CONDOR Monday Dec 13 2004 03:19 PM cond 107_115 Mp_107 Allen Press • DTPro System GALLEY File # 15TQ

The Condor 107:107±113 ᭧ 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: asociacion࿞[email protected] of Natural History (FMNH), Chicago. Because 4 Corresponding author: E-mail: [email protected] current Peruvian law does not allow the removal

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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 ϭ 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 Њ 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 • 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 Ϯ SD.

Species 1234 5 Perugyps diazi (MUSM 261) 22.5 25.7 23.2 35.1 140.0 Coragyps atratus (n ϭ 5) 13.4 Ϯ 1.3 9.3 Ϯ 0.9 10.6 Ϯ 0.7 18.5 Ϯ 0.7 87.3 Ϯ 3.4 Cathartes melambrotus (n ϭ 2) 10.6 Ϯ 0.5 9.5 Ϯ 0.3 10.0 Ϯ 0.0 14.6 Ϯ 1.3 75.0 Ϯ 1.9 Cathartes aura (n ϭ 5) 11.2 Ϯ 1.1 8.3 Ϯ 0.9 9.1 Ϯ 0.2 14.3 Ϯ 0.8 70.2 Ϯ 1.7 Sarcoramphus papa (n ϭ 5) 17.0 Ϯ 0.7 13.8 Ϯ 0.4 13.6 Ϯ 0.4 23.8 Ϯ 0.6 90.7 Ϯ 1.8 Vultur gryphus (n ϭ 2) 24.0 Ϯ 0.8 18.9 Ϯ 0.4 18.8 Ϯ 1.3 38.3 Ϯ 1.7 131.2 Ϯ 4.2 Gymnogyps californianus (n ϭ 4) 18.6 Ϯ 1.5 17.4 Ϯ 0.6 16.2 Ϯ 0.9 37.5 Ϯ 0.9 129.5 Ϯ 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. The scar is longer and pronounced locale of the Pisco Formation (Fig. 1). Right in Sarcoramphus. The intermetacarpal tuber- mandible with distal symphysis, MUSM 261 osity is low in Perugyps, similar to Vultur, (Fig. 3A); cervical vertebra, MUSM 263; sternal Gymnogyps californianus, and Breagyps, and end of right coracoid, MUSM 205 (Fig. 3B); dis- higher in Sarcoramphus, Cathartes, Co- tal left ulna, MUSM 423 (Fig. 2C); proximal ragyps, and G. kofordi Emslie 1988. right carpometacarpus, MUSM 206 (Fig. 2B),

FIGURE 3. (A) Right mandible with distal symphysis (MUSM 261) of Perugyps diazi new genus and species in dorsal (top) and lateral (bottom) views. (B) Sternal end of right coracoid (MUSM 205) of Perugyps diazi new genus and species in ventral (left) and dorsal (right) views. Scale bar ϭ 1 cm. CONDOR Monday Dec 13 2004 03:19 PM cond 107_115 Mp_110 Allen Press • DTPro System GALLEY File # 15TQ

110 MARCELO STUCCHI AND STEVEN D. EMSLIE

long and proximally curved os metacarpalis alu- lare, with a rounded extensor process; a broad contour of the carpal trochlea, and rounded fac- ets on the distal articular surface. The carpo- metacarpus of Perugyps (MUSM 206, 213) pre- sents at least three clear condor characteristics including (1) a proximally curved process ex- tensorius, (2) low intermetacarpal tuberosity, and (3) large size (Hertel 1992). In addition to characters above in the generic diagnosis, further characters distinguish Pe- rugyps diazi. Mandible. In dorsal view, the distal symphy- sis of MUSM 261 (Fig. 3A) is distinctly longer than in Coragyps, Sarcoramphus,or Gymnogyps, exceeded in length only by Vultur; the symphysis is relatively broader in Perugyps compared to all living Vulturidae. The symphy- FIGURE 4. (A) Distal left tibiotarsus (MUSM 260) sis in Cathartes aura is proportionally the same of Perugyps diazi new genus and species in anterior length as in Perugyps. Vultur and Gymnogyps view. (B) Right tarsometatarsus (MUSM 204) of Pe- also have a longer and more pronounced (high- rugyps diazi new genus and species in anterior (left) er) proximal end and articular (Table 1). In ad- and posterior (right) views. Scale bar ϭ 2 cm. dition, the ventral surface of the mandible in lat- eral view curves downward more sharply to- distal left tibiotarsus, MUSM 260 (Fig. 4A); wards the distal end in Gymnogyps compared to right tarsometatarsus, MUSM 204 (Fig. 4B). Perugyps (Fig. 3A) and Vultur. Measurements. See Table 1 for mandibular Cervical vertebra. MUSM 263 is more similar measurements. Measurements that are approxi- to Vultur than Gymnogyps in size, shape, and mate due to erosion on the bone are indicated robustness. MUSM 263 also is similar in char- by the ϳ symbol. Holotype right carpometacar- acters to Cathartes, though much larger. In an- pus (MUSM 213): total length, 147.2 mm; prox- terior view, Perugyps has external borders of the imal breadth, 38.4 mm; least breadth and depth prezygopophyses positioned at the same height of shaft, 26.9 and 23.1 mm, respectively. Prox- as the diapophyses, similar to Sarcoramphus and imal right carpometacarpus (MUSM 206): prox- Cathartes; these borders are relatively higher in imal breadth, 40.2 mm. Distal left ulna (MUSM Coragyps. No vertebrae of Breagyps or Gero- 423): distal breadth and depth, 17.8 mm and nogyps were available for comparison. 24.9 mm, respectively. Distal left tibiotarsus Coracoid. MUSM 205 (Fig. 3B) has an inter- (MUSM 260): distal breadth and depth, 27.6 nal distal angle that is more robust, curved, and mm and 26.2 mm, respectively. Right tarso- with a lower-hanging rim and the line for muscle metatarsus (MUSM 204): total length, ϳ141 attachment that is more distinct and extends far- mm; proximal breadth, 28.7 mm; least breadth ther up the shaft in Perugyps, compared to all 15.7 mm and depth of shaft 8 mm; middle troch- living genera of Vulturidae. Geronogyps (ROM lea breadth ϳ11.3 mm and depth ϳ14.7 mm; 12991, 12992, 12993) is more like Vultur in the distal breadth, ϳ40 mm. sterno-coracoidal impression, pneumatization, and shaft of internal distal angle. DESCRIPTION Ulna. MUSM 423 (Fig. 2C) has a very pro- The Family Vulturidae is characterized by the nounced and robust process lateral to the inter- presence of a deep and often pneumatic anterior nal condyle with a small pneumatic area at the carpal fossa in the carpometacarpus, which is base, similar to Vultur and Gymnogyps (less ro- not present or shallow in Ciconiidae and Tera- bust with no or little pneumatic area in Sarco- tornithidae (Emslie 1988b). Other characteristics ramphus, relatively robust with larger pneumatic for Vulturidae are a massive os metacarpale mi- area in Coragyps and Cathartes). The external nus and majus with a broad distal symphysis, a condyle in internal view extends relatively far- CONDOR Monday Dec 13 2004 03:19 PM cond 107_115 Mp_111 Allen Press • DTPro System GALLEY File # 15TQ

A NEW CONDOR FROM PERU 111

ther up the shaft, and is narrower, in Gymnogyps more robust, with broader and deeper distal than in Perugyps and Vultur. This condyle also trochleae, than in Perugyps. is relatively narrower in Perugyps than in Breagyps and the internal condyle is more prom- DISCUSSION inent in the latter. The Pisco Formation consists of tuffaceous Tibiotarsus. MUSM 260 (Fig. 4A) is more sandy siltstones, medium and coarse-grained similar to Gymnogyps than Vultur in the mor- sandstones, shelly sandstones, and to a lesser ex- phology of the distal condyles; the tendinal tent, conglomerates, bedded tuffs, and coquinas opening is relatively higher (more proximal) on that represent littoral environments that were the shaft in Perugyps than in these two species partially protected and close to shore (de Mui- or Coragyps, Cathartes, and Sarcoramphus.In zon and DeVries 1985). Six vertebrate-bearing anterior view, the intercondylar area is deeply levels were identi®ed by de Muizon and DeVries grooved and symmetrical in Perugyps (area is (1985), including those at Montemar and Sacaco shallow and asymmetrical in Vultur and Sur where fossils of Perugyps were recovered. Gymnogyps, very shallow in Cathartes). There The sediments at these two localities re¯ect a is a distinct bony shelf on the internal side of littoral paleoenviroment with protected beaches the tendinal opening in Perugyps, similar to and reefs exposed to marine currents (Marocco Gymnogyps (this shelf is small in Vultur and and de Muizon 1988). Other avian families so nearly absent in Coragyps). The external con- far identi®ed from the Pisco Formation include dyle is relatively more robust in Perugyps com- Spheniscidae, Sulidae, Phalacrocoracidae, Pela- pared to Gymnogyps or Vultur. The tendinal gornithidae, Laridae, Scolopacidae, Procellarii- groove passing below the supratendinal bridge dae and Diomedeidae (de Muizon 1981, de Mui- is straight in Perugyps, Cathartes, and Sarco- zon and DeVries 1985, Cheneval 1993, Stucchi ramphus, but curved internally and proximally 2003). Perugyps is the eighth genus of fossil condors in Coragyps, Gymnogyps, and Vultur (see and condor-like vultures to be described. Of the Campbell 1979). Geronogyps (ROM 13007) has other seven genera, three (Dryornis, Gero- a higher tendinal opening on the shaft, similar nogyps, and Wingegyps) are known from South to Perugyps, as well as a large shelf on the in- America (Brodkorb 1967, Campbell 1979, Al- ternal side. The intercondylar area is deeper in varenga and Olson 2004). Wingegyps cartellei is Geronogyps than in all living vultures, but not a small enigmatic condor from the late Pleisto- as deep as in Perugyps. Perugyps does not have cene of Brazil, no larger than a raven, but with any obvious features that differ from Breagyps characters strikingly similar to Gymnogyps (Al- in this element. varenga and Olson 2004). Dryornis pampeanus Tarsometatarsus. MUSM 204 (Fig. 4B) has Moreno and Mercerat 1891 (early to middle Pli- an anterior metatarsal groove that is deep and ocene, Argentina) is not known by any elements distinct, extending half way down the shaft (ex- shared with Perugyps and cannot be compared tends farther down the shaft in Vultur, (Moreno and Mercerat 1891). In addition, an un- Gymnogyps, Geronogyps, Hadrogyps Emslie described condor also from the middle Pliocene 1988, and Pliogyps Tordoff 1959). The external of Argentina is known by only a proximal ulna border of this groove also is slightly larger than and radius (Tambussi and Noriega 1999) and is the internal in Perugyps, Vultur, and Coragyps; not comparable to Perugyps. these borders are similar in size in Cathartes, Other fossil condors include Hadrogyps ai- Sarcoramphus, and Gymnogyps. The shaft is rel- gialeus from the middle Miocene, California atively robust as in Gymnogyps and Vultur, nar- (Emslie 1988a), and Pliogyps charon Tordoff rower in Coragyps, Cathartes and Sarcoramp- 1959 and P. ®sheri Becker 1986 from the late hus. The shaft also ¯ares only slightly outward Miocene and middle Pliocene, respectively, of at the proximal and distal ends in Perugyps Florida and Kansas (Tordoff 1959, Becker (shaft ¯ares distinctly outward at ends in Vultur, 1986). These two genera are smaller, condor-like Gymnogyps, Geronogyps, Breagyps, Hadrogyps, vultures that represent a parallel lineage of vul- and Pliogyps; more columnar in Hadrogyps and tures to the larger condors. Aizenogyps toomeyae Aizenogyps Emslie 1998). The tarsometatarsus was a large, robust condor from the Pliocene of of Aizenogyps toomeyae is relatively larger and Florida (Emslie 1998). Breagyps clarki is well CONDOR Monday Dec 13 2004 03:19 PM cond 107_115 Mp_112 Allen Press • DTPro System GALLEY File # 15TQ

112 MARCELO STUCCHI AND STEVEN D. EMSLIE

represented by fossils from the late Pleistocene During the late Miocene/early Pliocene, the Rancho la Brea, and it is distinct in morphology Peruvian coast was characterized by a diverse from all other genera (Miller 1910, Miller and assemblage of marine mammals and birds (de Howard 1938, Howard 1974, Emslie 1988b). Muizon and DeVries 1985, Cheneval 1993, One other fossil genus, Antillovultur Arredondo Stucchi 2003). This fauna is associated with re- 1976 from the late Pleistocene of Cuba, is now mains of Perugyps and we believe that this con- considered to be congeneric with Gymnogyps dor fed on coastal carcasses of marine mammals, (Emslie 1988b, SuaÂrez 2000, SuaÂrez and Emslie and perhaps took live chicks of seabirds, similar 2003). An indeterminate genus and species of to the Andean Condor's habits along the north large condor from the early Pliocene Lee Creek coast of Peru today (Pennycuick and Scholer Mine, North Carolina, is known by a humeral 1984, Wallace and Temple 1987). As no other end of a coracoid, distal tibiotarsus, and pedal scavenging species are known among the fauna phalanx (Olson and Rasmussen 2001). This ma- of the Pisco Formation, it is reasonable to as- terial is too fragmentary to provide diagnostic sume that Perugyps ®lled this niche. We expect characters in comparison to other condors, that additional material of this condor will be though the distal tibiotarsus (USNM 430883) recovered from vertebrate-bearing units of the has a relatively shallower intercondylar fossa Pisco Formation. compared with Perugyps (MUSM 260). Perugyps diazi indicates that condors were ACKNOWLEDGMENTS present in South America by the late Miocene, We thank Messrs. Ana MarõÂa and Santiago E. Stucchi, at least 2.0 Ma earlier than suggested by Emslie Mario Urbina, Rodolfo Salas and Thomas DeVries for their important contributions to this research. S. Olson (1988b), who proposed that condors may have and J. Dean allowed access to comparative collections arrived by the middle Pliocene and near the be- at the U. S. National Museum, Washington, DC. David ginning of the Great American Biotic Inter- Willard and John Bates allowed access to comparative change. In addition, Tonni and Noriega (1998) collections at the Field Museum Natural of History, report a fossil of the living Andean Condor from Chicago. We also thank Paul Velazco, Gonzalo CaÂr- denas, Judith Figueroa, Michael McGowan and Jose the early Chapadmalalan (4 Ma), RõÂo QuequeÂn Tello for their help in Chicago, and S. Olson, H. James Salado (Buenos Aires), Argentina. Thus, it is and T. DeVries for their help in Washington DC. This now apparent that condors reached South Amer- paper was improved by comments from K. Campbell ica by the late Miocene to early Pliocene. and S. Olson. If condors did evolve in North America, then LITERATURE CITED they were able to reach South America early in ALVARENGA,H.M.F.,AND S. L. OLSON. 2004. A new the evolution of this group. We hypothesize that genus of tiny condor from the Pleistocene of Bra- an ancestral condor was able to expand south- zil (Aves: Vulturidae). Proceedings of the Biolog- ward following coastal corridors on the western ical Society of Washington 117:1±9. side of the Andes. The California Condor can BAUMEL,J.J.[ED.]. 1993. Handbook of avian anatomy: range hundreds of kilometers in a single day, at nomina anatomica avium. 2nd ed. Publications of the Nuttall Ornithological Club 23. ground speeds up to 70±95 kph (Snyder and BECKER, J. J. 1986. A new vulture (Vulturidae: Snyder 2000), while the Andean Condor can Pliogyps) from the late Miocene of Florida. Pro- reach speeds averaging 65 kph (McGahan 1971) ceedings of the Biological Society of Washington and can ¯y 200 km across deserts from the An- 99:502±508. BRODKORB, P. 1967. Catalogue of fossil birds: Part 3 dean foothills to the coast to forage in a single (Ralliformes, Ichthyornithiformes, Charadrii- day. Moreover, Pennycuick and Scholer (1984) formes). Bulletin of the Florida State Museum Bi- found that the latter species is almost entirely ological Sciences 11:99±220. dependent on slope uplifts to sustain prolonged CAMPBELL, K. E., JR. 1979. The non-passerine Pleis- soaring ¯ight. We believe these ¯ight capabili- tocene avifauna of the Talara tar seeps, north- western Peru. Royal Ontario Museum, Life Sci- ties of condors, along with coastal winds and ences Contribution 118. updrafts common along the western slope of the CHENEVAL, J. 1993. LAvifaune Mio-PlioceÁnedela Andes, may have allowed an ancestral condor to Formation Pisco (PeÂrou). Etude PreÂliminaire. cross the marine barrier that existed between Document Laboratoire GeÂologie Lyon NЊ 125:85± North and South America (the submerged Pa- 95. DE MUIZON, C. 1981. Les Vertebres fossiles de la For- namanian land bridge) in the late Miocene and mation Pisco (PeÂrou). PremieÂre Partie: deux nou- early Pliocene. veaux Monachinae (Phocidae: Mammalia) du Pli- CONDOR Monday Dec 13 2004 03:19 PM cond 107_115 Mp_113 Allen Press • DTPro System GALLEY File # 15TQ

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ocene Inferior de Sud-Sacaco, p. 1±160. In Re- MORENO,F.P.,AND A. MERCERAT. 1891. Catalogo de cherches sur les grandes civilisations, MeÂmoire 6. los pajaros fosiles de la Republica Argentina. ADPF, Ed., Paris. Anales del Museo de la Plata, Paleontologia Ar- DE MUIZON, C., AND T. J. DEVRIES. 1985. Geology and gentina 1:7±71. paleontology of the Late Cenozoic marine depos- OLSON, S. 1985. The fossil record of birds, p. 79±256. its in the Sacaco area (Peru). Geolische Runds- In D. S. Farner, J. R. King, and K. C. Parkes chau 74:547±563. [EDS.], Avian biolog. Vol. VIII. Academic Press, EMSLIE, S. D. 1988a. An early condor-like vulture from Orlando, FL. North America. Auk 105:529±535. OLSON, S., AND P. R ASMUSSEN. 2001. Miocene and Pli- EMSLIE, S. D. 1988b. The fossil history and phyloge- ocene birds from the Lee Creek Mine, North Car- netic relationships of condors (Ciconiiformes: olina. Smithsonian Contributions to Paleobiology Vulturidae) in the New World. Journal of Verte- 90:233±365. brate Paleontology 8:212±228. PENNYCUICK, C. J., AND K. D. SCHOLEY. 1984. Flight EMSLIE, S. D. 1998. Avian community, climate, and behavior of Andean Condor Vultur gryphus and sea-level changes in the Plio-Pleistocene of the Turkey Vultures Cathartes aura around the Para- Florida Peninsula. Ornithological Monographs 50. cas Peninsula, Peru. Ibis 126:253±256. NYDER AND NYDER HERTEL, F. 1992. Morphological diversity of past and S , N., H. S . 2000. The California present New World vultures, p. 413±418. In K. Condor. Academic Press, San Diego, CA. STUCCHI, M. 2003. Los Piqueros (Aves: Sulidae) de la Campbell [ED.], Papers in avian paleontology hon- oring Pierce Brodkorb. Los Angeles County Mu- FormacioÂn Pisco, PeruÂ. BoletõÂn de la Sociedad seum Science Series, No. 36. GeoloÂgica del Peru 95:75±91. SUA REZ, W. 2000. ContribucioÂn al conocimiento del HOWARD, H. 1929. The avifauna of Emeryville shell- estatus geneÂrico del coÂndor extinto (Ciconiifor- mound. University of California Publications in mes: Vulturidae) del Cuaternario Cubano. Orni- Zoology 32:301±394. tologõÂa Neotropical 11:109±122. HOWARD, H. 1974. Postcranial elements of the extinct SUA REZ,W.,AND S. D. EMSLIE. 2003. New fossil ma- condor Breagyps clarki (Miller). Natural History terial with a redescription of the extinct condor Museum of Los Angeles County, Contributions in Gymnogyps varonai (Arredondo, 1971) from the Science 256. Quaternary of Cuba (Aves: Vulturidae). Proceed- MAROCCO, R., AND C. DE MUIZON. 1988. Los vertebra- ings of the Biological Society of Washington 116: dos del NeoÂgeno de la Costa Sur del PeruÂ: am- 29±37. biente sedimentario y condiciones de fosilizacioÂn. TAMBUSSI,C.P.,AND J. I. NORIEGA. 1999. The fossil Bulletin Institut Francais Etudes Andines XVII: record of condors (Ciconiiformes: Vulturidae) in 105±117. Argentina. Smithsonian Contributions to Paleobi- MCGAHAN, J. 1971. The condor, soaring spirit of the ology 89:177±184. Andes. National Geographic Magazine 139:684± TONNI, E., AND J. NORIEGA. 1998. Los coÂndores (Ci- 709. coniiformes, Vulturidae) de la RegioÂn Pampeana MILLER, L. 1910. The condor-like vultures of Rancho de la Argentina durante el Cenozoico tardõÂo: dis- la Brea. University of California Bulletin of the tribucioÂn, interacciones y extinciones. Ameghinia- Department of Geology 6:1±19. na 35:141±150. MILLER, L., AND H. HOWARD. 1938. The status of the TORDOFF, H. B. 1959. A condor from the upper Plio- extinct condor-like birds of the Rancho la Brea cene of Kansas. Condor 61:338±343. Pleistocene. Publications of the University of Cal- WALLACE,M.P.,AND S. A. TEMPLE. 1987. Competitive ifornia at Los Angeles in Biological Sciences 1: interactions within and between species in a guild 169±176. of avian scavengers. Auk 104:290±295.