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Description of the Earliest Fossil Penguin from South America and First Paleogene Vertebrate Locality of Tierra Del Fuego, Argentina

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Description of the Earliest Fossil Penguin from South America and First Paleogene Vertebrate Locality of Tierra Del Fuego, Argentina PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3423, 18 pp., 9 ®gures, 1 table December 9, 2003 Description of the Earliest Fossil Penguin from South America and First Paleogene Vertebrate Locality of Tierra Del Fuego, Argentina JULIA A. CLARKE,1 EDUARDO B. OLIVERO,2 AND PABLO PUERTA3 ABSTRACT We report the discovery of the ®rst vertebrate from the Paleogene of Tierra del Fuego (Isla Grande), Argentina, in southernmost South America. The specimen consists of parts of an associated pelvic girdle and limb that are identi®ed as belonging to the penguin stem clade (Aves: Pansphenisciformes). The specimen, from an exposure of the Leticia Formation (late middle Eocene), is the earliest known penguin (pansphenisciform) from South America. It is more than 20 million years older than the earliest previously recorded South American pen- guins (from the late Oligocene±early Miocene) and, thus, almost doubles their known record on the continent. A detailed description of the new specimen and a discussion of its implications for the understanding of penguin morphological evolution are provided. The new specimen and other fossil penguins do not currently point to the origin of extant, or crown clade, penguin lineages (Spheniscidae), by the Eocene, only to the divergence of the penguin stem lineage from its sister taxon by this time. The new fossil has several morphologies that differ from all extant penguins but are shared with other fossil penguin taxa, suggesting they may be outside Sphen- iscidae. However, in a discussion of the current status of penguin systematics, we suggest the urgent need for comprehensive phylogenetic analysis of fossil and extant penguins to clarify the timing and pattern of penguin diversi®cation. The specimen was recovered from a newly identi®ed fossil vertebrate locality, an exposure of the Eocene Leticia Formation at Punta Torcida on the Atlantic shore of southeastern Tierra del Fuego, Argentina. The new locality is introduced, and a brief geologic description is made, 1 Division of Paleontology, American Museum of Natural History. e-mail: [email protected] 2 Centro-Austral de Investigaciones Cientõ®cas (CADIC) of the Consejo Nacional de Investigaciones Cientõ®cas y TeÂcnicas (CONICET), CC 92, 9410 Ushuaia, Tierra del Fuego, Argentina. 3 Museo PaleontoloÂgico Egidio Feruglio, Avda. Fontana 140, 9100 Trelew, Chubut, Argentina. Copyright q American Museum of Natural History 2003 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3423 highlighting the potential of the shallow marine sediments of the Leticia Formation for con- tributing to our knowledge of the Paleogene vertebrate fossil record of Tierra del Fuego, and of southern South America, generally. INTRODUCTION go, Eocene outcrops are restricted to the western and northern Andean margin and The associated partial penguin pelvic gir- consist mostly of deep marine deposits, with dle and limb described here was recovered the exception of the shallow marine Leticia in the friable, silty, very ®ne sandstones of Formation. Of these exposed deposits, only the Leticia Formation exposed at Punta Tor- the Leticia Formation has produced a fossil cida on the Atlantic coast of southeastern Ti- vertebrate, that which is described here. erra del Fuego, Argentina (®gs. 1±3). The Fossil penguins are known from the late specimen was collected by MarõÂa I. LoÂpez- Paleocene to late Eocene of Seymour Island, Cabrera and Eduardo B. Olivero during ®eld Antarctica, which lies off of the northeastern study of the stratigraphy, sedimentology, and tip of the Antarctic Peninsula. The Eocene paleontology of the Eocene foreland succes- La Meseta Formation exposed on Seymour sion of the Austral Basin. The vertebrate- Island has produced a taxonomically diverse bearing Leticia Formation sandstones repre- penguin assemblage (Wiman, 1905; Simp- sent estuarine and proximal shelf deposits son, 1971a; Cione et al., 1976; Case, 1992). that have also produced abundant gastropods, Most of this material, including at least six bivalves, and solitary corals (Olivero and LoÂ- species, is from the upper part of the For- pez-Cabrera, 2001; Olivero et al., 2002). Fo- mation, which is late Eocene in age. This up- raminifera and nannoplankton from the Le- per part is currently included in ``Telm 7'' of ticia Formation indicate a late middle Eocene Sadler (1988), or the Allomember Submeseta age for the deposits (MalumiaÂn et al., 1994; of Marenssi et al. (1998), and has yielded a Olivero and MalumiaÂn, 1999; see below). Sr isotopic age of ;34.2 Ma (Dingle and Prior to the recovery of the new specimen, Lavelle, 1998). Sadler (1988) recognized the earliest fossil penguins from South seven informal stratigraphic packages in the America were latest Oligocene or early Mio- La Meseta Formation, ``Telm 1'' to ``Telm cene specimens from Patagonia (Simpson, 7'', and Marenssi et al. (1998) recognized six 1972; Fordyce and Jones, 1990). At the turn unconformity-bounded allomembers. Less of the 19th century, Carlos Ameghino col- abundant penguin material was also recorded lected penguin material at the mouth of the from ``Telm 3'' and ``Telm 4/5'' or Allo- Santa Cruz River in Santa Cruz Province, members Campamento and Cucullaea, re- Argentina, probably from the Monte LeoÂn spectively. These units are assigned a middle Formation, also of late Oligocene±early Mio- Eocene age based on the ;52.4/54.3 (late cene age (Acosta Hospitaleche et al., 2001). early Eocene) Sr isotopic age yielded by the Fossil penguins are additionally known from basal part of the underlying Acantilados Al- latest Oligocene±early Miocene shallow ma- lomember (Reguero et al., 2002). New dis- rine deposits assigned to the Gaiman or Pa- coveries from Seymour Island include pen- tagonia Formation and exposed in the RõÂo guin bones from the late early Eocene Acan- Chubut valley, near the city of Trelew, Pa- tilados Allomember or Telm 2±3 (La Meseta tagonia (Simpson, 1946; Simpson, 1981; Formation) as well as from the late Paleo- MalumiaÂn et al., 1999 and the bibliography cene Cross Valley Formation (S. Marenssi, therein). personal commun. to E.B. Olivero). Although middle and upper? Eocene ma- Paleogene penguins have been described rine sediments have a regional distribution in from the late middle and late Eocene through southern Patagonia, they outcrop only along Oligocene of New Zealand (Fordyce, 1991), the western margin of the Patagonian Andes. and further undescribed material is known These deposits are exposed in the Lago Ar- from the early Eocene (Fordyce, personal gentino±RõÂo Turbio area (MalumiaÂn et al., commun.). They are also known from the 1999). In southern Chile and Tierra del Fue- late Eocene (Simpson, 1957; Jenkins, 1974) 2003 CLARKE ET AL.: FOSSIL PENGUIN 3 and possibly Oligocene (Glaessner, 1955) of is also recommended that the name ``Sphen- Australia (reviewed in Fordyce and Jones, iscidae'' be formally applied to the clade 1990). A partial skeleton and other more comprised of the most recent common an- fragmentary material from ``proto-pen- cestor of all extant penguins and all of its guins'', or wing- propelled divers proposed descendants. However, a formal de®nition of to be closely related to penguins, have ad- Spheniscidae identifying this name as that ditionally been reported from the late Paleo- for the penguin crown clade should await cene or early Eocene of New Zealand (e.g., further consensus on the phylogenetic rela- Fordyce et al., 1986), but have not yet been tionships among extant penguins (e.g., com- described (Fordyce and Jones, 1990; Fordy- pare Kennedy and Page [2002] and clado- ce, personal commun.). gram of O'Hara [1989] published in Wil- liams [1995]) because of the required use of MATERIALS AND METHODS species or specimens as speci®ers for clade names (Cantino and de Queiroz, 2000). In the description of the new specimen, the INSTITUTIONAL ABBREVIATIONS: AMNH, English equivalents of the Latin osteological American Museum of Natural History; nomenclature of Baumel and Witmer (1993) CADIC, Centro Austral de Investigaciones are used. The terms of orientation for the an- Cientõ®cas, Tierra del Fuego, Argentina; atomical position of a bird, as speci®ed by MoNZ, Museum of New Zealand (formally Clark (1993), were followed with one excep- the Dominion Museum; e.g., as cited in tion. The ``time-honored'' terms (Clark, Simpson, 1971b), Wellington, New Zealand; 1993) of zoological nomenclature ``anterior'' OM, Otago Museum, Dunedin, New Zea- and ``posterior'' were used, rather than ``cra- land. nial'' (and ``rostral'') and ``caudal'' as pro- posed by Clark (1993) in the Handbook of GEOLOGIC SETTING OF THE PUNTA Avian Anatomy: Nomina Anatomica Avium TORCIDA LOCALITY (Baumel et al., 1993; see further discussion in Clarke and Norell, 2002). Appendix 1 lists In southeastern Tierra del Fuego, the Late the comparative materials used in preparation Cretaceous±Cenozoic sedimentary ®lling of of the description and systematic discussion. the foreland Austral Basin consists of four All taxonomic names above the species major, unconformity-bounded, synorogenic level used in this paper are used as clade clastic wedges, which are mainly character- names, although most await formal de®nition ized by very thick, deep marine siliciclastic under a system of phylogenetic nomenclature deposits (Olivero and MalumiaÂn, 2002). The (de Queiroz and Gauthier, 1990, 1992) and youngest, Oligocene±Miocene clastic wedge according to the PhyloCode (Cantino and de is mainly exposed outside the folded belt, to Queiroz, 2000). ``Pansphenisciformes'' is the north of Punta Gruesa (®g. 1), and con- used here as a name for all taxa more closely sists of subhorizontal, faulted strata. Within related to extant penguins than to any other the fold and thrust belt, to the south of Punta extant avian taxa. ``Sphenisciformes'' is sug- Gruesa, the deposits of the older clastic gested as a name for all parts of this lineage wedges of Maastrichtian±Danian, late Paleo- with a loss of aerial ¯ight homologous with cene±early Eocene, and late middle to late that of extant penguins.
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