First Articulated Skeleton of Palaeeudyptes Gunnari
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Antarctic Science 22(3), 289–298 (2010) & Antarctic Science Ltd 2010 doi:10.1017/S0954102009990769 First articulated skeleton of Palaeeudyptes gunnari from the late Eocene of Isla Marambio (Seymour Island), Antarctica CAROLINA ACOSTA HOSPITALECHE* and MARCELO REGUERO Divisio´n Paleontologı´a Vertebrados, Museo de La Plata, Paseo del Bosque s/n, B1900FWA, La Plata, CONICET, Argentina *[email protected] Abstract: The first articulated skeleton of a penguin from the late Eocene of Antarctica is described. MLP 96-I-6-13 comes from the upper Submeseta Allomember (La Meseta Formation) of Isla Marambio (locality DPV 10/84). The significance of this finding in the context of the Anthropornis nordenskjoeldi biozone is discussed. An osteologic description of the recovered elements and a brief discussion of its systematic determination are provided. MLP 96-I-6-13 is the first articulated skeleton with sure specific assignment to Palaeeudyptes gunnari (Wiman, 1905), a species previously known only through isolated tarsometatarsi and included in the groups of Wiman. Received 26 August 2009, accepted 12 November 2009 Key words: Anthropornis nordenskjoeldi biozone, fossil penguin, Palaeogene, Spheniscidae Introduction In the present contribution, the first articulated skeleton from Antarctica is studied. An osteologic description of the The number of penguin species (fourteen) and size classes recovered elements and a brief discussion about its (six) obtained from the late Eocene localities of Isla Marambio systematic assignment are given. The significance of this (La Meseta Formation, Elliot & Trautmant 1982), Antarctica, finding in the context of the Anthropornis nordenskjoeldi exceed the highest number of sympatric species (four) and size biozone is also discussed. classes currently occurring in regional breeding from the cold sub-Antarctic to cool temperate waters south of the Subtropical Convergence. There may be palaeobiological or taphonomical Material and methods reasons for this high percentage, but it is probable that it also reflects a real abundance of penguins, which appear to have During the 2008 field season on Isla Marambio one of the been the dominant coastal birds in the late Eocene of authors (M.R.) collected in situ a partially complete skeleton Antarctica (Case 1996). Even though La Meseta Formation of a fossil spheniscid. The fossil studied here comes from is one of the richest units in terms of diversity and abundance the lower level of the Submeseta Allomember, La Meseta of penguins, no articulated or associated skeleton has Formation (Marenssi et al. 1998a) or TELM 7 (Elliot & previously been described. The Anthropornis nordenskjoeldi Trautman 1982), which outcrops at several localities. The biozone defined within the Submeseta Allomember constitutes penguin-bearing horizon is composed of thick shelly the unit with highest specific diversity of fossil penguins ever conglomerates, well-sorted sands and interlaminated sand/ known, with fourteen recorded species (Tambussi et al. 2006, mud channel-fills with thin shelly conglomeratic intervals. however see Jadwiszczak 2006b). This type of deposit, with Specimen MLP 96-I-6-13 and the comparison materials high abundance of isolated elements, is not exclusive of are housed at the Divisio´n Paleontologı´a de Vertebrados of Antarctic sediments, but it is also characteristic of most fossil the Museo de La Plata (MLP), Argentina. Diagnostic penguin assemblages in the world. characters for Antarctic penguin species are taken from As in other cases, the remains are largely accumulated Myrcha et al. (2002). Osteological terminology follows close to nesting areas of the breeding colonies. These Baumel & Witmer (1993). Measurements were taken with deposits are usually represented by isolated and sometimes a Vernier Caliper (0.01 mm accuracy). fragmented bones. This taphonomic characteristic of the record has compelled palaeontologists to propose a Geological and depositional setting systematic scheme entirely based on isolated skeletal remains. A consequence of this is that most species are A series of prolific fossil sites and quarries are now known known only through a single element, the tarsometatarsus from La Meseta Formation in Isla Marambio, James Ross in most cases, or the humerus in a few others. Only a small Basin (del Valle et al. 1992), Antarctic Peninsula, as a number of fossil species, whose diagnoses are recent (Slack result of work conducted by various research groups since et al. 2006, Acosta Hospitaleche et al. 2007, Clarke et al. the mid 1980s (Fig. 1). The La Meseta Formation is 2007), have been described from articulated elements. composed of sandstones and mudstones with interbedded 289 http://journals.cambridge.org Downloaded: 11 Dec 2013 IP address: 62.173.43.73 290 C. ACOSTA HOSPITALECHE & M. REGUERO Fig. 1. a. Map showing the location of Isla Marambio, Antarctic Peninsula. b. Sketch map of the northern part of Isla Marambio showing the distribution of the Submeseta Allomember and the fossil penguin-bearing locality cited in the text. shell-rich conglomerates, organized into six erosionally- parasequences where this specimen was deposited based internal units, named from base to top Valle de Las (Marenssi 2006). The age of the fossiliferous interval is Focas, Acantilados, Campamento, Cucullaea I, Cucullaea constrained by the age of the basal unconformity (Marenssi II and Submeseta allomembers (Fig. 2). These units 2006, 36 Ma) and the Sr-derived age of Dingle & Lavelle were deposited during the Eocene in deltaic, estuarine and (1998) at 34.2 Ma. shallow marine settings, mostly within a north-west– Marine vertebrates such as the gadiform ‘‘Mesetaichthys’’, south-east trending valley (Marenssi et al. 1998a, 1998b). afewsharks(Pristiophorus and Carcharias), and a primitive The fossil presented here comes from the upper-level mysticete whale (Llanocetus denticrenatus)aswellas Submeseta Allomember in the south-western slope of the archaeocetes are also present in this zone (personal plateau (meseta) of the island (Fig. 1). Almost all fossil observations). A small number of penguin bones assigned to vertebrate specimens from the Submeseta Allomember were Anthropornis and Delphinornis were found close to the found in a stratigraphic interval that is easily distinguishable by specimen described here. the massive occurrence of penguin bones and the phosphatic Sedimentologically, the locality DPV 10/84 is included brachiopod ‘‘Lingula’’, and stratigraphically located 30–35 m in a facies association (facies association II of Marenssi below the top of the 145 m thick Submeseta Allomember, et al. 1998b), and is characterized by conglomeratic beds to within the Anthropornis nordenskjoeldi biozone (Tambussi mudstones with a diverse and abundant macrofauna that et al. 2006, but see Jadwiszczak 2006b). The locality DPV correspond to a valley-confined estuary mouth/inner 10/84 is located 70–75 m below this rich vertebrate-bearing estuary complex. Tidal channels and mixed flats, tidal horizon (Fig. 2b), and approximately 20 m above the contact inlets and deltas, and washover and beach environments between the Cucullaea II and Submeseta allomembers. The represent the interfingering of high and low energy base of the Submeseta Allomember is correlated with environments (Marenssi et al. 1998b). Penguin bones the main lowstand at 36 Ma, and the first part of this unit is recovered are usually disarticulated; some are broken; characterized by estuarine heterolithic sediments followed most are complete albeit with various degrees of abrasion. by a set of aggradational-retrogradational shoreface Most if not all of them were transported at least for a short http://journals.cambridge.org Downloaded: 11 Dec 2013 IP address: 62.173.43.73 FIRST ARTICULATED EOCENE PENGUIN FROM ANTARCTICA 291 Fig. 2. a. Stratigraphic section of the La Meseta Formation, Isla Marambio, Antarctic Peninsula (modified from Reguero et al. 2002). b. Measured section from the eastern flank of the plateau (stratotype of the Anthropornis nordenskjoeldi biozone) showing stratigraphic levels of the Cucullaea II and Submeseta Allomembers (modified from Marenssi 1995). Shaded rectangular area delimits the Anthropornis nordenskjoeldi biozone. time before burial and therefore their accumulations to be the first fossil species of penguin to be described, and represent parauthoctonous assemblages. consequently became the type species of Palaeeudyptes. A partial associated skeleton that included elements of the wing and leg was described by Hector (1872) and The Palaeeudyptes dilemma assigned to P. antarcticus. Subsequently, other remains, Before dealing with the descriptive analysis of the specimen which were originally attributed to this same individual under study, we need to introduce the taxonomic problem that (Hector 1873), were reviewed by Oliver (1930) and pervades this genus. Four species have been nominated for assigned to Pachydyptes ponderosus Oliver, 1930. this genus. Palaeeudyptes antarcticus Huxley, 1859 happened However, the presence of an associated tarsometatarsus http://journals.cambridge.org Downloaded: 11 Dec 2013 IP address: 62.173.43.73 292 C. ACOSTA HOSPITALECHE & M. REGUERO among these remains is not mentioned in these later works. any element other than a tarsometatarsus to P. klekowskii and The subsequent references to this skeleton made by P. gunnari, at least until other articulated remains are found. Lambrecht (1933) and Marples (1952) confirm the However, Jadwiszczak (2006a) assigns different skeletal