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The birds of CIEM, Tinigua National Park, Colombia: an overview of 13 years of ornithological research C. Daniel Cadena, Mauricio Alvarez, Juan L. Parra, Iván Jiménez, Carlos A. Mejia, Marcela Santamaría, Ana M. Franco, Carlos A. Botero, Germán D. Mejia, Ana M. Umaña, Alejandro Calixto, Juanita Aldana and Gustavo A. Londoño Cotinga 13 (2000): 46– 54

Presentamos un panorama general de la investigación ornitológica desarrollada durante trece años en el Centro de Investigaciones Ecológicas Macarena (CIEM), una estación biológica de la Universidad de los Andes (Bogotá, Colombia) y Miyagi University of Education (Japón). El CIEM está ubicado en un bosque húmedo tropical de tierras bajas al norte de la Amazonia colombiana (02°40'N 74°10'W), un lugar de gran importancia biogeográfica por la confluencia de elementos andinos, amazónicos, de los llanos y del escudo guyanés. Hasta la fecha, hemos registrado un total de 441 especies de aves incluyendo 18 migratorias Neárticas y cinco migratorias australes. Los registros reportados amplían el rango de distribución conocido de 114 de las especies incluyendo el primer registro para Colombia de Cacicus sclateri y el hallazgo de especies poco conocidas en el país como Deconychura stictolaema, Myrmotherula sunensis, Drymophila devillei, Rhegmatorhina melanosticta, Hylophylax punctulata y Turdus lawrencii. Además de inventarios de aves, varios proyectos en ornitología se han desarrollado en el CIEM desde su creación en 1986. Entre éstos se incluyen estudios de campo sobre ecología y comportamiento de paujiles Mitu salvini y Crax alector, densidades poblacionales de aves de gran tamaño, ecología de aves migratorias Nearticas, biología reproductiva de aves playeras, comportamiento de aves frugívoras, asociaciones de insectívoros en bandadas mixtas y selección de hábitat a nivel de la comunidad. El CIEM es un sitio bien establecido, con instalaciones adecuadas y buena información básica sobre su avifauna, lo cual lo convierte en un lugar ideal para cualquier persona interesada en desarrollar investigaciones de campo.

Introduction

influences. These factors point to the conservation priority for the area, as well as the need for biological research to provide basic information concerning its co n stitu en t com m unities and ecological processes. Sadly, political instability in the region has largely deterred such studies. The Centro de Investigaciones Ecológicas Macarena biological station (CIEM) is located on the west bank of the whitewater río Duda (02°40'N 74°10'W, 350-400 m asl.), which forms the boundary between Tinigua and La Macarena National Parks (Fig. 1). The río Duda meets the río Guayabero, a tributary of the río Orinoco, c.13 km downstream. The temperature at CIEM is relatively constant throughout the year (25°C) but precipitation is strongly seasonal with a dry season from December to February–March, and rainy season during the remainder of the year, with c.2600 mm mean annual rainfall17. CIEM encompasses lowland humid forest with several distinctive h a b itats4,13,27, 30. Low-lying seasonally flooded forests are characterised by a discontinuous canopy dom inated by Guarea guidonia, Laetia corymbulosa, Ficus spp., Inga spp. and Cecropia spp. trees, and a dense understorey with abundant Heliconia spp. herbs. Tessaria integrifolia, Gynérium sagittatum, young Cecropia spp. and various gram inoids dom inate early successional riparian forests on recently formed sandbanks along river m argins. Rolling hills dissected by streams support terra firme forest with

Tinigua National Park is located in the northern Colombian Amazon, betw een th e C ordillera Oriental of the Andes and Sierra de La Macarena (Fig. 1). Tinigua adjoins two other Colombian national parks: Los Picachos and La Macarena, which protect a 12 763 km2 altitudinal continuum that represents one of the key conservation areas in Colombia. The three protected areas encompass all life-zones, ranging from páramo in the Cordillera Oriental to lowland humid forests of the Amazonian plains surrounding the río Guaviare11. Although the region belongs to Amazonia sensu lato, it has a particularly interesting geological history and geographic position. Despite the close proximity of the Sierra de La Macarena to the E a ste rn Andes, th e ir origins are completely independent. The sierra, as with most of northern Amazonia, is geologically part of the Guyana Shield, characterised by poor soils, whereas the Andes are considerably younger with rich volcanic soils9,19. In addition, the geographic position of Tinigua and La Macarena National Parks is strongly influenced by the convergence of Amazonia and O rinoquia (Llanos) ecosystems. These major influences are all recognised centres of biodiversity and significant sources of species composition, posing problems w hen a tte m p tin g to classify th e area biogeographically and zoogeographically11. The region’s great biodiversity is probably largely a consequence of the interaction and overlap of these 46

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a continuous 20–25-m-high canopy with emergent trees to 30 m on ridges. Lower, eroded hill slopes and small valleys formed by creeks are covered by open degraded forest, dominated by vines, lianas, bamboo and thick understorey. One 426 ha forest plot surrounding one of three CIEM base camps encompasses primary forest (53% of the total area), open degraded forest (34%), seasonally flooded forest (11%), and riparian and secondary forests with overt human intervention (2%)28. CIEM was created in 1986 th ro u g h an agreem ent between Universidad de los Andes, Bogotá, Colombia and Miyagi U niversity of

Education, Japan. The Colombian and Japanese coordinators since that time have been CAM and K. Izawa. Since the establishm ent of Tinigua National Park in 1989, Universidad de los Andes has signed special agreem ents w ith local environm ental au th o ritie s in order to work cooperatively in research and education programmes. Since 1988, CIEM has published papers on different projects carried out at the study site in its own journal: Field studies of New World monkeys, La Macarena. The journal, which is published annually or biannually, recently changed its name to Field studies of fauna and flora, La Macarena, Colombia. Back issues can be obtained from CAM. Ornithological research at CIEM began in 1986 with non-systematic observations, mist-netting, and specimen collection conducted by CAM, IJ, MA, MS and several other students of Universidad de los Andes. Specimens have been deposited in the ornithological collections of the Instituto de Ciencias N atu rales (ICN), U niversidad N acional de Colombia, Bogotá and the Instituto Alexander von Humboldt (IAvH, formerly UNIFEM-INDERENA), Villa de Leyva. Research during the first seven years produced a prelim inary bird inventory of 373 species1. Bird censuses and mist-netting continue to be undertaken by various researchers and students. This paper presents a summary of the findings from 13 years of ornithological research together with the current inventory of 441 species (Appendix 1). Although the list is still far from com plete, w ith sam pling lacking in various m icrohabitats, such as oxbow lakes, bamboo thickets, tree-fall gaps, and forest edges on river banks, the number of species recorded at CIEM falls within the range reported for Neotropical lowland forest localities following long-term studies (e.g. Karr et al. 199016). The avifauna of CIEM includes 18 Nearctic and five austral migrants, while records of 114 species represent significant extensions of the distribution described by Hilty & Brown 198612. Others expand previously reported a ltitu d in a l ranges. The following records are particularly noteworthy because the species involved are poorly known in Colombia, with no or very few previous records. Spot-throated Woodcreeper Deconychura stictolaema A lthough suspected to be w idespread in the Colombian Amazonia, this species has only been reported from dpto. Putumayo7, two localities in dpto. Amazonas3(M. Kelsey pers. comm.), southern dpto. Vaupés5, and Chiribiquete National Park (F. G. Stiles pers. comm.). Four individuals have been captured (one collected by MA et al., ICN 32489) at CIEM in terra firme primary forest.

F ig ure 1. L o c a tio n o f T in ig u a N a tio n a l Park.

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Pseudobombax munguba in the middle of an oxbow lake. Unfortunately, no specimens were obtained due to the inaccessibility of the nest.

Rio Suno Antwren Myrmotherula sunensis The only previous records of th is species in Colombia were made in the extreme south-east on the Nariño–Putumayo border near the eastern base of the Andes12. A single specimen collected by CAM at CIEM (IAvH 7035) represents a very significant extension of its known range.

Specific projects at CIEM Aside from general bird inventories, several specific ornithological projects have been conducted at CIEM. A b rief sum m ary of some, largely unpublished, studies is presented, with further details being available from CDC. The most important project at CIEM has been the pioneer work on the ecology of curassows (Cracidae) in the wild. MS and AMF commenced the project in 1990, habituating a pair of Salvin’s Curassows Mitu salvini to the presence of human observers. This perm itted the observers (and subsequent researchers) to follow the curassow pair and their young for extended time periods at very close range (2–5 m) and make very detailed observations on their behaviour. After intensive fieldwork, valuable information was collected for this species on diet composition, foraging behaviour, spatial requirements, reproductive biology, parental care, seed dispersal and predation, and population density8,24. A complementary study focused on the importance of these birds as seed predators and dispersers6. Stevenson26 provides information on population densities of both Mitu salvini and Black Curassow Crax alector, along with other cracids occurring at CIEM: Speckled Chachalaca Ortalis g u tta ta , B lue-throated P iping-Guan Pipile cumanensis and Spix’s Guan Penelope jacquacu. His censuses also included Grey-winged Trumpeters Psophia crepitans. Since 1994, Crax alector has also been studied at CIEM, with researchers focusing on the species’ diet com position, sp atial requirements, reproductive biology and feeding of the young. Recent research on curassows at CIEM has been conducted by IJ et al. and has focused on foraging behaviour and use of space, its variation over different time periods, and its relationship to the availability of food. Future work on curassows will include a comparative study on the ecology of M. salvini and C. alector and relationships to fruit availability, and a doctoral dissertation th a t endeavours to determine how curassows select their diet. Details are available from IJ. CIEM was one of five Colombian localities selected for censuses and banding of Nearctic migrants in 1991 and 1992 by MA, IJ, K. Schuetze and G. W. Kaiser of Universidad de los Andes and the Programa Latino America of the Canadian Wildlife Service (CWS). Presently, AC in conjunction w ith Dr K. A. Hobson (CWS—U niversity of Saskatchewan) is studying the migration biology of four Nearctic passerines (Catharus ustulatus, Dendroica striata, Setophaga ruticilla, and Seiurus noveboracensis) at CIEM. This is part of a major project th a t plans to establish the connection

Striated Antbird Drymophila devillei Known in Colombia only from two specimens from uncertain localities in dpto. Putumayo29. A pair was observed and repeatedly heard singing in a bamboo thicket in seasonally flooded forest, very close to a river bank, in June 1997. Vocalisations were recorded by IJ and deposited in the Library of Natural Sounds at Cornell University, Ithaca, NY. The species’ specific bamboo m icro h ab itat requirements18,22probably account for its apparently patchy distribution. Hairy-crested Antbird Rhegmatorhina melanosticta This ‘professional’ ant follower was known in Colombia only from two lo calities in dpto. Putumayo7,32 and recently from Chiribiquete and La Paya National Parks (F. G. Stiles pers. comm.), and Serranía de los Churumbelos, dpto. Cauca (P. Salaman pers. comm.). Several sight records have been made at CIEM, both in terra firme primary and open degraded forests. Dot-backed Antbird Hylophylax punctulata Known from one Colombian specimen (FMNH, Chicago) from Río Guapaya, Sierra de La Macarena, collected by K. von Sneidern in 1957 and previously misidentified as H. naevia23 (D. E. Willard pers. comm.). This species is patchily distributed and its distribution poorly known22. At CIEM, a male was trapped and photographed in 1997 by JA and CAB in terra firme forest very close to seasonally flooded forest, which appears to be its usual habitat24. Lawrence's Thrush Turdus lawrencii Known in Colombia only from specimens obtained in L eticia20 and P utum ayo7 and sightings in Vaupés5,12and Amazonas3 (M. Kelsey pers. comm.). Turdus lawrencii is the least known of the Turdus that breed in Amazonia and is patchily distributed, often being absent from apparently appropriate habitat10. Three observed and one collected (ICN 32026) at CIEM by MA et al. in 1995 in terra firme forest represent the northernmost records for the species. Ecuadorian Cacique Cacicus sclateri The occurrence of this poorly known species was considered possible but unconfirmed in southern Colombia14. A pair observed by CAB at CIEM in April 1996 nested on an isolated aquatic ceiba 48

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between breeding and wintering sites for different populations of these and other Nearctic migrants. Further analyses including the use of genetic tags and stable isotopes will provide data on the origin of the birds wintering at CIEM, and long-term monitoring will aid in determining the importance of this region for the conservation of migrant birds. Various species nest on the extensive sand shores of the río Duda during the dry season. D etailed studies have been performed on the breeding biology of some of these: Pied Lapwing Vanellus cayanus by IJ and CAM15, and Large-billed Phaetusa simplex and Yellow-billed Terns Sterna superciliaris by J. C. Verhelst31. Two studies involving frugivores have been completed a t CIEM: the n a tu ra l history and reproductive biology of Purple-throated Fruitcrow Querula purpurata by J. D. Amaya2, and on the assem blages of diu rn al fru it-eatin g arboreal vertebrates at fig trees by IJ, MA and CAM. CAB and JA, co n cen tratin g on species composition and role within flocks, their daily activities, habitat use and home range size have studied mixed insectivorous species foraging flocks in the understorey. In progress, since 1995, is a long­ term project on habitat selection of forest birds, directed by MA, in which several students of U niversidad de los Andes have participated. Systematic censuses and mist-netting have been carried out in riparian, seasonally flooded, and terra firme forests. In addition, more than 130 hours of bird vocalisations have been recorded by MA and IJ. Once processed, at the Sound Bank of the Instituto Alexander von Humboldt, it is likely that several new species will be added to the area’s avifauna. As demonstrated, CIEM is a well-established site with facilities and good baseline bird data, ideal for anyone wishing to conduct research. Those wishing to visit CIEM should contact Carlos A. Mejia to arrange permission, accommodation and receive specific access details. Two or three days are required to reach CIEM from Bogotá. The trip has three main stages: Bogotá–Villavicencio (fourhour bus ride or 30-minute flight), Villavicencio–La Macarena (one- hour flight), and La Macarena– CIEM (c.8 hours upriver, depending on weather conditions).

A ckn ow led gem ents This paper was improved with comments and information provided by Paul Salaman, Gary W. Kaiser, Martin Kelsey, Pablo R. Stevenson, F. Gary Stiles, David E. Willard and Nicolás Reyes. Those involved in curassow projects aside from the authors and those cited above include: Ximena Bernal, Jaqueline Carmona, M aría Clara Castellanos, Marisol Escaño, Jimena Forero, Jorge Molina, Juan Carlos Sánchez, Juan Gabriel Soler and Paola 49

Zuluaga. Students participating in field courses at CIEM provided much useful information. For their support in the field we thank the late Gerardo S an ab ria, Alvaro S an ab ria and Ramiro Montealegre. The Japan-Colom bia agreem ent allowed us to work at CIEM.

R eferences 1. A lvarez, M., Jim énez, L, Mejia, C. A. & Santamaría, M. (1993) Bird species at the CIEM, Tinigua National Park: a partial list. Field studies o f New World Monkeys, La Macarena, Colombia 8: 33–40. 2. Amaya, J. D. (1997) Biología reproductiva y aspectos de historia n atu ral de Querula purpurata (Aves: Cotingidae) en un bosque del Parque Nacional N atural Tinigua (Meta, Colombia). Universidad de los Andes, Bogotá: BSc. thesis. 3. Andrade, G. I. & Rubio-Torgler, H. (1994) Sustainable use of the tropical rain forest: evidence from the avifauna in a shiftingcultivation habitat mosaic in the Colombian Amazon. Conserv. Biol. 8: 545–554. 4. Anonymous (1997) The study site: a brief description. Field studies of fauna and flora, La Macarena, Colombia 10: 5. 5. Bennett-Defler, S. (1994) Las aves de la estación Capará: una lista prelim inar de especies. Trianea (Act. Cient. Tecn. INDERENA) 5: 379– 400. 6. De los Ríos, P. T. (1997) Dispersión y depredación de semillas por paujiles Mitu salvini (Aves: Cracidae) en la Amazonia colombiana. Universidad de los Andes, Bogotá: BSc. thesis. 7. Fitzpatrick, J. W. & Willard, D. E. (1982) Twentyone bird species new or little known from the Republic of Colombia. Bull. Brit. Orn. Club 102: 153–158. 8. Franco, A. M. & Santamaría, M. (1997) Biología reproductiva del paujil Mitu salvini. In: Strahl, S. D., Beaujon, S., Brooks, D. M., Begazo, A., Sedaghatkish, G. & Olmos, F. (eds.) The Cracidae: their biology and conservation. Blaine, WA: Hancock House Publishers. 9. Gentry, A. H. (1990) La región Amazónica. In: Villegas, J. B. & Uribe, C. (eds.) Selva Húmeda de Colombia. Bogotá: Villegas Editores. 10. Hardy, J. W. & Parker, T. A. (1997) The nature and probable function of vocal copying in Lawrence’s Thrush, Turdus lawrencii. Orn, Monogr. 48: 307–320. 11. Hernández, J. L, Sánchez, H., Rodríguez, J. V, Castaño, C., Cano, M. & Mejia, I. (1998) P arque N acional N a tu ra l S ierra de La Macarena. In: Castaño-Uribe, C. & Cano, M. (eds.) El sistema de Parques Nacionales Naturales de Colombia, Bogotá: Ed. Nomos.

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12. Hilty, W. L. & Brown, S. L. (1986) A guide to the birds o f Colombia. Princeton: Princeton University Press. 13. Hirabuki, Y. (1990) Vegetation and landform structure in the study area of La Macarena: a physiognomic investigation. Field studies of New World monkeys, La Macarena, Colombia 3: 35–48. 14. Jaramillo, A. & Burke, P. (1999) New World blackbirds: the icterids. Princeton: Princeton University Press. 15. Jiménez, I. & Mejia, C. A. (1998) Breeding biology of H oploxypterus cayanus (Charadriidae) at the río Duda. Field studies of fauna and flora, La Macarena, Colombia 12: 3–7. 16. Karr, J. R., Robinson, S. K., Blake, J. G. & B ierreg aard , R. O. (1990) B irds of four Neotropical forests. In: Gentry, A. H. (ed.) Four Neotropical rainforests. New Haven: Yale University Press. 17. Kimura, K., Nishimura, A., Izawa, K. & Mejia, C. A. (1994) Annual changes of rainfall and temperature in the tropical seasonal forest at La Macarena Field Station, Colombia. Field studies o f New World monkeys, La Macarena, Colombia 9: 1–3. 18. Kratter, A. W. (1997) Bamboo specialization in Amazonian birds. Biotropica 29: 100– 110. 19. Mejia, C. A. (1995) Fauna de La Serranía de La Macarena. Bogotá: Amazonas Editores & Ediciones Uniandes. 20. Morales, J. E. (1979) Prim er registro para Colombia de Turdus lawrencii Coues (Aves: Turdidae). Lozanía (Acta Zoológica Colombiana) 29: 2–4. 21. Parker, T. A., Stotz, D. F. & Fitzpatrick, J. W. (1996) Ecological and distributional databases. In: Stotz, D. F , Fitzpatrick, J. W., Parker, T. A. & Moskovits, D. K. Neotropical birds: ecology and conservation. Chicago: U niversity of Chicago Press. 22. Parker, T. A., Stotz, D. F. & Fitzpatrick, J. W. (1997) Notes on avian bamboo specialists in southwestern Amazonian Brazil. Orn. Monogr. 48: 543–547. 23. Ridgely, R. S. & Tudor, G. (1994) The birds of South America, 2. Austin: University of Texas Press. 24. Santamaría, M. & Franco, A. M. (1997) Hábitos alimenticios del paujil M itu salvini en el Parque Nacional Natural Tinigua, Colombia. In: Strahl, S. D., Beaujon, S., Brooks, D. M., Begazo, A., Sedaghatkish, G. & Olmos, F. (eds.) The Cracidae: their biology and conservation. Blaine, WA: Hancock House Publishers. 25. Sibley, C. G. (1996) Birds of the world. Version 2.0. Cincinnati: Thayer Birding Software.

26. Stevenson, P. R. (1996) Censos diurnos de mamíferos y algunas aves de gran tamaño en el P arque N acional T inigua, Colombia. Universitas Scientiarum 3: 67–81. 27. Stevenson, P. R., Quiñones, M. J. & Ahumada, J. A. (1994) Ecological strategies of woolly monkeys (Lagothrix lagotricha) at Tinigua National Park, Colombia. Amer. J. Primatology 3 2 : 123–140. 28. Stevenson, P. R., Castellanos, M. C. & Medina, A. P. (1999) Elementos arbóreos de los bosques de un plano inundable en el Parque Nacional Natural Tinigua, Colombia. Caldasia 21: 38– 49. 29. Stotz, D. F.(1990) First specimens ofDrymophila devillei from Colombia. Bull. Brit. Orn. Club 110: 37–38. 30. Takehara, A. & Hirabuki, Y. (1991) Population structure of Tessaria integrifolia (Asteraceae) on newly deposited riverine soils. Field studies o f New World monkeys, La Macarena, Colombia 5: 25-29. 31. Verhelst, J. C. (1996) Biología de anidación de Phaetusa sim plex y Sterna superciliaris (Laridae) en las playas del río Duda, Parque N acional N a tu ra l Tinigua, Colombia. Universidad de los Andes, Bogotá: BSc. thesis. 32. Willis, E. O. (1969) On the behavior of five species of R hegm atorhina, ant-following antbirds of the Amazon basin. Wilson Bull. 81: 363–395. C. Daniel Cadena (1) D epartam ento de Ciencias Biológicas, Universidad de los Andes, Apartado Aéreo 110229, Bogotá, Colombia. E-mail: [email protected]. Mauricio Alvarez (2) Instituto Alexander von Humboldt, Claustro de San Agustín, Villa de Leyva, Boyacá, Colombia, Email: [email protected]. Juan L. Parra 1 E-mail: [email protected]. Iván Jiménez Department of Biology, University of Missouri–St. Louis, 8001 Natural Bridge Road, St. Louis MO 63121-4499, USA. E-mail: [email protected]. Carlos A. Mejía 1 E-mail: [email protected]. Marcela Santamaría E-mail: [email protected]. Ana M. Franco 2 E-mail: [email protected]. 50

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Carlos A. Botero 1 E-mail: [email protected].

M uscovy D uck Cairina moschata T u rk ey V ulture Cathartes aura G re a te r Y ellow -headed V ulture Cathartes melambrotus Black V ulture Coragyps atratus King V ulture Sarcoramphus papa O sp re y Pandion haliaetus (N) Sw allow -tailed Kite Elanoides forficatus Pearl Kite Gampsonyx swainsonii Snail Kite Rostrhamus sociabilis * D o u b le -to o th e d Kite Harpagus bidentatus P lum beous Kite Ictinia plumbea C ra n e H awk Geranospiza caerulescens Tiny H awk Accipiter superciliosus W h ite H aw k Leucopternis albicollis G re a t Black Haw k Buteogallus urubitinga Savanna H aw k Buteogallus meridionalis * B lack-collared H aw k Busarellus nigricollis * G rey-lined H awk Buteo nitidus * R oadside H aw k Buteo magnirostris Z o n e-tailed H aw k Buteo albonotatus C re s te d Eagle Morphnus guianensis * O rn a te H aw k-Eagle Spizaetus ornatus Black C arac a ra Daptrius ater R e d -th ro a te d C arac a ra Daptrius americanus Y ellow -headed C arac a ra Milvago chimachima * Laughing Falcon Herpetotheres cachinnans Lined F o rest-Falcon Micrastur gilvicollis C o llared F o re s t-Falcon Micrastur semitorquatus * Bat Falcon Falco rufigularis O ra n g e -b re a ste d Falcon Falco deiroleucus Speckled C hachalaca Ortalis guttata Spix’s G uan Penelope jacquacu B lu e-th ro ated Piping-Guan Pipile cumanensis Salvin’s C u ra sso w Mitu salvini * Black C u ra sso w Crax alector * M arbled W o o d - Quail Odontophorus gujanensis Limpkin Aramus guarauna * G rey-w inged T r u m p e te r Psophia crepitans Purple G allinule Porphyrio martinicus A zure G allinule Porphyrio flavirostris G rey -n eck ed W o o d - Rail Aramides cajonea S u n b ittern Eurypyga helias W a ttle d Jacana Jacana jacana S o u th ern Lapwing Vanellus chilensis Pied Lapwing Vanellus cayanus C o llared Plover Charadrius collaris Solitary S andpiper Tringa solitaria (N) G re a te r Yellowlegs Tringa melanoleuca (N) S p o tted Sandpiper Tringa macularia (N) D o u b le-strip ed T hick-knee Burhinus bistriatus Large-billed T e rn Phaetusa simplex Yellow-billed T e rn Sterna superciliaris Black Skim m er Rynchops niger Scaled Pigeon Columba speciose Pale-vented Pigeon Columba cayennensis Ruddy Pigeon Columba subvinacea Plum beous Pigeon Columba plumbea Blue G ro u n d Dove Claravis pretiosa G rey -fro n te d dove Leptotila rufaxilla Ruddy Q u ail-Dove Geotrygon montana S carlet M acaw Ara macao C h e s tn u t-fro n te d M acaw Ara severa R ed-bellied M acaw Ara manilata W h ite -e y ed P ara k e et Aratinga leucophthalmus M aroon-tailed P a ra k e et Pyrrhura melanura C obalt-w inged P a ra k e et Brotogeris cyanoptera B lack-headed P a rro t Pionites melanocephala O ra n g e -c h ee k e d P a rr o t Pionopsitta barrabandi * B lue-headed P a rr o t Pionus menstruus Y ello w -cro w n ed P a rr o t Am azona ochrocephala

Germán D. Mejía 1 E-mail: [email protected]. Ana M. U m aña E-mail: [email protected]. Alejandro Calixto Centro de Investigaciones Ecológicas Macarena, Apartado Aéreo 270175, Bogotá, Colombia. E-mail: [email protected]. Juanita Aldana 1 E-mail: j [email protected]. Gustavo A. Londoño 1 E-mail: [email protected].

A p p e n d ix 1. Bird species recorded at CIEM, Tinigua

National Park, Colombia. English and scientific names fo llo w Sibley ( 1996)25. Sequence follow s H ilty & Brown ( 1986)12. (*) = Range extension (N ) = N earctic migrant (A) = Austral migrant Subspecies are mentioned only when relevant to migratory status. G re a t T inam ou Tinamus major W h ite -th ro a te d T inam ou Tinamus guttatus C in e re o u s T inam ou Crypturellus cinereus Little T inam ou Crypturellus soui U nd ulated T inam ou Crypturellus undulatus V ariegated T inam ou Crypturellus variegatus * N e o tro p ic C o rm o ra n t Phalacrocorax brasilianus Anhinga Anhinga anhinga G re a t Blue H ero n Ardea herodias * C o co i H e ro n Ardea cocoi G re a t E gret Ardea alba Snow y Egret Egretta thula Little Blue H ero n Egretta caerulea G re e n H ero n Butorides virescens (N) S triated H e ro n Butorides striatus Agami H e ro n Agamia agami C attle E gret Bubulcus ibis W histling H ero n Syrigma sibilatrix * C ap ped H ero n Pilherodius pileatus B lack-crow ned N ight-He ro n Nycticorax nycticorax R u fescent T ig er-h e ro n Tigrisoma lineatum B oat-billed H ero n Cochlearius cochlearia W o o d S to rk Mycteria americana G re e n Ibis Mesembrinibis cayennensis * W h isp erin g Ibis Phimosus infuscatus S carlet Ibis Eudocimus ruber R o seate Spoonbill Ajaia ajaja H o rn e d S cre a m e r Anhima cornuta W h ite-faced W histling-Duck Dendrocygna viduata Black-bellied W histling-Duck Dendrocygna autumnalis * O rin o c o G o o se Neochen jubata B lue-w inged Teal Anas discors (N)

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Cotinga 13

The birds o f CIEM, Tinigua N ational Park, C olom bia

Mealy P a rro t Am azona farinosa Black-billed C u ck o o Coccyzus erythropthalmus (N ) Yellow-billed C u ck o o Coccyzus americanus (N) D ark-billed C u ck o o Coccyzus melacoryphus (A) Squirrel C u ck o o Piaya cayana Black-bellied C u ck o o Piaya melanogaster Little C u ck o o Piaya minuta G r e a te r Ani Crotophaga major S m ooth-billed Ani Crotophaga ani S triped C u ck o o Tapera naevia H oatzin Opisthocomus hoazin T ropical S cre e ch -Owl Otus choliba Taw ny-bellied S cre e ch -Owl Otus watsonii * Spectacled O w l Pulsatrix perspicillata F erruginous Pygmy-owl Glaucidium brasilianum M ottled O w l Strix virgata O ilbird Steatornis caripensis * G re a t P o to o Nyctibius grandis C o m m o n P o to o Nyctibius griseus S an d -co lo red N ighthaw k Chordeiles rupestris N acu n d a N ighthaw k Podager nacunda Pau raq u e Nyctidromus albicollis L adder-tailed N ightjar Hydropsalis climacocerca * W h ite -c o lla re d Swift Streptoprocne zonaris S h o rt-tailed Swift Chaetura brachyura R u fo u s-b reasted H e rm it Glaucis hirsuta Pale-tailed B arb th ro a t Threnetes leucurus Long-tailed H e rm it Phaethornis superciliosus W h ite -b e a rd e d H e rm it Phaethornis hispidus S o o ty -cap ped H e rm it Phaethornis augusti * R eddish H e rm it Phaethornis ruber * G rey -ch in ned H e rm it Phaethornis griseogularis Little H e rm it Phaethornis longuemareus G re y -b re a ste d Sabrew ing Campylopterus largipennis W h ite -n e ck e d Jacobin Florisuga mellivora B lack -th roated Mango Anthracothorax nigricollis Festive C o q u e tte Lophornis chalybeus * Black-bellied T h o rn tail Popelairia langsdorffi * B lue-tailed Em erald Chlorostilbon mellisugus Fork-tailed W o o d n y m p h Thalurania furcata W h ite -c h in n ed S apphire Hylocharis cyanus * G o ld en -tailed Sapphire Chrysuronia oenone V ersico lo red Em erald Amazilia versicolor * G litte rin g -th ro a te d Em erald Amazilia fimbriata B lack-eared Fairy Heliothryx aurita Long-billed S ta rth ro a t Heliomaster longirostris * Black-tailed T ro g o n Trogon melanurus W h ite -ta ile d T ro g o n Trogon viridis B lack -th ro ated T ro g o n Trogon rufus * B lu e-cro w ned T ro g o n Trogon curucui * V iolaceou s T ro g o n Trogon violaceus Ringed Kingfisher Megaceryle torquata A m azon Kingfisher Chloroceryle am azona G re e n Kingfisher Chloroceryle americana G re e n -a n d -ru fo u s Kingfisher Chloroceryle inda A m erican Pygmy Kingfisher Chloroceryle aenea R ufous-capped M o tm o t Baryphthengus ruficapillus B lu e-cro w ned M o tm o t M om otus m om ota B row n Jacam ar Brachygalba lugubris * Y ellow -billed Jacam ar Galbula albirostris W h ite -c h in n ed Jacam ar Galbula tombacea B ronzy Jacam ar Galbula leucogastra * G re a t Jacam ar Jacamerops aureus W h ite -n e ck e d Puffbird Notharchus macrorhynchus * Pied Puffbird Notharchus tectus * C h estn u t-ca p p e d Puffbird Bucco macrodactylus C o llared Puffbird Bucco capensis W h ite -c h e s te d Puffbird Malacoptila fusca L an ceolated M onklet Micromonacha lanceolata

B row n N u n le t Nonnula brunnea B lack-fronted N u n b ird Monasa nigrifrons W h ite -fro n te d N u n b ird M onasa morphoeus Y ellow -billed N u n b ird Monasa flavirostris Sw allow -w inged Puffbird Chelidoptera tenebrosa S carle t-c ro w n e d B arb et Capito aurovirens * B lack -sp o tted B arb et Capito niger L e m o n -th ro a ted B arb et Eubucco richardsoni C h e s tn u t-e a re d A ra cari Pteroglossus castanotis M any-banded A racari Pteroglossus pluricinctus L e tte re d A racari Pteroglossus inscriptus Ivory-billed A racari Pteroglossus azara G o ld en -co llared T o u c a n e t Selenidera reinwardtii Y ellow -ridged T o u can Ramphastos culminatus Red-billed T o u can Ramphastos tucanus R ufou s-b reasted Piculet Picumnus rufiventris * Lafresnaye’s Piculet Picumnus lafresnayi * S p o t-b re aste d W o o d p e c k e r Colaptes punctigula Y ello w -th ro ated W o o d p e c k e r Piculus flavigula G o ld en -g reen W o o d p e c k e r Piculus chrysochloros C h e s tn u t W o o d p e c k e r Celeus elegans Scaly-breasted W o o d p e c k e r Celeus grammicus C re a m -c o lo re d W o o d p e c k e r Celeus flavus Ringed W o o d p e c k e r Celeus torquatus * Lineated W o o d p e c k e r Dryocopus lineatus Y ellow -tu fted W o o d p e c k e r M elanerpes cruentatus S m oky-brow n W o o d p e c k e r Veniliornis fumigatus * Little W o o d p e c k e r Veniliornis passerinus R ed-ru m p ed W o o d p e c k e r Veniliornis kirkii * C rim s o n -c re ste d W o o d p e c k e r Campephilus melanoleucos R ed-n eck ed W o o d p e c k e r Campephilus rubricollis Plain-brow n W o o d c r e e p e r Dendrocincla fuliginosa W h ite -c h in n ed W o o d c r e e p e r Dendrocincla merula * S p o t-th ro a te d W o o d c r e e p e r Deconychura stictolaema * O livaceo u s W o o d c r e e p e r Sittasomus griseicapillus W edg e-b illed W o o d c r e e p e r Glyphorynchus spirurus Long-billed W o o d c r e e p e r Nasica longirostris * C in n a m o n -th ro a te d W o o d c r e e p e r Dendrexetastes rufigula * Strong-billed W o o d c r e e p e r Xiphocolaptes promeropirhynchus * B arred W o o d c r e e p e r Dendrocolaptes Certhia B lack-banded W o o d c r e e p e r Dendrocolaptes picumnus * Straight-billed W o o d c r e e p e r Xiphorhynchus picus Spix’s W o o d c r e e p e r Xiphorhynchus spixii B uff-throated W o o d c r e e p e r Xiphorhynchus guttatus C urve-billed Scythebill Campylorhamphus procurvoides * D usky Spinetail Synallaxis moesta P lain-crow ned Spinetail Synallaxis gujanensis Ruddy Spinetail Synallaxis rutilons C h estn u t-w in g ed H ookbill Ancistrops strigilatus S triped W o o d h a u n te r Hyloctistes subulatus C in n am o n -ru m p ed F oliage-gleaner Philydor pyrrhodes * C h estn u t-w in g ed Foliage-gleaner Philydor erythropterum * Rufous-tailed Foliage-gleaner Philydor ruficaudatus O live-backed Foliage-gleaner Automolus infuscatus Ruddy Foliage-gleaner Automolus rubiginosus * B uff-throated Foliage-gleaner Automolus ochrolaemus C h e s tn u t-c ro w n e d Foliage-gleaner Automolus rufipileatus * Plain X en o p s X enops minutus S hort-billed L eafto sser Sclerurus rufigularis Black-tailed L eafto sser Sclerurus caudacutus Fasciated A n tsh rik e Cymbilaimus lineatus G re a t A n tsh rik e Taraba major B ar-c re ste d A n tsh rik e Thamnophilus multistriatus * Lined A n tsh rik e Thamnophilus tenuepunctatus Plain-winged A n tsh rik e Thamnophilus schistaceus M o u se-co lo red A n tsh rik e Thamnophilus murinus * E astern Slaty-An tsh rik e Thamnophilus punctatus * A m azonian A n tsh rik e Thamnophilus amazonicus D u sk y -th ro a te d A n tsh rik e Tham nom anes ardesiacus

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Cotinga 13

The birds o f CIEM, Tinigua N ational Park, C olom bia Slaty-capped Flycatcher Leptopogon superciliaris * Sepia-capped Flycatcher Leptopogon amaurocephalus Y ellow T y ran n u let Capsiempis flaveola * Ringed A n tp ip it Corythopis torquata * S ho rt-tailed Pygmy-Ty ra n t Myiornis ecaudatus * Y ello w -b ro w ed T o d y -Fly catch er Todirostrum chrysocrotaphum C o m m o n T o d y -Fly catch er Todirostrum cinereum * S late-headed T o d y -Fly catch er Todirostrum sylvia * R usty -fro n ted T o d y -Fly catch er Todirostrum latirostre * B row nish Flycatcher Cnipodectes subbrunneus R ufous-tailed Flatbill Ramphotrigon ruficauda * L arge-headed Flatbill Ramphotrigon megacephala * O livaceous Flatbill Rhynchocyclus olivaceus G re y -c ro w n e d Flycatcher Tolmomyias poliocephalus Y ello w -b reasted Flycatcher Tolmomyias flaviventris G o ld e n -c ro w n e d Spadebill Platyrinchus coronatus * Royal Flycatcher Onychorhynchus coronatus * R uddy-tailed Flycatcher Myiobius erythrurus S ulp h u r-ru m p ed Flycatcher Myiobius barbatus D rab W a te r - Ty ra n t Ochthornis littoralis Pied W a te r - Ty ra n t Fluvicola pica * Long-tailed T y ran t Colonia colonus * B right-rum ped A ttila Attila spadiceus * C innam on A ttila Attila cinnamomeus * G reyish M o u rn e r Rhytipterna simplex C in e re o u s M o u rn e r Laniocera hypopyrra Sirystes Sirystes sibilator * S h o rt-c re ste d Flycatcher Myiarchus ferox B ro w n -c re sted Flycatcher Myiarchus tyrannulus * G re a t K iskadee Pitangus sulphuratus L esser K iskadee Philohydor lictor B oat-billed F lycatcher Megarynchus pitangua R usty-m argined Flycatcher M yiozete te s cayanensis Social Flycatcher M yiozetetes similis G rey -cap p ed Flycatcher M yio zetetes granadensis S treak ed F lycatcher Myiodynastes maculatus solitarius (A) G o ld e n -c ro w n e d Flycatcher Myiodynastes chrysocephalus * Piratic Flycatcher Legatus leucophaius V ariegated Flycatcher Empidonomus varius (A) Sulphury Flycatcher Tyrannopsis sulphurea Fork-tailed F lycatcher Tyrannus savana E astern Kingbird Tyrannus tyrannus (N) T ropical Kingbird Tyrannus melancholicus W h ite-w in g ed Swallow Tachycineta albiventer B ro w n -ch ested M artin Progne tapera fusca (A) W h ite -b a n d ed Sw allow Atticora fasciata W h ite-th ig h ed Swallow Neochelidon tibialis S o u th ern R ough-w inged Sw allow Stelgidopteryx ruficollis V iolaceous Jay Cyanocorax violaceus Black-capped D o n aco b iu s Donacobius atricapillus T hrush-like W re n Campylorhynchus turdinus C o ray a W re n Thryothorus coraya H o u se W re n Troglodytes aedon W h ite -b re a ste d W o o d -Wren Henicorhina leucosticta S o u th e rn N ightingale-Wren Microcerculus marginatus T ropical M ockingbird M imus gilvus S w ain so n ’s T h ru sh Catharus ustulatus (N) Pale-eyed T h ru sh Platycichla leucops * Black-billed T h ru sh Turdus ignobilis L a w ren ce's T h ru sh Turdus lawrencii * B are-eyed T h ru sh Turdus nudigenis * W h ite -n e ck e d T h ru sh Turdus albicollis Long-billed G n a tw re n Ramphocaenus melanurus * T ropical G n a tc a tc h e r Polioptila plumbea * R ufo u s-b ro w ed P e p p ersh rik e Cyclarhis gujanensis R ed-eyed V ireo Vireo olivaceus (N) B ro w n -h ead ed G re e n le t Hylophilus brunneiceps * T aw n y -cro w n ed G re e n le t Hylophilus ochraceiceps Shiny C o w b ird Molothrus bonariensis

C in e re o u s A n tsh rik e Tham nom anes caesius Pygmy A n tw re n Myrmotherula brachyura S treak ed A n tw re n Myrmotherula surinamensis C h e r r i e s A n tw re n Myrmotherula cherriei * P lain -th ro ated A n tw re n Myrmotherula hauxwelli S trip e -th ro a te d A n tw re n Myrmotherula haem atonota * R ufous-tailed A n tw re n Myrmotherula erythrura * O r n a te A n tw re n Myrmotherula ornata W h ite-flan k ed A n tw re n Myrmotherula axillaris Slaty A n tw re n Myrmotherula schisticolor Long-w inged A n tw re n Myrmotherula longipennis * Rio Suno A n tw re n Myrmotherula sunensis* G re y A n tw re n Myrmotherula menetriesii B anded A ntbird Dichrozona cincta W h ite-frin g ed A n tw re n Formicivora grísea * S triated A ntbird Drymophila devillei * G rey A ntbird Cercomacra cinerascens D usky A ntbird Cercomacra tyrannina Blackish A ntbird Cercomacra nigrescens * Black A ntbird Cercomacra serva * J et A n tbird Cercomacra nigricans W h ite -b ro w e d A ntbird Myrmoborus leucophrys Black-faced A ntbird Myrmoborus myotherinus W arb lin g A ntbird Hypocnemis cantator S pot-w inged A ntbird Percnostola leucostigma Silvered A ntbird Sclateria naevia * W h ite -s h o u ld e re d A ntbird M yrmeciza melanoceps B lack -throated A ntbird M yrmeciza atrothorax W h ite -p lu m e d A ntbird Pithys albifrons W h ite -c h e e k e d A ntbird Gymnopithys leucaspis H a iry -crested A ntbird Rhegmatorhina melanosticta * B lack -spotted B are-eye Phlegopsis nigromaculata * R eddish-w inged B are-eye Phlegopsis erythroptera * S p o t-b acked A ntbird Hylophylax naevia D o t-b ack ed A ntbird Hylophylax punctulata * Scale-backed A ntbird Hylophylax poecilinota S triated A n tth ru sh Cham aeza nobilis * R ufous-capped A n tth ru sh Formicarius colma Black-faced A n tth ru sh Formicarius analis T hru sh -like A n tp itta Myrmothera campanisona G o ld en -h ead ed Manakin Pipra erythrocephala B lu e-crow ned Manakin Lepidothrix coronata W ire -ta ile d M anakin Pipra filicauda W h ite -b e a rd e d M anakin Manacus manacus S triped Manakin Machaeropterus regulus G re e n M anakin Chloropipo holochlora D w arf T yrant-M anakin Tyranneutes stolzm anni T h ru sh -like Schiffornis Schiffornis turdinus * W h ite -b ro w e d P u rp letu ft Iodopleura isabellae * Scream ing Piha Lipaugus vociferans C h e s tn u t-c ro w n e d Becard Pachyramphus castaneus W h ite-w in g ed B ecard Pachyramphus polychopterus P in k -th ro ated Becard Pachyramphus minor Black-tailed T ityra Ti tyra cayana B lack-crow ned T i t yra Ti tyra inquisitor P u rp le-th ro a te d C otinga Porphyrolaema porphyrolaema * P lu m -th ro ated C otinga Cotinga maynana * Spangled C otinga Cotinga cayana B are-n ecked F ru itcro w Gymnoderus foetidus P u rp le-th ro a te d F ru itcro w Querula purpurata A m azonian U m brellabird Cephalopterus ornatus S len d er-fo o ted T y ran n u let Zimmerius gracilipes * S o u th e rn B eardless-Ty ra n n u le t Camptostoma obsoletum M o u se-co lo red T y ran n u let Phaeomyias murina * Y ello w -crow ned T y ran n u let Tyrannulus elatus F o re s t Elaenia Myiopagis gaimardii Small-billed Elaenia Elaenia parvirostris (A) M o ttle-backed Elaenia Elaenia gigas O c h re-b ellied Flycatcher Mionectes oleagineus

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Cotinga 13

The birds o f CIEM, Tinigua N ational Park, C olom bia Y ellow -bellied T an ag er Tangara xanthogastra M asked T an ag er Tangara nigrocincta T u rq u o ise T an ag er Tangara mexicana Blue-grey T an ag er Thraupis episcopus Palm T an ag er Thraupis palmarum Silver-beaked T an ag er Ramphocelus carbo M asked C rim so n T an ag er Ramphocelus nigrogularis S um m er T an ag er Piranga rubra (N) S carlet T an ag er Piranga olivacea (N) R ed -cro w n ed A n t-tan ag er Habia rubica W h ite-lin ed T an ag er Tachyphonus rufus * F ulvo u s-crested T an ag er Tachyphonus surinamus W h ite -s h o u ld e re d T an ag er Tachyphonus luctuosus Y ellow -backed T an ag er Hemithraupis flavicollis Magpie T an ag er Cissopis leverianus Blue-black G ro s b ea k Cyanocompsa cyanoides * B uff-throated S a ltato r Saltator maxim us G reyish S altato r Saltator coerulescens S late-co lo red G ro s b e a k Saltator grossus * R ed-capped C ardinal Paroaria gularis B lack-striped S p arro w Arremonops conirostris * P ecto ral S p a rro w Arremon taciturnus * L esser Seed-finch Oryzoborus angolensis Lined S e e d e a te r Sporophila lineola * L e sso n ’s S e e d e a te r Sporophila bouvronides * B lack-and-w hite S e e d e a te r Sporophila luctuosa Y ellow -bellied S e e d e a te r Sporophila nigricollis R udd y -b reasted S e e d e a te r Sporophila minuta C h estn u t-b ellied S e e d e a te r Sporophila castaneiventris Y ello w -b ro w ed S p a rro w Am m odram us aurifrons Blue-black G rassq u it Volatinia jacarina

G iant C o w b ird Molothrus oryzivora C re ste d O ro p e n d o la Psarocolius decumanus G re e n O ro p e n d o la Psarocolius viridis R usset-backed O ro p e n d o la Psarocolius angustifrons A m azonian O ro p e n d o la Gymnostinops bifasciatus Y ellow -rum ped C acique Cacicus cela R ed -ru m ped C acique Cacicus haemorrhous * Solitary C acique Cacicus solitarius Ecuadorian C acique Cacicus sclateri * Y ello w -h ooded Blackbird Agelaius icterocephalus * M oriche O rio le Icterus chrysocephalus Blackpoll W a rb le r Dendroica striata (N) A m erican R ed s ta rt Setophaga ruticilla (N) N o rth e rn W a te rth ru s h Seiurus noveboracensis (N) M ourning W a rb le r Oporornis Philadelphia (N) C anada W a rb le r Wilsonia canadensis (N) Buff-rum ped W a rb le r Basileuterus fulvicauda * B ananaquit Coereba flaveola G re e n H o n e y c re e p e r Chlorophanes spiza Purple H o n e y c re e p e r Cyanerpes caeruleus * Blue D acnis Dacnis cayana Black-faced D acnis Dacnis lineata Y ellow -bellied D acnis Dacnis flaviventer * Sw allow T anager Tersina viridis W h ite -v e n te d Euphonia Euphonia minuta Thick-billed Euphonia Euphonia laniirostris R ufous-bellied Euphonia Euphonia rufiventris W h ite -lo re d Euphonia Euphonia chrysopasta * O p a l-ru m p e d T anager Tangara velia O p a l-c ro w n e d T anager Tangara callophrys * Paradise T anager Tangara chilensis G reen -an d -g o ld T anager Tangara schrankii *

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