correlations which were impossible until this Cretaceous deposits have delivered a very abundant record of finding. Based on Mayr and Rubilar-Rogers (2010), it is highly dinosaur prints that allow us to know about the fauna of the time probable that all Bahía Inglesa Formation material (several isolated despite the absence of skeletal remains. Some formations of this bones) belong to the genus Pelagornis, coinciding with earlier period overlie Jurassic deposits, such that it becomes possible to identifications (Chávez et al., 2007). understand changes in faunal composition across these periods. Undescribed fossil birds from the Pliocene of the Horcón Additionally, the titanosaurs represent the best-documented non- Formation have been mentioned (Carrillo-Briceño et al., 2011)as avian dinosaurs in Chile. Doubtless, new forms of these dinosaurs Charadriiformes, Sphenisciformes and Phalacrocoracidae, but at will continue to be discovered in sites like La Higuera ravine in the least one fossil of Spheniscus from this unit is recognized at the Atacama Region, where new and more complete specimens are genus level (Yury-Yáñez et al., 2010b). likely to be collected. Two records of Neogene birds are ascribed as the only non- As mentioned above, teeth possibly belonging to a dromaeo- marine birds, the Falconidae Milvago sp. from La Portada Forma- saurid are the only cranial remains of non-avian dinosaurs tion (Emslie and Guerra Correa, 2003) and M. chilensis described as currently known from Chile. If this identification is correct, this flightless and big sized snake bird (Anhingidae; Alvarenga, 1995). evidence would be the first record of this kind of dinosaur from Chile and one of the few currently known for Gondwana. 7. Quaternary The only record of birds from the cretaceous in Chile is two isolated tarsometatarsi from the Quiriquina Formation. This A large number of unpublished materials (hundreds of isolated material is important since it is one of the few occurrences of specimens) have been recovered from the Pleistocene of San Mesozoic birds known for Gondwana. New field work in this Vicente de Taguatagua, mainly from plaster jackets of vertebrate formation is necessary in order to find new evidence of these birds. remains recovered by Lautaro Núñez and Rodolfo Casamiquela in Until now, there is no information from Chile about other the 1980’s. These comprise well-preserved and diagnostic remains terrestrial taxa that must have co-existed with dinosaurs during the that will allow determining temporal variation in the taxa Mesozoic, such as mammals and ophidians. This fact is defined composition of the zone from the Pleistocene to present. mainly by the lack of systematic search for other components of the Bird remains have also been found at the Pleistocene locality of Mesozoic biota. Reports of lizards and amphibians must be Pilauco Bajo, in the city of Osorno which have been informally corroborated since current mentions are unclear and the destiny of mentioned and are currently under study (Montero et al., 2008). The these materials is unknown. collection of the department of Geology of the University of Chile Continental Mesozoic vertebrates are very important to narrow houses undetermined fragments from the Pleistocene of Mocha down possible ages of continental levels (Blanco et al., 2000; Island of penguins of similar size to the modern genus Spheniscus that Rubilar-Rogers, 2003). Thus they will continue to be a good tool for were collected by Tavera and Veyl in 1958, but remain unstudied. establishing biochrones of Mesozoic continental levels in Chile. D. Rubilar-Rogers et al. / Journal of South American Earth Sciences 37 (2012) 242e255 253

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