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Palaeoenvironmental and Faunal Inferences Based on the Avian Fossil Record of Patagonia and Pampa: What Works and What Does Not

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Palaeoenvironmental and Faunal Inferences Based on the Avian Fossil Record of Patagonia and Pampa: What Works and What Does Not Biological Journal of the Linnean Society, 2011, 103, 458–474. With 5 figures Palaeoenvironmental and faunal inferences based on the avian fossil record of Patagonia and Pampa: what works and what does not CLAUDIA P. TAMBUSSI* CONICET, División Palaeontología Vertebrados, Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina Received 28 February 2011; accepted for publication 28 February 2011bij_1658 458..474 Analysing the effect of climatic/environmental changes on bird communities during the South American Cenozoic is quite complicated. Taking into consideration the extremely complex evolution of such environmental conditions and the incomplete and episodic fossil bird record in this part of the continent, any generalization should be considered with caution. However, some aspects may be noted: (1) certain typically South American bird groups evolved in total isolation, i.e. terrestrial or poorly flying birds, incapable of crossing important water barriers (Rheiformes, Tinamiformes, Phorusrhacidae, Brontornithidae, Anhimidae); (2) other good flyers did not cross until immediately before the definitive connection between both Americas (Teratornithidae, Passeriformes Suboscines); (3) most of the families established important intercontinental relationships (Anhingidae, Pelecanidae, Ciconiidae, Anatidae, Presbyornithidae, Rallidae, Falconidae and Accipitridae); (4) several taxa that are presently important members of the rich South American bird fauna are unknown for certain geological time periods (Throchilidae); and (5) there is a high prevalence of carnivorous birds over all other trophic habits, regardless of the association or age analysed. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474. ADDITIONAL KEYWORDS: Aves – biogeography – Cenozoic – faunal associations – palaeoenvironments. Analizar el efecto que los cambios climáticos y ambientales tuvieron en las comunidades de aves durante el Cenozoico sudamericano es complicado y cualquier generalización debe tomarse con cautela. Sin embargo pueden señalarse algunos aspectos: (1) algunos grupos de aves típicamente sudamericanas evolucionaron en total aislamiento (Rheiformes, Tinamiformes, Phorusrhacidae, Brontornithidae, Anhimidae); (2) otros grupos buenos voladores no cruzaron a América del Norte hasta establecido el puente Panameño entre las dos Américas (Teratornithidae, Passeriformes Suboscines); (3) la mayoría de las familias establecieron importantes relaciones intercontinentales (Anhingidae, Pelecanidae, Ciconiidae, Anatidae, Presbyornithidae, Rallidae, Falconidae, Accipi- tridae); (4) importantes miembros de la avifauna sudamericana actual son desconocidos en el registro fósil (Throchilidae); (5) hay una prevalencia de aves carnívoras en todas las asociaciones cualquiera fuere su antigüedad. PALABRAS CLAVE: Aves – biogeografía – Cenozoico – Asociaciones faunísticas – palaeoambientes. INTRODUCTION stunted and dwarf plants; in the valleys the same thorn- bearing bushes grow. Everywhere we see the same birds and The country remains the same: terribly uninteresting. The insects []. The curse of sterility is on the land. great similarity in productions is a very striking feature in all Charles Darwin’s Diary of the Voyage of H.M.S. Beagle, Patagonia. The level plains of arid shingle support the same April 22nd, 1834 Patagonia is currently an arid (yearly precipitations *E-mail: [email protected] under 300 mm), cold (mean annual temperature, 458 © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474 PATAGONIAN AND PAMPEAN FOSSIL BIRDS 459 12 °C) and windy territory with low-growing, steppe- sentatives of the earliest Neogene. These exposures adapted shrub vegetation. Only towards the west, in are emblematic as they have been known since the the vicinity of the Andean mountain range, do pre- late 19th century thanks to the work of the brothers cipitations become more abundant and allow the Carlos and Florentino Ameghino, the former recog- development of forests with dense undergrowth nized by his essential field work and the latter by his (Barreda & Palazzesi, 2007) and more abundant and vast written production (Ameghino, 1891, 1895, 1906, varied vegetation. According to phytogeographical 1908). and ecological criteria, Argentina can be divided into Accordingly, the oldest Neornithes (or modern) 18 eco-regions, seven of which are represented in birds from Patagonia (and the whole of Argentina) Patagonia and in the marine coastal sectors and come from the San Jorge Gulf Basin (Chubut and neighbouring islands (Fig. 1). Each of these eco- Santa Cruz Provinces), corresponding fundamentally regions is also characterized by the presence of to the Riochican (middle to late Palaeocene), Casa- certain bird species, both residents and visitors. mayoran (late Eocene) and Deseadan (late Oligocene) Taken as a whole, Patagonia currently hosts about 40 periods (Fig. 1A), whereas the Neogene records origi- nonpasseriform and 14 passeriform bird families nate from the Santacrucian (late early Miocene) of the (Narovsky & Babarskas, 2000) with heterogeneous Austral Basin, up to the most modern Huayquerian concentration in each eco-region, with the prevalence (late Miocene), located on the northern San Jorge of marine birds near the Atlantic Ocean coast, low Gulf Basin, in Chubut Province (Dozo et al., 2010), as bird diversity on the steppe and highest diversity in well as north of Patagonia, in La Pampa Province the mountain forests. (Montalvo et al., 2008; M. N. Cenizo, C. P. Tambussi & Modern-day Patagonia has been moulded by a C. Montalvo, unpubl. data). The vertebrate fossil transformation process resulting from the interaction record in Patagonia is very scarce from the end of the of global tectonics, with palaeogeographical and Miocene to the present. Information for this period palaeoclimatic changes. Two events are key and may thus needs to be complemented with extra- be considered as iconic for this transformation: first, Patagonian (Pampasian) records to better understand the opening of the Drake Passage at some point the macro-events in the Patagonian region (Tonni & during the Eocene–Oligocene transition (c. 28 Ma ago; Carlini, 2008). Madden et al., 2010) isolated Antarctica from Patago- At least two marine transgressions affected the nia, bringing about a trend towards cooler conditions entire South American continent along different cor- (Reguero & Marenssi, 2010), and, second, the rise of ridors during the Miocene. The first transgression the Andes mountain range during the Miocene, which took place between 15 and 13 Ma and was dominated blocked the passage of the humid East winds, induced by eustatic and tectonic factors. The second transgres- a process of desertification. The trend towards global sion took place between 10 and 5 Ma and resulted cooling, which was already evident after the Mid- mainly from tectonic events (Hernandez et al., 2005). Eocene Climate Optimum, became more marked Thus, the continental sediments of the Palaeogene towards the Miocene, although with some inter- and Neogene are interdigitated with highly fossilifer- spersed warmer periods, such as the event known as ous sediments of marine origin containing both ver- the Middle Miocene Climate Optimum (Zachos et al., tebrates and invertebrates. Among birds, penguins 2001; Madden et al., 2010, and references cited are the most frequent (Acosta Hospitaleche et al., therein). According to Pascual & Odreman Rivas 2007, 2008). Marine sediments are represented in the (1971), two areas can be recognized in Patagonia: to San Julián, Monte León and Gaiman Formations the north, the San Jorge Gulf Basin extends between (late Eocene to early Miocene) and the more modern the Somun Curá massif and the Deseado massif Puerto Madryn Formation (middle to late Miocene) (Fig. 1B), and the Austral Basin to the south of the outcropping in the Valdés Peninsula (Dozo et al., Deseado massif (Tonni & Carlini, 2008). This scheme 2010). The greatest of the Miocene transgressions is provides a useful basis for the arrangement of the known as the Paranaensean; it began in the latest bird records discussed here. Oligocene and ended during the early middle Miocene Although the San Jorge Gulf Basin is dominated (15–13 Ma). The final phases of this transgression by Palaeogene sediments (Palaeocene–Oligocene, are coincident with the Neogene Climatic Optimum. approximately between 65 and 23 Ma, which com- The Paranaensean covered much of the Chaco- prises the first part of the Cenozoic era, according to Paranaensean lowlands and eastern Patagonia, Gradstein, Ogg & Smith, 2004), the Austral Basin affecting the current territories of Argentina, comprises mostly sediments from the initial part of Uruguay, parts of Brazil and Paraguay, southern the Neogene (Tonni & Carlini, 2008). The wonderful Bolivia, Venezuela, Colombia and Ecuador. exposures of the Santa Cruz Formation (late early Although the fossil record of birds in the Cenozoic is Miocene) have undoubtedly provided the best repre- incomplete, scarce and consisting mostly of isolated © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103, 458–474 460 C. P. TAMBUSSI Figure 1. A, Palaeocene to Recent time scale following Gradstein et al. (2004) including time scale for Cenozoic mammalian faunas of South America (SALMA). B, Map of Patagonia showing the present-day major ecosystems (1, High Andes; 2, Araucanian Forest; 3, Steppe; 4, Shrub) and
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