A New Genus for the Cuban Teratorn (Aves: Teratornithidae)
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 122(1):103–116. 2009. A new genus for the Cuban teratorn (Aves: Teratornithidae) William Sua´rez and Storrs L. Olson* (WS) Departamento de Paleogeografı´a y Paleobiologı´a, Museo Nacional de Historia Natural, Obispo 61, Plaza de Armas, La Habana, CH 10100, Cuba, e-mail: [email protected]; (SLO) Department of Vertebrate Zoology, National Museum of Natural History, P. O. Box 37012, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A, e-mail: [email protected] Abstract.—Study of the type series and much new fossil material of the Cuban teratorn, Teratornis olsoni Arredondo & Arredondo, shows that this species possessed unique characters within the family Teratornithidae, including a shorter and more flattened humerus and femur, and a tarsometatarsus with a long trochlea II. The differences are so great as to merit a new genus, Oscaravis, for the species. Some osteological characters of Oscaravis suggest that it was less derived than Teratornis and possibly more similar to Argentavis. As the only insular member of the Teratornithidae, Oscaravis shows that teratorns were capable of overwater dispersal, so that the expansion of the family into North America from the south need not have been dependent on the presence of a land bridge. Resumen.—El estudio de la serie tipo y de nuevo material fo´sil del teratorno cubano, Teratornis olsoni Arredondo & Arredondo, muestran que este taxon presento´caracteres u´nicos dentro de la familia Teratornithidae, que incluyen hu´meroy fe´mur ma´s cortos y aplanados, y tarsometatarso con tro´clea II alargada. La magnitud de las diferencias existentes permite erigir un nuevo ge´nero, Oscaravis, para esta especie. Algunos caracteres osteolo´gicos de Oscaravis indican que divergio´ menos que Teratornis,y por ello estuvo ma´s cercano morfolo´gicamente a Argentavis.Comou´nico miembro insular en Teratornithidae, Oscaravis evidencia que los teratornos fueron capaces de dispersarse sobre barreras acua´ticas, por lo que la expansio´n de la familia a Norte Ame´rica, desde el sur, no dependio´dela presencia de un puente terrestre. The extinct family Teratornithidae L. of the genera, was long known only from Miller (1909), the ordinal affinities of two tarsometatarsi from Rancho La Brea, which remain obscure, is known only California, that were regarded as belong- from the New World and at present ing to the Vulturidae (L. Miller 1910, includes five genera: Teratornis L. Miller 1925), although the genus was later (1909), Cathartornis L. Miller (1910), assigned to the Teratornithidae (L. Miller Argentavis Campbell & Tonni (1980), & Howard 1938). Campbell et al. (1999) Aiolornis Campbell et al. (1999), and considered the grounds for separating Taubatornis Olson & Alvarenga (2002). Cathartornis from Teratornis to be weak, With the exception of Teratornis, all are although additional specimens of C. monotypic (see Campbell & Stenger gracilis L. Miller from Rancho La Brea 2002). Cathartornis, the most problematic were recently reported by Campbell & Stenger (2002:3). Additional elements * Corresponding author. continue to be identified, although it is 104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON uncertain whether their differences from A subset of those fossils was later named as Teratornis warrant separate generic status a new species, Teratornis olsoni Arredondo (K. E. Campbell, Jr., pers. comm.). & Arredondo (2002), which was described The Teratornithidae appears to have in comparison with T. merriami. Addi- originated in South America and was tional material of T. olsoni was collected in thought to have colonized North America Cuba, or identified in collections by WS, during the Great American Faunal Inter- always in association with remains of the change in the late Pliocene (Campbell & Cuban condor Gymnogyps varonai (Arre- Tonni 1981) or perhaps earlier (Olson & dondo, 1971) and with large, extinct Alvarenga 2002). Taubatornis campbelli members of the Accipitridae (see Sua´rez Olson & Alvarenga (2002), from deposits 2000a, b, 2004). Comparison of T. olsoni of the Tremembe´ Formation (Upper with other teratorns, especially T. mer- Oligocene or Lower Miocene, ca. riami, shows that it should be referred to a 25 mya), Taubate´ Basin, Sa˜o Paulo, new genus, described herein. This brings to Brazil, is the oldest and smallest known six the number of genera in the Terator- teratorn (Olson & Alvarenga 2002). It nithidae and provides additional informa- was followed temporally by the largest tion on the evolution and biogeography of known flying bird, Argentavis magnificens the family. Campbell & Tonni (1980), from four localities of late Miocene age (Huayquer- Materials and Methods ian, ca. 6 mya) in Argentina (Campbell & Tonni 1980, Campbell 1995). In addition, Specimens of the Cuban teratorn de- a fragmentary tarsometatarsus apparently scribed herein are housed in the following related to Teratornis was reported from institutional and private collections in La La Carolina, Ecuador (Campbell & Habana, Cuba: Museo Nacional de His- Tonni 1980), which is the only evidence toria Natural de Cuba (MNHNCu); of the family in the Quaternary of South Instituto de Ecologı´a y Sistema´tica America. (CZACC: Colecciones Zoolo´gicas de la Apart from three specimens referred to Academia de Ciencias de Cuba); Oscar Aiolornis incredibilis (Howard 1952) from Arredondo collection (OA); Osvaldo Ji- late Pliocene (Blancan) deposits (Camp- me´nez collection (OJ); William Sua´rez bell et al. 1999), the North American re- collection (WS). We examined all the cord of Teratornithidae is entirely Pleisto- Cuban specimens previously described cene in age. Our knowledge of the mor- or referred to Teratornithidae, including phology of the family is based mainly on the holotypical right femur (CZACC 400- remains of Teratornis merriami L. Miller 649) of Teratornis olsoni. Comparisons from the asphalt deposits at Rancho La with skeletons of large modern members Brea, although the osteology of this of the Vulturidae (Vultur and Gymnogyps) species is still only incompletely described. were made in the skeletal collections of All other taxa of Teratornithidae are the National Museum of Natural History, known from much less complete material Smithsonian Institution (USNM), Wash- that usually can be interpreted only ington, D.C. Comparisons were made through comparisons with Teratornis. with extensive material of Teratornis That teratorns existed outside of the merriami in the collections of the George continental Americas was shown by C. Page Museum, a branch facility of the Sua´rez & Arredondo (1997), who record- Natural History Museum of Los Angeles ed seven specimens that were tentatively County, including the following speci- identified as Teratornis sp. from Quater- mens sent on loan: left quadrate LACM nary cave deposits in western Cuba (Fig. 1). HC F3452; left coracoid LACM K3303; VOLUME 122, NUMBER 1 105 Fig. 1. Map of western Cuba showing Quaternary fossil localities in which the Cuban teratorn Oscaravis olsoni has been recovered. right scapula LACM B103; left humerus Gymnogyps) and agree with Teratornis in LACM B261; right tibiotarsus LACM the following characters. Quadrate larger B1044; left tarsometatarsus LACM B566. and massive, with L-shaped mandibular A cast of right femur LACM B788 was articulation; socket for quadratojugal used for Fig. 7. We also examined the much larger. Coracoid with procoracoid holotypical distal end of right tibiotarsus process projecting more dorsad (rather (MHNT-VT 5154) of Taubatornis camp- than laterad); procoracoid foramen larger belli from the Museu de Historia Natural and more sternally placed (not true of de Taubate´, Sa˜o Paulo, Brazil. Compar- Argentavis, however). Scapula with longer isons with Cathartornis gracilis (L. Miller and more pointed acromion; shorter 1910), Argentavis magnificens (Campbell and curved furcular articulation; shaft & Tonni 1980, Campbell 1995), and straighter ‘‘in both planes’’ as noted by L. Aiolornis incredibilis (Howard 1952, Miller (1925:90). Humerus with long, Campbell et at. 1999) relied on published straight, deep ligamental furrow; pectoral descriptions and illustrations. Osteologi- crest ‘‘a thick, lumpy ridge with almost no cal terminology follows Baumel et al. [distal] tubercle’’ that ‘‘overhangs the (1979) and Howard (1929), with some bicipital furrow’’ (L. Miller 1925:90); necessary modifications. Measurements shaft straighter and very concave antero- were taken with digital calipers to the proximally. Ulnare with attachment for nearest 0.1 mm. lig. ulno-ulnocarpale long, diagonal, and ridge-like, located close to the external Systematic Paleontology prominence (Howard 1952:51). Femur with head relatively larger and very Order Incertae Sedis indistinctly set off from the shaft; tro- Family Teratornithidae L. Miller, 1909 chanter weakly developed without a deep The Cuban fossils differ from compa- depression at its base; pneumatic open- rable bones of Vulturidae (Vultur and ings (proximal end) small and not aligned 106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in a large oval depression below the trochanter; anterior surface of shaft with a distinct anterior intermuscular line near the midline; shaft much more robust, with flat posterior face of the proximal end. Tibiotarsus with shaft above condyles wider; distal articular surface of the external condyle flatter with the intercon- dylar groove wider and less distinct; internal condyle (anterior view) much shorter or reduced. Tarsometatarsus