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CONDOR Monday Dec 13 2004 03:19 PM cond 107_115 Mp_107 Allen Press x DTPro System GALLEY File # 15TQ The Condor 107:107±113 q The Cooper Ornithological Society 2005 A NEW CONDOR (CICONIIFORMES, VULTURIDAE) FROM THE LATE MIOCENE/EARLY PLIOCENE PISCO FORMATION, PERU MARCELO STUCCHI1,3 AND STEVEN D. EMSLIE2,4 1AsociacioÂn Ucumari. Jr. Los AgroÂlogos 220, Lima 12, Peru 2Department of Biological Sciences, University of North Carolina, Wilmington, NC 28403 Abstract. We report the oldest fossil condor (Vulturidae) from South America and the ®rst from the Pisco Formation (14.0±2.0 Ma) of Peru, described herein as Perugyps diazi new genus and species. The Pisco Formation, exposed on the southern coast of Peru, has produced well-preserved and abundant marine and terrestrial vertebrate fossils from the late Miocene/early Pliocene (6.0±4.5 Ma) Montemar and Sacaco Sur localities, from where P. diazi was recovered. The new condor adds to our knowledge on the evolution and biogeo- graphic distribution of New World vultures. The age of this new species supports the hy- pothesis that condors probably evolved in North America and entered South America by the late Miocene/early Pliocene. We believe it is likely that the ®rst condors to reach South America probably did so via a coastal corridor along the western side of the Andes where they became part of the diverse coastal fauna in southern Peru. Key words: condor, Miocene, Peru, Perugyps diazi, Pisco Formation, Pliocene, Vultur- idae. Un Nuevo CoÂndor (Ciconiiformes, Vulturidae) del Mioceno TardõÂo-Plioceno Temprano de la FormacioÂn Pisco, PeruÂ. Resumen. Se reporta el coÂndor maÂs antiguo de AmeÂrica del Sur y el primero para la FormacioÂn Pisco (14±2 Ma), y se describe como Perugyps diazi. De esta formacioÂn, situada en la costa sur del PeruÂ, provienen gran cantidad de aves marinas en muy buen estado de conservacioÂn, en especial de los niveles Montemar y Sacaco Sur (Mioceno tardõÂo/Plioceno temprano, 6.0±4.5 Ma), justamente de donde procede Perugyps. Este nuevo coÂndor anÄade importante informacioÂn sobre la evolucioÂn y distribucioÂn biogeogra®ca de estas aves, pues su edad apoya la hipoÂtesis de que los coÂndores probablemente evolucionaron en AmeÂrica del Norte y entraron a AmeÂrica del Sur entre el Mioceno tardõÂo y el Plioceno temprano. Sugerimos que su llegada pudo realizarse por el corredor costero del lado occidental de los Andes, en donde pasaron a formar parte de la diversa fauna del sur del PeruÂ. INTRODUCTION cene tar seeps at Talara on the northern coast The oldest record of the family Vulturidae in the (Campbell 1979). Recently, eight disassociated Americas is the early Oligocene genus Phas- bones of a new species of condor were found in magyps Wetmore 1927 from the United States deposits of the Montemar (6±4.5 Ma) and Sa- (see Emslie 1988a), though the validity of this caco Sur (5 Ma), vertebrate-bearing levels of the taxon has been questioned (Olson 1985). New Pisco Formation on the south-central coast of World vultures appeared in South America by Peru (Fig. 1). These fossils are assumed to rep- the late Oligocene/early Miocene in Brazil with resent a single genus and species of condor de- Brasilogyps Alvarenga 1985, a species very scribed herein. similar in size and features to the living Co- ragyps (Emslie 1988a). In Peru, fossils of ®ve METHODS modern genera (Vultur, Gymnogyps, Sarcoramp- All fossils were compared with skeletal material hus, Coragyps, and Cathartes) and one extinct of living genera of condors and vultures (Vultur genus (Geronogyps Campbell 1979) have been Linnaeus 1758, Sarcoramphus DumeÂril 1806, identi®ed and described, all from late Pleisto- Cathartes Illiger 1811, Gymnogyps Lesson 1842, Coragyps Saint-Hilaire 1853) at the U. S. National Museum of Natural History, Smithso- Manuscript received 29 September 2003; accepted 4 October 2004. nian Institution (USNM), and the Field Museum 3 E-mail: [email protected] of Natural History (FMNH), Chicago. Because 4 Corresponding author: E-mail: [email protected] current Peruvian law does not allow the removal [107] CONDOR Monday Dec 13 2004 03:19 PM cond 107_115 Mp_108 Allen Press x DTPro System GALLEY File # 15TQ 108 MARCELO STUCCHI AND STEVEN D. EMSLIE FIGURE 2. (A) Holotype right carpometacarpus (MUSM 213) of Perugyps diazi new genus and species in internal (left) and external (right) views. (B) Prox- imal right carpometacarpus (MUSM 206) of Perugyps FIGURE 1. Map of the Pisco Formation in Peru diazi new genus and species in internal view. (C) Dis- showing the location of the Montemar and Sacaco Sur tal left ulna (MUSM 423) of Perugyps diazi new genus localities where fossils of Perugyps diazi new genus and species in internal view. Scale bar 5 2 cm. and species were recovered (after de Muizon and DeVries 1985). Inset map depicts the location of the Pisco Formation in relation to Lima, the capital of RESULTS Peru. SYSTEMATIC PALEONTOLOGY Order Ciconiiformes (Bonaparte 1854) Family Vulturidae (Illiger 1811) of original fossil material from the country, nor Perugyps gen. nov. could casts be made due to the fragile nature of Perugyps diazi sp. nov. (Fig. 2±4) the bones, all comparisons outside of Peru were Holotype. Right carpometacarpus missing proximal portion of os metacarpale minus completed with high-quality digital photographs (MUSM 213; Fig. 2A), collected by M. Stucchi, (SONY Cyber-shot DSc-P92, 5.0 megapixels) of July 2000. the fossils, in 2±3 views per element, as well as 8 Locality/horizon. Pisco Formation, Montemar with 360 digital videos of each specimen, with vertebrate-bearing locality (late Miocene/early modern skeletal material. Thus, we are con®dent Pliocene; 6.0±4.5 Ma; de Muizon and DeVries that all characters that distinguish the new fossil 1985; DeVries, pers. comm.). species could be discerned from these compari- Diagnosis. Perugyps is diagnosed as a condor sons. by the following characters: Comparisons to other fossil taxa were com- 1. Mandible with symphysis proportionately pleted using published illustrations and descrip- larger than in Coragyps, Cathartes melam- tions, and casts of Royal Ontario Museum fos- brotus Wetmore 1964, and Gymnogyps; sils (ROM 12991, 12992, 12993, 13007) of Ge- smaller than in Sarcoramphus and Vultur; ronogyps reliquus Campbell 1979. Terminology and similar in proportions to Cathartes aura follows that of Howard (1929) and Baumel (Linnaeus 1758). Symphysis is proportion- (1993); measurements were completed with ver- ately broader and the dentary has a relatively nier calipers to the nearest 0.1 mm. All fossils higher coronoid process than in all living described here are housed at the Departamento Vulturidae. Mandible longer than in Vultur de PaleontologõÂa de Vertebrados, Museo de His- gryphus Linnaeus 1758 and Gymnogyps ca- toria Natural, Universidad Nacional Mayor de lifornianus (Shaw 1798, Table 1). San Marcos (MUSM), Lima, Peru, and are cat- 2. Sixth cervical vertebra with prezygopophyses alogued with MUSM numbers. angled more anteriorly, rounder and more ro- CONDOR Monday Dec 13 2004 03:19 PM cond 107_115 Mp_109 Allen Press x DTPro System GALLEY File # 15TQ A NEW CONDOR FROM PERU 109 TABLE 1. Mandibular meseasurements (mm) of living Vulturidae in comparison with Perugyps diazi new genus and species. Measurement codes are: (1) symphysis length, (2) symphysis width at proximal end, (3) coronoid process height, (4) coronoid process-articular length, and (5) total length of mandible. All measurements are mean 6 SD. Species 1234 5 Perugyps diazi (MUSM 261) 22.5 25.7 23.2 35.1 140.0 Coragyps atratus (n 5 5) 13.4 6 1.3 9.3 6 0.9 10.6 6 0.7 18.5 6 0.7 87.3 6 3.4 Cathartes melambrotus (n 5 2) 10.6 6 0.5 9.5 6 0.3 10.0 6 0.0 14.6 6 1.3 75.0 6 1.9 Cathartes aura (n 5 5) 11.2 6 1.1 8.3 6 0.9 9.1 6 0.2 14.3 6 0.8 70.2 6 1.7 Sarcoramphus papa (n 5 5) 17.0 6 0.7 13.8 6 0.4 13.6 6 0.4 23.8 6 0.6 90.7 6 1.8 Vultur gryphus (n 5 2) 24.0 6 0.8 18.9 6 0.4 18.8 6 1.3 38.3 6 1.7 131.2 6 4.2 Gymnogyps californianus (n 5 4) 18.6 6 1.5 17.4 6 0.6 16.2 6 0.9 37.5 6 0.9 129.5 6 6.3 bust, than in Vultur and Gymnogyps; postzy- 5. Tibiotarsus with tendinal furrow more cen- gopophyses rounder and angled more later- tered than in Vultur, Gymnogyps, Sarcoramp- ally than in Vultur and Gymnogyps. hus, Coragyps, and Cathartes. 3. Coracoid with deep and rounded sternocora- 6. Tarsometatarsus robust with distal external coidal impression that is not pneumatic (shal- trochlea placed as low or lower than middle lower and not rounded, often pneumatic in trochlea (external trochlea higher than middle Vultur, Gymnogyps, Sarcoramphus, Co- in Vultur, Gymnogyps, Sarcoramphus, Co- ragyps, and Cathartes). ragyps, and Cathartes). 4. Carpometacarpus with anterior carpal fossa less pneumatic than in Vultur, Gymnogyps, Genus etymology. From Peru, where the fos- Sarcoramphus, Coragyps, and Cathartes. sils were found, and gyps from Greek, mascu- Proximal symphysis short, as in Vultur; sym- line, vulture. physis relatively longer in Cathartes, Co- Species etymology. Named for Mr. Eusebio ragyps, and Gymnogyps. MUSM 206 has a DõÂaz, in recognition of his contributions to Pe- muscle scar for the ¯exor metacarpi short and ruvian vertebrate paleontology. pronounced, as in Vultur and Breagyps (L. Referred material. MUSM 205, 206, and 260 Miller 1910); scar in less pronounced in are from the Sacaco Sur locale while MUSM Gymnogyps and more so in Cathartes and 204, 261, 263, and 423 are from the Montemar Coragyps.