(ISSN 0892-1016) The Journal of Raptor Research March 1998 Volume 32 Number 1 Contents Preface: Conservation and Ecology of Raptors in the Tropics. Richard t. Watson 1 Conservation Status of Tropical Raptors. Keith l. Biidstein, Wendy Scheisky, jorje Zalles and Susie Ellis 3 Conservation Status of Birds of Prey in the South American Tropics. Richard O. Bierregaard, Jr. 19 Raptors in the East African Tropics and Western Indian Ocean Islands: State of Ecological Knowledge and Conservation Status. Munir virani and Richard T. Watson 28 Current Status and Conservation of Falconiformes in Tropical Asia, jean- Marc Thiollay 40 Tropical Forest Raptors in Indonesia: Recent Information on Distribution, Status, and Conservation, s. (Bas) van Baien 56 Status and Conservation of Raptors in Australia’s Tropics. Nick Mooney 64 BOOK Review. Edited by Jeffrey S. Marks 74 Manuscript Referees 76 The Raptor Research Foundation, Inc. gratefully acknowledges a grant and logistical support provided by Boise State University to assist in the publication of the journal. Persons interested in predatory birds are invited to join The Raptor Research Foundation, Inc. Send requests for information concerning membership, subscriptions, special publications, or change of address to OSNA, P.O. Box 1897, Lawrence, KS 66044-8897, U.S.A. The Journal of Raptor Research (ISSN 0892-1016) is published quarterly and available to individuals for $33.00 per year and to libraries and institutions for $50.00 per year from The Raptor Research Foundation, Inc., 14377 117th Street South, Hastings, Minnesota 55033, U.SA (Add $3 for destinations outside of the continental United States.) Periodicals postage paid at Hastings, Minnesota, and additional mailing offices. POSTMASTER: Send address changes to The Journal of Raptor Research, OSNA, P.O. Box 1897, Lawrence, KS 66044-8897, U.SA Printed by Allen Press, Inc., Lawrence, Kansas, U.SA Copyright 1998 by The Raptor Research Foundation, Inc. Printed in U.SA © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). THE JOURNAL OF RAPTOR RESEARCH A QUARTERLY PUBLICATION OF THE RAPTOR RESEARCH FOUNDATION, INC. Vol. 32 March 1998 No. 1 J. Raptor Res. 32(l):l-2 © 1998 The Raptor Research Foundation, Inc. PREFACE CONSERVATION AND ECOLOGY OF RAPTORS IN THE TROPICS Richard T. Watson The Peregrine Fund, Inc., 566 W. Flying Hawk Lane, Boise, ID 83709 U.S.A. Tropical ecosystems contain the most biological- ly diverse habitats on earth (Wilson 1988). Over 75% of the world’s endemic species occur in trop- ical regions (Bibby et al. 1992) and over 25% of the world’s ecoregions are in the tropics (Bailey 1996). One such ecoregion, tropical rainforest, covers less than 10% of the earth’s land surface yet contains more than half of the species on earth. This is alarming because tropical ecosystems are disappearing faster than all other habitats around the world (Whitmore 1997). Their loss has pro- found implications for the conservation of the world’s biological diversity (Wilson 1988) . The loss of tropical forests is the most conspicuous threat, but the conversion of savanna areas to intensive agriculture is also a threat by reducing available habitat to nonforest species. With an estimated 90% of all species of raptors occurring, wholly or in part, in the tropics (Ken- nedy 1986), and 45% of all raptor species found in tropical rainforests (Thiollay 1994), the loss of tropical ecosystems is especially alarming for these birds. As top-order predators with low population densities, raptors are particularly sensitive to habi- tat loss. In addition to this sensitivity, their high visibility and general allure, make them ideal spe- cies for conserving critically-endangered habitats. Nevertheless, the distribution and status of many raptors are poorly known because many are secre- tive, forest-dwelling species (Meyburg and van Bal- en 1994, Thiollay 1994, van Balen 1994). Because of the need for more information on the current status of tropical raptors and threats that jeopardize their future survival, I organized this symposium as part of the 1996 joint meeting of the Raptor Research Foundation and American Ornithologists’ Union held in Boise, Idaho. Its pur- pose was to bring together currently active re- searchers from each of the tropical biogeograph- ical regions of the world to give first-hand accounts of the status of raptors in their region. Together, these papers represent a complete synthesis of the current state of knowledge and conservation status of the world’s raptors in the tropics. They provide information on distributions and population den- sities, taxonomy, breeding behavior, habitat re- quirements, and sensitivity to habitat modification that can be used to influence the future of raptor research and conservation in many of the world’s tropical areas. Acknowledgments This symposium was supported by the Peregrine Fund, Inc. Publication of this issue was funded in part by the Peregrine Find, Inc. and the Raptor Research Founda- tion, Inc. Literature Cited BAILEY, R.G. 1996. Ecosystem geography. Springer, New York, NY U.S.A. Bibby, C.J., N.J. Collar, M.J. Crosby, M.F. Heath, Ch. Imboden, T.H. Johnson, A.J. Long, A.J. Statters- field and S.J. Thirgood. 1992. Putting biodiversity on the map: priority areas for global conservation I.C.B.P., Cambridge, U.K. Kennedy, R.S. 1986. Raptors in the tropics — the next 50 years. Raptor Res. Rep. 5:17-25. Meyburg, B.-U. and S. van Balen. 1994. Raptors on Su- lawesi (Indonesia): the influence of rainforest de- struction and human density on their populations. Pages 269-276 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. 1 2 Watson Vol. 32, No. 1 Thiollay, J.-M. 1994. A world review of tropical forest raptors: current trends, research objectives and con- servation strategy. Pages 231-239 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation to- day. World Working Group for Birds of Prey and Owls, London, U.K. van Balen, S. 1994. The status and conservation of birds of prey in the Sondaic and Wallacean regions of In- donesia. Pages 245-254 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. Whitmore, T.C. 1997. Tropical forest disturbance, dis- appearance, and species loss. In W.F. Laurence and R.O. Bierregaard, Jr. [Eds.], Tropical forest remnants: ecology, management, and conservation of fragment- ed communities. Univ. Chicago Press, Chicago, IL U.S.A. Wilson, E.O. 1988. Biodiversity. National Academy Press, Washington, DC U.S.A. J. Raptor Res. 32(1):3-18 © 1998 The Raptor Research Foundation, Inc. CONSERVATION STATUS OF TROPICAL RAPTORS 1 Keith L. Bildstein, Wendy Schelsky and Jorje Zalles Hawk Mountain Sanctuary, 1700 Hawk Mountain Road, Kempton, PA 19529 U.S.A. Susie Ellis IUCN Conservation Breeding Study Group, Minneapolis Zoo, Minneapolis, MN U.S.A. Abstract. — Seventy-six percent (222) of the world’s 292 species of diurnal raptors are found mainly or completely in the tropics. Forty-six percent of all tropical raptors are threatened by habitat loss, 11% by environmental contaminants, and 19% by direct persecution. Seventeen percent are threatened by two of these factors, 2% by all three factors. Regionally, 42% of all Neotropical, 60% of all Afrotropical, 60% of all Indomalayan, and 77% of all Australotropical raptors are threatened by one of more of these factors. IUCN classifies 27% of all tropical raptors (59 species) as Near Threatened, Vulnerable, Endan- gered, or Critically Endangered. Similar statistics are 23% (17 of 73 species), 14% (11 of 80 species), 33% (21 of 63), and 34% (12 of 35), for the Neotropical, Afrotropical, Indomalayan, and Australotrop- ical regions, respectively. Thirty percent of all tropical raptors are endemics. Fifty-seven percent of all tropical raptors are complete, partial, or irruptive migrants. The degree of endemism and migration behavior varies among the four regions. Although some of the regional differences in conservation status reflect regional differences in knowledge, many appear to reflect ecological differences among the four regions. Key WORDS: Conservation status ; tropics ; tropical raptors', endemics', migration. Estado de conservacion de las aves rapaces tropicales Resumen. — Setenta y seis por ciento (222) de las especies de aves rapaces diurnas del mundo (292) se encuentran en los tropicos. Cuarenta y seis por ciento de todas las aves rapaces tropicales estan amen- azadas por la perdida de habitat, 11% por contaminantes ambientales y 19% por persecution directa. Diez y siete por ciento estan amenazadas por dos de estos factores, 2% por los tres factores. Regional- mente, 42% de todas las aves rapaces neotropicales, 60% de todas las afrotropicales, 60% de todas las indomalayas y 77% de todas las especies tropicales de Australia estan amenazadas por uno o mas de estos factores. La IUCN clasifica el 27% de todas las aves rapaces tropicales (59 especies) como Cercan- amente Amenazadas, Vulnerables, Amenazadas, o Crfticamente Amenazadas. Estadisticas similares in- dican 23% (17 de 73 especies), 14% (11 de 80 especies), 33% (21 de 63) y 34% (12 de 35), para las regiones neotropical, afrotropical, indomalayay australiana respectivamente. Treinta por ciento de todas las aves rapaces tropicales son endemicas. Cincuenta y siete por ciento de todas las aves rapaces tropi- cales son migratorias totalmente, parcialmente o en forma irruptiva. El grado de endemismo y com- portamiento migratorio varia entre las cuatro regiones. Aunque las diferencias regionales en el estado de conservacion reflejan las diferencias regionales de conocimiento, parece que tambien son el reflejo de las diferencias ecologicas entre las cuatro regiones. [Traduction de Cesar Marquez] Tropical ecosystems are some of the most bio- logically diverse habitats on earth (Wilson 1988). Seventy-six percent of all centers of avian endem- ism (e.g., “endemic bird areas,” Bibby et al. 1992) occur in tropical regions, and the same is true for many plant and animal taxa (Gentry 1986, Mc- 1 This paper is dedicated to the memory of Hawk Moun- tain Sanctuary’s longtime friend, Roger Tory Peterson. Neely et al. 1988, Myers 1988, Thirgood and Heath 1994). Biogeographically, 27% (8 of 30) of the world’s ecoregions are represented in the tropics (Bailey 1996). Tropical rainforests, which cover only 7% of the earth’s land surface, are estimated to contain more than half of the world’s species (Wilson 1988). Previous studies suggest that 90% of all species of raptors occur, wholly or in part, in the tropics 3 4 Bildstein et al. Vol. 32, No. 1 (Kennedy 1986) , and that 45% of all species of rap tors occur in tropical rainforests (Thiollay 1985a, 1994). Half of all tropical countries have at least 50 species of raptors (Table 1). Many tropical raptors, especially forest dwellers, are secretive and difficult to study. The distribution and status of many species are poorly known (Mey- burg and van Balen 1994, Thiollay 1994, van Balen 1994; but see Thiollay 1985b, Kennedy 1986, Burn- ham et al. 1994, Watson and Lewis 1994). Even so, several patterns are apparent. The major threat to tropical raptors is habitat destruction (Thiollay 1989a, 1992, 1994), followed by environmental contamination and shooting (Thiollay 1985a). Overall, endemic species are more likely to be threatened, as are nonmigratory and Old World species. Previous studies of tropical raptors have focused on individual species, countries, regions, or habitat types (e.g., Meyburg and Chancellor 1989, 1994, Baker-Gabb 1994, Thiollay 1994, van Balen 1994, and references therein). We use these and other published sources, including del Hoyo et al. (1994), together with the unpublished IUCN Raptor Con- servation Assessment and Management Plan and Hawk Mountain Sanctuary Hawks Aloft Worldwide databases, to proride an overview of raptor diver- sity and conservation status in the tropics. Specifically, we assess regional differences and the influence of ecological circumstances on the conservation status of tropical species. We detail the distribution of the world’s tropical raptors re- gionally, assess regional differences in diversity and conservation status, and determine the extent to which endemism and migratory behavior are as- sociated with conservation status. We also discuss how conservationists can better protect tropical raptors. Because our focus is global and regional, individual species are mentioned in the text only as examples of general patterns. A list of the con- servation status of individual species appears in Ap- pendix 1. Study Area and Methods The Tropics Defined. Geographically, the tropics in- clude regions on both sides of the Equator extending to 23.5°N and S (the Tropics of Cancer and Capricorn, re- spectively). Terrestrial ecosystems within the region are characterized by near constant day lengths, relatively high solar radiation, and, when adjusted for altitude, rel- atively high diurnal and nocturnal temperatures (Desh- mukh 1986). Although the tropics have a somewhat more complex ecological definition that takes climate, as well as geography, into account (Deshmukh 1986, Bailey 1996), the regions defined both ways are largely coinci- dent. We define the earth’s four tropical regions, the Neotropics, Afrotropics, Indomalayan Region, and Aus- tralotropics, geographically as the land masses between the Tropics of Cancer and Capricorn in each of these four regions. Approximately 85% of the Neotropics is in the Humid Tropical Domain ecoregion (Bailey 1996), of which slightly more than half is savanna; the remainder is trop- ical rainforest. Dry Domain ecoregions in the Neotropics include both steppe and desert. Overall, approximately 40% of the Neotropics is in the Tropical Rainforest ecore- gion. The Afrotropics include the southern half of the Ara- bian Peninsula and most of Africa and Madagascar. Ap- proximately 55% of the region is in the Humid Tropical Domain ecoregion (Bailey 1996). Dry Domain ecore- gions in the Afrotropics are mainly deserts, including the southern Sahara and the Namib, with some steppe. Ap- proximately 10% of the Afrotropics is in the Tropical Rainforest ecoregion. The Indomalayan tropics cover much of southeast Asia and neighboring islands. More than 95% of the region is in the Humid Tropical Domain ecoregion (Bailey 1996) , of which half is savanna, and half is rainforest. Dry Domain ecoregions are limited to eastern India. Overall, approximately 50% of the Indomalayan region is in the Tropical Rainforest ecoregion. The Australotropics include the Moluccas, parts of Aus- tralia, and all of New Guinea. Approximately 60% of the region is part of the Humid Tropical Domain (Bailey 1996), slightly more than half of which is savanna. Dry Domain areas of the Australotropics include tropical por- tions of the Great Sandy Desert of interior Australia, and its associated steppes. Approximately 30% of the Austral- otropics is in the Tropical Rainforest ecoregion. Tropical Raptors Defined. We consider raptors to be members of the avian families Accipitridae, Sagittariidae, and Falconidae; and the avian subfamily Cathartinae (Sibley and Monroe 1990), the birds that most authori- ties call diurnal raptors (Brown and Amadon 1968, Ama- don and Bull 1988, del Hoyo et al. 1994, Griffiths 1994). For our purposes, tropical raptors include the subset of these 292 species (Amadon and Bull 1988) whose com- bined breeding and nonbreeding distributions, as de- tailed in del Hoyo et al. (1994), occur wholly or mainly (>50%) within the tropics. Species (e.g., Swainson’s Hawk [Buleo swainsoni\) , whose migratory routes include the tropics, but whose breeding and nonbreeding ranges lie wholly or mainly outside of the tropics, are not in- cluded as tropical raptors. Endemic Raptors Defined. As a group, raptors are rel- atively large, frequently migratory birds that occur at low densities across large areas. Few can be characterized as restricted-range, or endemic species (e.g., species with breeding ranges below 50 000 km 2 , Bibby et al. 1992). Even so, some species have relatively small ranges com- pared with others. Such species often are used as “um- brella” or “flagship” species for local and regional con- servation efforts (e.g., Philippine Eagle [Pithecophaga jef- feryi, Kennedy 1983], Barred Forest-falcon [Micrastur ruf- icollis, C. Marquez, pers. comm.], Mauritius Kestrel [Falco punctatus, Jones 1981]). Here, we consider endemic spe- March 1998 Status of Tropical Raptors 5 Table 1. Wholly and mainly tropical countries with at least 50 species of raptors; together with the migration, distribution, and conservation status ( sensu Collar et al. 1994) of those raptors. Number of Species Near Critically Country Total (Migrants) Endemics Vulnerable Threatened Endangered Endangered Neotropics Argentina 61 (39) 1 10 Bolivia 68 (40) 1 7 Brazil 66 (39) 1 2 7 Colombia 75 (43) 1 10 1 Costa Rica 53 (34) 6 Ecuador 72 (37) 4 1 9 2 French Guiana 55 (31) 5 Guyana 50 (30) 6 Mexico 54 (41) 4 Panama 57 (34) 7 Paraguay 51 (37) 1 6 Peru 57 (38) 2 9 1 Venezuela 75 (38) 8 Afro tropics Angola 58 (47) 1 1 Botswana 54 (47) 2 1 Burundi 54 (34) 1 1 Cameroon 59 (47) 1 1 Central African Republic 55 (43) 1 1 Chad 52 (47) 1 1 Djibouti 50 (47) 2 1 Ethiopia 71 (61) 4 1 Ivory Coast 50 (38) 1 1 Kenya 72 (62) 3 2 Madagascar 16 (4) 11 3 2 Malawi 56 (46) 2 1 Mozambique 60 (50) 3 2 Namibia 56 (47) 2 2 Nigeria 59 (47) 1 1 Senegal 51 (44) 1 1 Sudan 74 (64) 3 1 Tanzania 69 (58) 2 2 Uganda 71 (59) 1 1 Zaire 65 (52) 1 Zambia 58 (49) 2 1 Zimbabwe 56 (47) 2 2 Indomalayan India 63 (59) 3 4 14 Indonesia 62 (51) (39 [39] ) a 16 (8) 2 (1) 8 (6) 1 (1) Myanmar 52 (47) 4 10 Australotropical Indonesia 62 (33) (39 [22] ) a 16 (9) 2 (1) 8 (2) 1 (0) a Indonesia, which spans the Indomalayan and the Australotropical regions, appears twice in the table. The data presented above represent country- and (tropical-region) totals. 6 Bildstein et al. Vol. 32, No. 1 Table 2. Distribution of the world’s diurnal raptors as a function of tropical distribution. Taxonomy based on Amadon and Bull (1988); distribution based on del Hoyo et al. (1994). Number of Species (%) a Extent of Range in Tropics Worldwide Neotropics Afrotropics Indomalayan Australotropics All of range 78 (27%) 20 (22%) 17 (15%) 22 (26%) 20 (41%) More than half of range 144 (49%) 53 (58%) 63 (56%) 41 (49%) 15 (31%) Less than half of range 60 (21%) 19 (21%) 32 (29%) 21 (25%) 14 (29%) None of range 10 (3%) Total 292 92 112 84 49 a Worldwide percentages represent proportions of all species, including those that occur only outside of the tropics. Regional per- centages represent proportions of species of raptors from that region. cies to be raptors whose overall ranges are limited to one or two countries. We consider endemic genera to be gen- era that occur in only one tropical region. Threats to Tropical Raptors. Human threats to tropical raptors include habitat destruction and fragmentation, pollution and pesticide use, and direct human persecu- tion (Newton and Chancellor 1985, Diamond and Love- joy 1985, Meyburg and Chancellor 1989, 1994, and ref- erences therein). Statistical Tests. We used log-likelihood ratio (G) tests with Yates correction in tests where df = 1 (Zar 1984), to assess regional differences in the endemic, migration and conservation status of tropical raptors, as well as to determine the extent to which endemism and migration were associated with differences in the conservation sta- tus of the birds. In all analyses, differences were consid- ered significant if P < 0.05. Results Distribution, Endemism, and Migratory Status. Seventy-eight species of raptors (27%) occur whol- ly within the tropics. An additional 144 species (49%) occur mainly within the tropics. Overall, 282 species (97%) of the world’s raptors have breeding or wintering distributions that occur, at least in part, within one or more tropical regions (Table 2). Seven countries have 70 or more species of tropical raptors (Table 1). The distribution and status of the world’s 222 wholly or mainly tropical raptors appear in Appendix 1. Raptor faunas differ considerably among the four regions. Eighty-nine percent of tropical rap- tors occur in a single tropical region, and except for the contiguous Indomalayan and Australotrop- ical regions, there is little species overlap among regions (Table 3). Regional faunal specificity is greatest in the Neotropics, where 72 of 73 species have tropical distributions restricted to that region; the single exception being the nearly cosmopolitan Osprey ( Pandion haliaetus) (Table 3). Twenty-six Neotropical genera (50 species) occur only in that tropical region, compared with 12 (19), 5 (10), and 4 (5), genera (species), respectively, in the Af- rotropics, Indomalayan Region, and Australotrop- ics (Table 4). Table 3. Regional distribution of tropical raptors. Species with region-restricted ranges are those that occur in only one tropical region. Widespread species occur in more than one tropical region. Number of Species (%) Extent of Range Worldwide Neotropics Afrotropics Indomalayan Australotropics Wholly tropical raptors Region-restricted 77 (99%) 20 (100%) 17 (100%) 21 (95%) 19 (95%) Widespread 1 (1%) 0 0 1 (5%) 1 (5%) Mainly tropical raptors Region-restricted 121 (84%) 52 (98%) 48 (76%) 18 (44%) 3 (20%) Widespread 23 (16%) 1 (2%) 15 (24%) 23 (56%) 12 (80%) Wholly and mainly tropical raptors Region-restricted 198 (89%) 72 (99%) 65 (81%) 39 (62%) 22 (63%) Widespread 24 (11%) 1 (1%) 15 (19%) 24 (38%) 13 (37%) March 1998 Status of Tropical Raptors 7 Table 4. Distribution of tropical endemics. Endemic species are species whose range is restricted to one, or at most, two countries; endemic genera are genera restrict- ed to one tropical region. Region Number (%) Endemic Species Number (%) Endemic Genera Neotropics 7 (10%) (a)* 26 (76%) (a) Afrotropics 12 (15%) (a) 12 (38%) (b) Indomalayan 18 (29%) (b) 5 (21%) (b) Australotropics 22 (63%) (c) 4 (25%) (b) All regions 67 (30%) 47 (70%) a Regions with the same letters are not significantly different from one another (log-likelihood ratio [G] test; species: df = 1; P < 0.05; genera: df = 1; P < 0.05). Endemism is high in tropical raptors. Fifty-six percent of all wholly tropical raptors (44 of 78 spe- cies) occur in only one or two countries, compared with only 12% of all other raptors (log-likelihood ratio test: G — 54.3, df — 1, P < 0.001). The Aus- tralotropics have more endemic species than any other tropical region (vs. Neotropics: log-likeli- hood ratio test: G = 30.7, df = 1, P < 0.001; vs. Afrotropics: G = 23.6, df = 1 , P < 0.001; vs. In- domalayan: G — 9.6, df = 1, P < 0.005; Table 4). Ninety-one percent of all endemic tropical rap- tors (51 of 58 species) occur on islands, mainly in the Indian and South Pacific oceans. Three coun- tries, Indonesia (16), Papua New Guinea (14), and Madagascar (11), together have 41 species of en- demic raptors. Nine percent of all wholly, and 32% of all mainly tropical raptors, are complete or partial migrants (Kerlinger 1989). An additional 6% and 24%, re- spectively, are local or irruptive migrants (Table 5) . There are significantly more migratory raptors (lo- cal-irruptive species included) in the Afrotropics than in any other tropical region (vs. Neotropics: log-likelihood ratio test: G = 17.4, df = 1, P < 0.001; vs. Indomalayan: G = 4.8, df = 1, P < 0.05; vs. Australotropics: G = 11.5, df = 1, P < 0.001; Table 5). Conservation Status. More than half of all trop- ical raptors are threatened by habitat loss, environ- mental contaminants, direct persecution, or com- binations of these factors (Table 6). Habitat loss appears to be the greatest threat (habitat loss vs. environmental contaminants: log-likelihood ratio test: G = 69.3, df = 1, P < 0.001; habitat loss vs. direct persecution: G = 35.2, df = 1, P < 0.001; Table 6). Neotropical raptors appear to be less threatened than those from other tropical regions (Table 6) . Direct persecution, especially, appears to occur less in the Neotropics than in other tropical regions (Table 6). Environmental contaminants, including pesticides, appear to be particularly im- portant threats in the Australotropics (Table 6) . Thirty-seven species of wholly or mainly tropical raptors (17% of the world’s tropical raptors) are listed as Critically Endangered, Endangered, or Vulnerable (Collar et al. 1994; Table 7). An addi- tional 22 species (10%) are listed as Near Threat- ened. Two species are listed as data deficient (Ta- ble 7) . Wholly tropical raptors appear be at greater risk than are mainly tropical raptors (log-likeli- hood ratio test: G = 17.1, df = 1 , P< 0.001; Table 7). Nonmigratory species appear to be at greater risk than migratory ones (log-likelihood ratio test: G = 19.3, df = 1, P< 0.001; Table 8). Forty (68%) of all Threatened and Near Threatened species are forest-dependent (Appendix 1). Twenty-eight at- risk species (47%) are restricted to islands. Twenty- one at-risk species (36%) are both forest-depen- dent and island-restricted. Countries with Endangered or Critically Endan- gered tropical raptors include Colombia (1 Endan- gered, 0 Critically Endangered), Cuba (1,0), Do- minican Republic (1,0), Ecuador (2,0), Haiti (1,0), Peru (1,0), Madagascar (0,2), Mauritius (1,0), In- Table 5. Migratory status of wholly and mainly tropical raptors. Status is based primarily on Kerlinger (1989) , as modified by del Hoyo et al. (1994) and Hawks Aloft Worldtvide. Migratory Status Worldwide Neotropics Afrotropics Indomalayan Australotropics Complete 11 (5%) 2 (3%) 7 (9%) 7 (11%) 3 (9%) Partial 61 (27%) 16 (22%) 33 (41%) 21 (33%) 7 (20%) Local-irruptive 54 (24%) 16 (22%) 24 (30%) 11 (17%) 6 (17%) Total migratory 126 (57%) 34 (47%) (a) a 64 (80%) (b) 39 (62%) (a) 16 (46%) (a) Nonmigratory 96 (43%) 39 (53%) 16 (20%) 24 (38%) 19 (54%) a Regions with the same letters are not significantly different from one another (log-likelihood ratio [G] test; df = 1; P < 0.05). 8 Bildstein et al. Vol. 32, No. 1 Table 6. Threats to tropical raptors. Threats are based, principally, on information in del Hoyo et al. (1994). Number of Species Affected (%) Threat Worldwide Neotropics Afrotropics Indomalayan Australotropics Wholly tropical raptors Habitat loss Environmental contami- 38 (49%) 7 (35%) 10 (59%) 8 (36%) 14 (70%) nants 6 (8%) 0 2 (12%) 0 4 (20%) Direct persecution Total number of species 10 (13%) 1 (5%) 2 (12%) 2 (9%) 5 (25%) affected 40 (51%) 7 (35%) 11 (65%) 8 (36%) 15 (75%) Number of species affected by: two threats 12 (15%) 1 (5%) 3 (18%) 2 (9%) 6 (30%) three threats 1 (1%) 0 0 0 1 (5%) Mainly tropical raptors Habitat loss Environmental contami- 64 (44%) 20 (38%) 25 (40%) 26 (63%) 8 (53%) nants 18 (13%) 6 (11%) 10 (16%) 7 (17%) 5 (33%) Direct persecution Total number of species 33 (23%) 5 (9%) 22 (35%) 13 (32%) 8 (53%) affected 81 (56%) 24 (45%) 37 (59%) 30 (73%) 12 (80%) Number of species affected by: two threats 26 (18%) 6 (11%) 16 (25%) 10 (24%) 4 (27%) three threats 4 (3%) 0 2 (3%) 3 (7%) 2 (13%) Wholly and mainly tropical raptors Habitat loss Environmental contami- 102 (46%) 27 (37%) (a) a 35 (44%) (a, b) l 34 (54%) (a, b) 22 (63%) (b) nants 24 (11%) 6 (8%) (a) 12 (15%) (a, b) l 7 (11%) (a) 9 (26%) (b) Direct persecution Total number of species 43 (19%) 6 (8%) (a) 24 (30%) (b) 15 (24%) (b) 13 (37%) (b) affected 121 (55%) 31 (42%) (a) 48 (60%) (b) 38 (60%) (b) 27 (77%) (b) Number of species affected by: two threats 38 (17%) 7 (10%) 19 (24%) 12 (19%) 10 (29%) three threats 5 (2%) 0 2 (3%) 3 (5%) 3 (9%) a Regions with the same letters are not significantly different from one another (log-likelihood ratio [G] test; df = l ;P< 0.05). donesia (1,0), the Philippines (0,1), Papua New Guinea (1,0), and the Solomon Islands (1,0) (Col- lar et al. 1994). Discussion Our analyses demonstrate several patterns that should be of use to conservationists. They show that there are high levels of within-country diver- sity and endemism in all tropical regions, there is high congruence of raptor species diversity and en- demism with those of other taxa, there is an indi- cation that habitat loss (as opposed to environ- mental contaminants and direct persecution) pos- es the greatest threat, there are associations be- tween at-risk conservation status and endemism, sedentary behavior, forest-dependence and island- dwelling, and there are significant regional differ- ences in the extent of migration behavior, endem- ism, and conservation status. Diversity and Endemism in Tropical Raptors. More than three-quarters of the world’s diurnal birds of prey occur wholly or mainly within the tropics, and more than one-third of these occur only within this region. Three “megadiversity” countries ( sensu Mittermeier 1988) in the Neotrop- ics (Colombia, Ecuador, and Venezuela) and four in the Afrotropics (Ethiopia, Kenya, Sudan, and Uganda) each contain 70 or more species of rap- March 1998 Status of Tropical Raptors 9 Table 7. Conservation status of tropical raptors based on Collar et al. (1994). Conservation Status Worldwide Neotropics Afrotropics Indomalayan Australo- TROPICS Wholly tropical raptors Critically endangered 3 (4%) 0 2 (12%) 1 (5%) 0 Endangered 6 (8%) 3 (15%) 1 (6%) 1 (5%) 1 (5%) Vulnerable 7 (9%) 1 (5%) 0 3 (14%) 3 (15%) Total Threatened 16 (21%) 4 (20%) 3 (18%) 5 (23%) 4 (20%) Near Threatened 18 (23%) 3 (15%) 1 (6%) 8 (36%) 6 (30%) Not Threatened 42 (53%) 13 (65%) 13 (76%) 8 (36%) 9 (45%) Insufficient data 2 (3%) 0 0 1 (5%) 1 (5%) Mainly tropical raptors Critically Endangered 0 0 0 0 0 Endangered 1 (1%) 0 0 0 1 (7%) Vulnerable 5 (3%) 2 (4%) 2 (3%) 0 1 (7%) Total Threatened 6 (4%) 2 (4%) 2 (3%) 0 2 (13%) Near Threatened 19 (13%) 8 (15%) 5 (8%) 8 (20%) 0 Not Threatened 119 (83%) 43 (81%) 56 (89%) 33 (80%) 13 (87%) Insufficient data 0 0 0 0 0 Wholly and mainly tropical raptors Critically Endangered 3(1%) 0 2 (3%) 1 (2%) 0 Endangered 7 (3%) 3 (4%) 1 (1%) 1 (2%) 2 (6%) Vulnerable 12 (5%) 3 (4%) 2 (3%) 3 (5%) 4 (11%) Total Threatened 22 (10%) 6 (8%) 5 (6%) 5 (8%) 6 (17%) Near Threatened 37 (17%) 11 (15%) 6 (8%) 16 (25%) 6 (17%) Not Threatened 161 (73%) 56 (77%) 69 (86%) 41 (65%) 22 (63%) Insufficient data 2 (1%) 0 0 1 (2%) 1 (3%) tors and have at least one-quarter of the world’s raptor species each. Fourteen rainforest “hot spots” (Myers 1988) which together comprise less than 1 % of the earth’s land surface contain more than 30% of the world’s raptor species (Thiollay 1994). Tropical islands, many of whose natural habitats have been or are being destroyed (Green and Suss- Table 8. Conservation status of nonmigratory and mi- gratory wholly and mainly tropical raptors compared (conservation status based on Collar et al. [1994]). Conservation Status Non- migratory Migratory Critically endangered 3 (3%) 0 Endangered 6 (6%) 1 (1%) Vulnerable 7 (7%) 5 (4%) Total threatened 16 (17%) 6 (5%) Near threatened 24 (25%) 13 (10%) Not threatened 54 (56%) 107 (85%) Insufficient data 2 (2%) 0 man 1990, Hannah et al. 1994), host 24% (18 spe- cies) of all Threatened and Near Threatened spe- cies of raptors. The tropics are home to most of the world’s en- demic raptors, as well as to most of its endangered species. Indonesia, alone, has 24% (16 of 67 spe- cies) of all endemic birds of prey. Indeed, one rea- son for the high species diversity of tropical raptors is high regional endemism. For example, while the Nearctic and Western Palearctic share 11% (8 spe- cies) of their combined raptor faunas, their tropi- cal counterparts (e.g., the Neotropics and Afro- tropics) share less than 1% (1 of 152) of their com- bined faunas. Although overlap among the Afro- tropics, Indomalayan Region, and Australo tropics is higher than that between these regions and the Neotropics (Table 3), differences in faunal com- position among tropical regions is extensive, over- all. Seventy percent of all tropical genera (47 of 67) occur in a single tropical region. Congruence of Raptor Diversity with Biodiver- sity in Other Groups of Plants and Animals . Geo- 10 Bildsteim et al. Vol. 32, No. 1 graphic centers of raptor diversity and species en- demism are highly congruent with those for many groups of plants and animals (Gentry 1986, Mc- Neely et al. 1988, Myers 1988, Bibby et al. 1992, Thirgood and Heath 1994). Countries with ex- tremely high raptor diversity, as well as centers of raptor endemism, are important conservation sites for plants and other animals, too. Principal Threats to Raptors. Habitat loss re- mains the principal threat to tropical raptors. Many natural ecosystems in the region are being con- verted to human-dominated landscapes (Myers 1984). A recent global analysis of human habitat disturbance (Hannah et al. 1994), for example, suggests that the Indomalayan and Afrotropical realms are the two most disturbed biogeographical realms ( sensu Udvardy 1975) on earth. Many bio- geographical provinces, including the Cuban and Greater Antillean provinces in the Neotropics, the Malagasy woodland-savanna province in the Afro- tropics, and the Java and Philippines province in the Indomalayan Realm, are overwhelmingly dom- inated by human landscapes (Hannah et al. 1994) . Many disturbed provinces harbor endemic and at- risk species of raptors. Because many tropical hab- itats are only now being threatened by human dis- turbance (Hannah et al. 1994), habitat loss will re- main a principal concern for some time. Overall, 68% of all Threatened and Near Threat- ened tropical raptors are forest-dependent species. Many of these species require enormous tracts of contiguous forest (Thiollay 1989a, 1993), and are thus particularly vulnerable to habitat loss (Dia- mond and Lovejoy 1985, Meyburg and Chancellor 1989, 1994, and references therein). All but one of Australia’s 15 Threatened raptors are forest dwell- ers (Baker-Gabb 1994). Almost half (28 of 59 species) of all Threatened and Near Threatened tropical raptors are restrict- ed to islands. Current levels of habitat degradation on tropical islands (Hannah et al. 1994), together with the recent history of human-induced avian ex- tinctions at such sites (Moors 1985, Temple 1986, Wotzkow 1994), suggests that these species, too, are at high risk. Gregarious and nomadic, open-habitat species, including those that depend upon superabundant prey associated with seasonally fluctuating wetlands (e.g., Snail Kites [Rostrhavnus sociabilis]) , and upon swarming locusts (e.g., Montagu’s [Circus py gar gus\ and Pallid Harriers [C. macrourus], Swainson’s Hawks, Western [Falco vespertinus ] and Eastern Red-footed Falcons [F amurensis ], and Lesser Kes- trels [F. naumanni ] ) , also appear to be particularly vulnerable to habitat loss and degradation. Be- cause members of these species operate over enor- mous ecological neighborhoods, local habitat loss can significantly affect regional, and even conti- nental populations of these raptors (Bucher 1992) . The social nature of these species, together with the fact that they frequently feed on prey that are considered agricultural pests, means that large numbers can be poisoned by a single pesticide ap- plication (Thiollay 1989b, del Hoyo et al. 1994, Woodbridge et al. 1995). Even when individuals are not poisoned, large-scale efforts to remove in- sect pests, including swarming locusts from tropical regions, can threaten these species by significandy reducing prey availability (Thiollay 1989b). Overall, the increasing use of agricultural pesti- cides in many tropical countries (Goulston 1996) suggests that this threat, which has long been a concern in temperate agriculturalized countries (Hickey 1969, Cooke et al. 1982), is likely to in- crease. Tropical raptors also are threatened by the use of pesticides to control insect vectors of human diseases (Thiollay 1989b). Although direct persecution as a threat to trop- ical raptors appears to be decreasing overall (Thiollay 1994), shooting remains a threat in cer- tain countries (van Balen 1994, Bildstein et al. 1995). Social species such as vultures, and species that flock on migration or that roost communally, as well as those that are considered to be agricul- tural or aquacultural pests, are especially vulnera- ble (Thiollay 1989b). Given the region’s increasing dependence upon aquaculture as a source of hu- man protein (World Resources Institute 1996), persecution of piscivorous raptors is likely to in- crease (FAO 1989 and references therein). Regional Differences. The current geographic distribution of tropical raptors is the result of com- plex interactions among climatic, geomorphologic, and ecological processes operating across numer- ous temporal and spatial scales (Sankovski and Pridnia 1995). In the Indomalayan and Australo- tropical regions, for example, raptor diversity is en- hanced because of numerous island endemics in Indonesia (16 species) and New Guinea (14 spe- cies), respectively (Table 4). In the Afrotropics, on the other hand, where species diversity is bolstered in part by island endemics from Madagascar, much of the region’s diversity results from the high num- ber of migrants from the Western and Eastern Pa- March 1998 Status of Tropical Raptors 11 learctic (Newton 1995) (Table 5) . In the Neotrop- ics, diversity appears to result from geographic fea- tures and evolutionary processes particular to the region (e.g., forest refugia and centers of endem- ism, Haffer 1974 and Cracraft 1985). As a result, while the Neotropics have fewer endemic species (7) than any other region, they have significantly more endemic genera (26) (Table 4) . The Neotropics, and to a lesser extent the Af- rotropics (particularly when only continental Afri- ca is considered), have proportionately fewer at- risk species than do the Indomalayan and Austral- otropics. This difference appears to be attribut- able, mainly, to two factors: the greater number of island species in the Indomalayan Region and the greater incidence of disturbed ecosystems there (Hannah et al. 1994). Final Thoughts and Suggestions. Many conser- vation biologists have suggested focusing conser- vation efforts on megadiversity countries (Mitter- rneier 1988), biodiversity “hot spots” (Myers 1988), specific habitats types (Wilson 1988 and ref- erences therein) , or endemic bird areas (Ribby et al. 1992). Thiollay (1994), for example, uses this approach in identifying four priority areas of trop- ical forest raptor diversity, which together contain 40% of all tropical forest raptor species. All four areas, however, occur in only two tropical regions (the Indomalayan and Australotropical) , and fo- cusing on these sites will do little to protect the very different raptor communities of other regions. Recently, several conservation biologists have suggested that when choices need to be made on which species to protect, efforts should focus on species with the greatest “phylogenetic diversity” (e.g., those most distantly related to other existing taxa, Forey et al. 1994) . Eleven tropical raptors be- long to genera in which all members of the genus are Threatened or Near Threatened. The Neotrop- ics have five such species ( Harpyhaliaetus solitarius, H. coronatus, Morphnus guianensis, Harpia harpyja, and Spizastur melanoleucus) , the Afrotropics have one ( Eutriorchis astur) , the Indomalayan Region has three ( Ichthyophaga humilis, /. icMhyaetus, and Pithe- cophaga jefferyt ) , and the Australotropics have two ( Lophoictinia isura and Harpyopsis novaeguineae) . Ten of these species are largely or entirely seden- tary, eight are large eagles (>1500 g), six are for- est-dependent species, and three are island forms. These species, especially, deserve focused conser- vation efforts. One of the greatest challenges facing conserva- tionists interested in protecting tropical raptors is our ignorance of the biology of many of these spe- cies. This is true not only of secretive forest-dwell- ing, range-restricted endemics (Meyburg and Chancellor 1994, and references therein), but also for wide-ranging migratory species whose ecology outside of the tropics is relatively well studied (Sen- ner and Fuller 1989, Newton 1995). Indeed, while we appear to have good information on species dis- tributions, we lack reasonable population estimates for many species of tropical raptors (del Hoyo et al. 1994), as well as information on factors limiting their distribution and abundance. We also know relatively litde about the extent to which tropical residents interact with migrants overwintering in the region (but see Kirk and Gosler 1994). We strongly recommend the expansion of field- work in tropical areas. Initiatives to gather infor- mation on species’ ecology, distributions, and abundances (Burnham et al. 1994) and Hawk Mountain Sanctuary’s Hawks Aloft Worldwide, which uses the spectacle of long-distance migration to strengthen local conservation activities along ma- jor migratory corridors in the tropics (Bildstein et al. 1995), together with meetings and publications that focus on tropical raptors (Meyburg and Chan- cellor 1989, 1994), should be continued and ex- panded. Because the fate of tropical raptors ultimately rests in the hands of the human inhabitants of these areas, individuals and organizations interest- ed in protecting tropical raptors need to encour- age and support training opportunities for conser- vationists and biologists living in the region. Acknowledgments We thank the hundreds of raptor biologists, conserva- tionists, and field-workers who contributed data essential to this manuscript. We especially thank the contributors and compilers of del Hoyo et al. (1994), including RO. Bierregaard, W.S. Clark, S.J.S. Debus, A. Elliot, D.C. Elouston, J. del Hoyo, A.C. Kemp, L.F. Riff, B.-U. Mey- burg, P.D. Olsen, J. Orta, A.F. Poole, J. Sargatal, J.-M. Thiollay, and C.M. White. We also thank the dozens of participants of IUCN raptor CAMP workshops in Bada- joz, Spain, Asuncion, Paraguay, and Duluth, MN U.S.A , the taxon and geographic editors associated with that ef- fort, and the 10 conservation organizations working to- gether on the project (BirdLife International, Fonds d’lntervention pour les Rapaces, The Hawk and Owl Trust, Hawk Mountain Sanctuary, The National Birds of Prey Centre, The Peregrine Fund, The Raptor Conser- vation Group, The Raptor Research Foundation, The World Working Group for Birds of Prey and Owls, IUCN- 12 Bildstein et al. Vol. 32, No. 1 SSC Conservation Breeding Specialist Group). Also, we thank the more than 400 individuals who have contrib- uted data to Hawk Mountain Sanctuary’s Hawks Aloft Worldwide initiative. We thank Rick Watson for inviting us to participate in the symposium on tropical raptors, and Rick Watson, Marc Bechard, and two anonymous referees, for editing and improving our ms. Eric Atkinson provided statistical advice and analyses. This is Hawk Mountain Sanctuary contribution number 54. Literature Cited Amadon, D. and J. Bull. 1988. Hawks and owls of the world: a distributional and taxonomic list. Proc. West. Found. Vertebr. Zool. 3:295-357. Bailey, R.G. 1996. Ecosystem geography. Springer, New York, NY U.S.A. Baker-Gabb, D.J. 1994. Threatened raptors of Australia’s tropical forests. Pages 241-244 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. Bibby, C.J., N.J. Collar, M.J. Crosby, M.F. Heath, Ch. Imboden, T.H. Johnson, A.J. Long, A.J. Sattersfield and S.J. Thirgood. 1992. Putting biodiversity on the map: priority areas for global conservation. ICBP, Cambridge, U.K. Bildstein, K.L., J.J. Brett, L.J. Goodrich and C. Viver- ette. 1995. Hawks Aloft Worldwide: a network to pro- tect the world’s migrating birds of prey and the hab- itats essential to their migrations. Pages 504—16 in D.A. Saunders, J.L. Craig and E.M. Mattishe [Eds.], Nature conservation. Surrey Beaty 8c Sons, Chipping Norton, N.S.W., Australia. Brown, L. and D. Amadon. 1968. Eagles, hawks and fal- cons of the world. McGraw-Hill, New York, NYU.S.A. Bucher, E.H. 1992. The causes of extinction of the Pas- senger Pigeon. Curr. Ornithol. 9:1-36. Burnham, W.A., D.F. Whitacre and J.P. Jenny [Eds.]. 1994. The Maya Project: use of raptors as tools for conservation and ecological monitoring of biological diversity. Pages 257-264 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. Collar, N.J., M.J. Crosby and A.J. Sattersfield. 1994. Birds to watch 2: the world list of threatened birds. BirdLife International, Cambridge, U.K. Cooke, A.S., A.A. Bell and M.B. Haas. 1982. Predatory birds, pesticides, and pollution. Institute of Terrestrial Ecology, Huntingdon, U.K. Cracraft,J. 1985. Historical biogeography and patterns of differentiation within the South American avifau- na: areas of endemism. Ornithol. Monogr. 36:49-84. del Hoyo, J., A. Elliot and J. Sargatal [Eds.]. 1994. Handbook of the birds of the world, Vol. 2. Lynx Ed- icions, Barcelona, Spain. Deshmukh, I. 1986. Ecology and tropical biology. Black- well Scientific, Palo Alto, CA U.S.A. Diamond, A.W. and T.E. Lovejoy [Eds.]. 1985. Conser- vation of tropical forest birds. ICBP, Tech. Publ. No. 4, Cambridge, U.K. FAO. 1989. Report of the EIFAC working party on pre- vention and control of bird predation in aquaculture and fisheries operations. Food and Agriculture Or- ganization of the United Nations, Rome, Italy, Forey, P.L., C.J. Humphries and R.I. Vane-Wright [Eds.]. 1994. Systematics and conservation evaluation. Clar- endon Press, Oxford, U.K. Gentry, A.H. 1986. Endemism in tropical vs. temperate plant communities. Pages 153—181 in M.E. Soule [Ed.], Conservation biology. Sinauer, Sunderland, MA U.S.A. Goulston, G. 1996. Crop protection in Latin America. PJB Publications, Ltd., London, U.K. Green, G.M. and R.W. Sussman. 1990. Deforestation his- tory of the eastern rainforests of Madagascar from sat- ellite images. Science 248:212-215. Griffiths, C.S. 1994. Monophyly of the Falconiformes based on syringeal morphology. Auk 111:787-805. Haffer, J. 1974. Avian speciation in tropical South America. Publ. Nuttall Ornithol. Club, No. 14. Nuttall Ornithological Club, Cambridge, MA U.S.A. Hannah, L., D. Lohse, C. Hutchinson, J.L. Carr and A. Lankerani. 1994. A preliminary inventory of human disturbance of world ecosystems. Ambio 23:246-250. Hickey, J.J. [Ed.] 1969. Peregrine Falcon populations: their biology and decline. Univ. of Wisconsin, Madi- son, WI U.S.A. Jones, C.G. 1981. The Mauritius Kestrel. Its biology and conservation. Hawk Trust Annu. Rep. 10:18-20. Kennedy, R.S. 1983. Can Filipinos learn to love this bird? Int. Wildl. 13(4):26-33. . 1986. Raptors in the tropics — the next 50 years. Raptor Res. Rep. 5:17-25. Kerlinger, P. 1989. Flight strategies of migrating hawks. Univ. Chicago Press, Chicago, IL U.S.A, Kirk, D.A. and A.G. Gosler. 1994. Body condition varies with migration and competition in migrant and resi- dent South American vultures. Auk 111:933-944. McNeely, J.A., K.R. Miller, W.V. Reid, R.A. Mittermeier and T.B. Werner. 1988. Conserving the world’s bio- logical diversity. IUCN, Gland, Switzerland. Meyburg, B.-U. and R.D. Chancellor. 1989. Raptors in the modern world. World Working Group for Birds of Prey and Owls, London, U.K. and . 1994. Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. and S. VAN Balen. 1994. Raptors on Sulawesi (In- donesia): the influence of rainforest destruction and human density on their populations. Pages 269-276 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor March 1998 Status of Tropical Raptors 13 conservation today. World Working Group for Birds of Prey and Owls, London, U.K. Mittermeier, R.A. 1988. Primate diversity and the trop- ical forest: case studies from Brazil and Madagascar and the importance of megadiversity countries. Pages 145-154 in E.O. Wilson [Ed.], Biodiversity. National Academy, Washington, DC U.S.A. Moors, PJ- 1985. Conservation of island birds. I.C.B.P., Tech. Publ. No. 3, Cambridge, U.K. Myers, N. 1984. The primary source. Norton, New York, NYU.S.A. . 1988. Threatened biotas: hot spots in tropical forests. The Environmentalist 10:243-256. Newton, I. 1995. Relationships between breeding and wintering ranges in Palaearctic-African migrants. Ibis 137:241-249. and R.D. Chancellor. 1985. Conservation stud- ies on raptors. ICBP, Tech. Publ. No. 5, Cambridge, U.K. Sankovski, A. and M. Pridnia. 1995. A comparison of the southern Appalachian (U.S.A.) and southwestern Caucasus (Russia) forests: influences of historical events and present environment./. Biogeogr. 22:1073— 1081. Senner, S.E. and M.R. Fuller. 1989. Status and conser- vation of North American raptors migrating in the Neotropics. Pages 53-58 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptors in the modern world. World Working Group for Birds of Prey and Owls, London, U.K. Sibley, C.G. and B.L. Monroe, Jr. 1990. Distribution and taxonomy of birds of the world. Yale Univ. Press, New Haven, CT U.S.A. Temple, S.A. [Ed.]. 1986. Endangered birds. Univ. Wis- consin Press, Madison, WI U.S.A. Thiollay, J.-M. 1985a. Falconiformes of tropical rain- forests: a review. Pages 155-165 in I. Newton and R.D. Chancellor [Eds.], Conservation studies on raptors. ICBP, Cambridge, U.K. . 1985b. The West African forest avifauna: a re- view. Pages 171-186 in I. Newton and R.D. Chancellor [Eds.], Conservation studies on raptors. ICBP, Cam- bridge, U.K. . 1989a. Area requirements for the conservation of rainforest raptors and game birds in French Guia- na. Conserv. Biol. 3:128-137. . 1989b. Distribution and ecology of Palearctic birds of prey wintering in West and Central Africa. Pages 99-107 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptors in the modern world. World Working Group for Birds of Prey and Owls, London, U.K. . 1992. Influence of selective logging on bird spe- cies diversity in a Guianan rainforest. Conserv. Biol 6 - 47-63. . 1993. Response of a raptor community to shrinking area and degradation of tropical rainforest in the south west Ghats (India). Ecogeography 16:97- 110 . . 1994. A world review of tropical forest raptors. current trends, research objectives and conservation strategy. Pages 231-239 in B.-U. Meyburg and R D Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. Thirgood, S.J. and M.F. Heath. 1994. Global patterns of endemism and the conservation of biodiversity. Pages 207-227 in P.L. Forey, C.J. Humphries and R.I. Vane-Wright [Eds.], Systematics and conservation evaluation, Clarendon Press, Oxford, U.K. Udvardy, M.D.F, 1975. A classification of the biogeo- graphical provinces of the world. IUCN, Morges, Swit- zerland. van Balen, S. 1994. The status and conservation of birds of prey in the Sondaic and Wallacean regions of In- donesia. Pages 245-254 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. Watson, R.T. and R. Lewis. 1994. Raptor studies in Mad- agascar’s rainforest. Pages 283-290 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation to- day. World Working Group for Birds of Prey and Owls, London, U.K. Wilson, E.O. [Ed.]. 1988. Biodiversity. National Acade- my Press, Washington, DC U.S.A. Woodbridge, B., K.K. Finley and S.T. Seager. 1995. An investigation of the Swainson’s Hawk in Argentina./ Raptor Res. 29:202-204. World Resources Institute. 1996. World resources, 1996-97. World Resources Institute, New York, NY U.S.A. Wotzkow, C. 1994. Status, distribution, current research and conservation of forest birds of prey in Cuba. Pages 291-299 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. Zar, J.H. 1984. Biostatistical analysis. Second Ed. Pren- tice Hall, Englewood Cliffs, NJ U.S.A. Received 16 October 1996; accepted 10 November 1997 14 Bildstein et al. Vol. 32, No. 1 Appendix 1. Conservation and distribution status of the world’s tropical raptors. Species IUCN Conservation Status Endemic Forest Dependent Island Dependent Tropical Distribution Neotropics Coragyps atratus Nt a No No No Mainly Cathartes aura Nt No No No Mainly Cathartes burrovianus Nt No No No Mainly Cathartes melambrotus Nt No Yes No Wholly Sarcoramphus papa Nt No No No Mainly Pandion haliaetud 0 ’ Nt No No No Mainly Leptodon cayanensis Nt No Yes No Mainly Chondrohierax undnatus Nt No Yes No Mainly Elanoides forficatus Nt No Yes No Mainly Gampsonyx swainsonii Nt No No No Mainly Elanus leucurus Nt No No No Mainly Rostrhamus sodabilis Nt No No No Mainly Rostrhamus hamatus Nt No Yes No Wholly Harpagus bidentatus Nt No No No Mainly Harpagus diodon Nt No Yes No Mainly Ictinia plumbea Nt No No No Mainly Circus buffoni Nt No No No Mainly Acdpiter poliogaster Near tThreatened No Yes No Mainly Acdpiter superdliosus Nt No Yes No Mainly Acdpiter collaris Near threatened No Yes No Wholly Acdpiter gundlachi Endangered Yes No Yes Wholly Acdpiter bicolor Nt No No No Mainly Geranospiza caerulescens Nt No No No Mainly Leucopternis schistacea Nt No Yes No Wholly Leucopternis plumbea Near threatened No Yes No Wholly Leucopternis princeps Nt No Yes No Wholly Leucopternis melanops Nt No Yes No Wholly Leucopternis kuhli Nt No Yes No Wholly Leucopternis lacernulata Vulnerable Yes Yes No Mainly Leucopternis semiplumbea Near threatened No Yes No Wholly Leucopternis albicollis Nt No Yes No Wholly Leucopternis ocddentalis Endangered Yes Yes No Wholly Leucopternis polionota Near threatened No Yes No Mainly Asturina nitida Nt No No No Mainly Buteogallus aequinoctialis Nt No Yes No Mainly Buteogallus subtilis Nt No Yes No Wholly Buteogallus anthracinus Nt No Yes No Mainly Buteogallus urubitinga Nt No No No Mainly Buteogallus meridionalis Nt No No No Mainly Parabuteo unidnctus Nt No No No Mainly Busarellus nigricollis Nt No No No Mainly Geranoaetus melanoleucus Nt No No No Mainly Harpyhaliaetus solitarius Near threatened No Yes No Mainly Harpyhaliaetus coronatus Vulnerable No No No Mainly Buteo magnirostris Nt No No No Mainly Buteo leucorrhous Nt No Yes No Mainly Buteo ridgivayi Endangered Yes No Yes Wholly Buteo platypterus Nt No Yes No Mainly Buteo brachyurus Nt No No No Mainly Buteo galapagoensis Vulnerable Yes No Yes Wholly Buteo albicaudatus Nt No No No Mainly March 1998 Status of Tropical Raptors 15 Appendix 1. Continued. Species IUCN Conservation Status Endemic Forest Dependent Island Dependent Tropical Distribution Buteo poecilochrous Nt No No No Mainly Buteo albonotatus Nt No No No Mainly Morphnus guianensis Near threatened No Yes No Mainly Harpia harpyja Near threatened No Yes No Mainly Spizastur melanoleucus Near threatened No No No Mainly Spizaetus ornatus Nt No Yes No Mainly Spizaetus tyrannus Nt No Yes No Mainly Spizaetus isidori Near threatened No Yes No Mainly Daptrius ater Nt No Yes No Wholly Daptrius americanus Nt No Yes No Mainly Phalcoboenus carunculatus Nt Yes No No Wholly Polyborus plancus Nt No No No Mainly Milvago chimachima Nt No No No Mainly Herpetotheres cachinnans Nt No No No Mainly Micrastur ruficollis Nt No Yes No Mainly Micrastur gilvicollis Nt No Yes No Wholly Micrastur mirandollei Nt No Yes No Wholly Micrastur semitorquatus Nt No Yes No Mainly Micrastur buckleyi Nt Yes Yes No Wholly Falco femoralis Nt No No No Mainly Falco rufigularis Nt No Yes No Mainly Falco deiroleucus Near threatened No Yes No Mainly Afro tropics Pandion haliaetu.P Nt No No No Mainly Aviceda cuculoides Nt No Yes No Mainly Aviceda madagascariensis Nt Yes Yes Yes Mainly Pernis apivorus Nt No No No Mainly Macheiramphus alcinuP Nt No Yes No Mainly Elanus caeruleufi Nt No No No Mainly Chelictinia riocourii Nt No No No Wholly Milvus migrant Nt No No No Mainly Haliaeetus vocifer Nt No No No Mainly Haliaeetus vociferoides Critically endangered Yes No Yes Wholly Aegypius tracheliotus Nt No No No Mainly Aegypius occipitalis Nt No No No Mainly Necrosyrtes monachus Nt No No No Mainly Gyps rueppellii Nt No No No Wholly Gyps africanus Nt No No No Mainly Neophron percnopterus h Nt No No No Mainly Gypohierax angolensis Nt No Yes No Mainly Circaetus gallicus h Nt No No No Mainly Circaetus cinereus Nt No No No Mainly Circaetus fasdolatus Near threatened No Yes No Mainly Circaetus cinerascens Nt No Yes No Wholly Terathopius ecaudatus Nt No No No Mainly Dryotriorchis spectabilis Nt No Yes No Wholly Eutriorchis astur Critically endangered Yes Yes Yes Wholly Polyboroides typus Nt No No No Mainly Polyboroides radiatus Nt Yes Yes Yes Mainly Melierax poliopterus Nt No No No Wholly Melierax metabates Nt No Yes No Mainly 16 Bildstein et al. Vol. 32, No. 1 Appendix 1. Continued. Species IUCN Conservation Status Endemic Forest Dependent Island Dependent Tropical Distribution Melierax canorus Nt No No No Mainly Melierax gabar Nt No No No Mainly Kaupifalco monogrammicus Nt No Yes No Mainly Butastur ruftpennis Nt No No No Wholly Circus macrourud Near threatened No No No Mainly Circus pygargu d Nt No No No Mainly Circus aeruginosud Nt No No No Mainly Circus ranivorus Nt No No No Mainly Circus maillardi Near threatened Yes No Yes Mainly Accipiter tachiro Nt No Yes No Mainly Accipiter castanilius Nt No Yes No Wholly Accipiter brevipes Nt No No No Mainly Accipiter badiud Nt No No No Mainly Accipiter francesii Nt Yes Yes Yes Mainly Accipiter erythropus Nt No Yes No Wholly Accipiter minullus Nt No No No Mainly Accipiter madagascariensis Near threatened Yes Yes Yes Mainly Accipiter ovampensis Nt No Yes No Mainly Accipiter rufiventris Nt No No No Mainly Accipiter melanoleucus Nt No Yes No Mainly Accipiter henstii Near threatened Yes No Yes Wholly Urotriorchis macrourus Nt No No No Wholly Buteo oreophilus Nt No Yes No Mainly Buteo brachypterus Nt Yes Yes Yes Mainly Buteo auguralis Nt No Yes No Wholly Buteo augur Nt No No No Mainly Aquila pomarind Nt No No No Mainly Aquila rapad Nt No No No Mainly Aquila verreauxii Nt No No No Mainly Hieraaetus wahlbergi Nt No No No Mainly Hieraaetus fasciatud Nt No No No Mainly Hieraaetus spilogaster Nt No No No Mainly Hieraaetus pennatud Nt No No No Mainly Hieraaetus ayresii Nt No No No Mainly Hieraaetus bellicosus Nt No No No Mainly Spizaetus occipitalis Nt No No No Mainly Spizaetus africanus Nt No Yes No Wholly Spizaetus coronatus Nt No Yes No Mainly Sagittarius serpentarius Nt No No No Mainly Polihierax semitorquatus Nt No No No Mainly Falco naumannd Vulnerable No No No Mainly Falco newtoni Nt Yes No Yes Mainly Falco punctatus Endangered Yes Yes Yes Wholly Falco alopex Nt No No No Wholly Falco ardosiaceus Nt No No No Wholly Falco dickinsoni Nt No No No Mainly Falco zoniventris Nt Yes No Yes Mainly Falco chicquerd Near threatened No No No Mainly Falco concolor Nt No No No Mainly Falco cuvieri Nt No No No Mainly Falco biarmicus Nt No No No Mainly Falco fasciinucha Vulnerable No No No Mainly March 1998 Status of Tropical Raptors 17 Appendix 1 . Continued. Species IUCN Conservation Status Endemic Forest Dependent Island Dependent Tropical Distribution Indomalayan Pandion haliaetuP Nt No No No Mainly Aviceda jerdoni Near threatened No Yes No Mainly Aviceda subcristatcP Nt No No Yes Mainly Aviceda leuphotes Nt No Yes No Mainly Pernis ptilorhynchus Nt No No No Mainly Pernis celebensis Nt Yes Yes Yes Wholly Macheirhamphus aldnus h Nt No Yes No Mainly Elanus caeruleu Nt No No No Mainly Milvus migrant Nt No No No Mainly Haliastur induP* Nt No No No Mainly Haliaeetus leucogasteP Nt No No No Mainly Ichthyophaga humilis Near threatened No No No Mainly Ichthyophaga ichthyaetus Near threatened No Yes No Mainly Aegypius calvus Near threatened No No No Mainly Gyps indicus Near threatened No No No Mainly Gyps bengalensis Near threatened No No No Mainly Neophron percnopterus h Nt No No No Mainly Circaetus gallicup Nt No No No Mainly Spibmis cheela Nt No No No Mainly Spibmis kinabaluensis Data deficient Yes Yes Yes Wholly Spilomis minimus Near threatened Yes Yes Yes Wholly Spilornis elgini Near threatened Yes No Yes Wholly Butastur teesa Nt No No No Mainly Butastur liventer Near threatened No No No Wholly Butastur indicuP Nt No Yes No Mainly Circus macrouruP Near threatened No No No Mainly Circus melanoleucos Nt No No No Mainly Circus pygarguP Nt No No No Mainly Circus aeruginosus h Nt No No No Mainly Accipiter trivirgatus Nt No Yes No Mainly Accipiter griseiceps Nt Yes Yes Yes Wholly Accipiter badiuP Nt No No No Mainly Accipiter butleri Near threatened Yes Yes Yes Wholly Accipiter sobensiP Nt No Yes No Mainly Accipiter trinotatus Nt Yes Yes Yes Wholly Accipiter JasciatuP Nt No No No Mainly Accipiter novaeholbndiaP Nt No Yes No Mainly Accipiter virgatus Nt No Yes No Mainly Accipiter nanus Near threatened Yes Yes Yes Wholly Accipiter erythraucherP Nt Yes Yes Yes Wholly Accipiter rhodogaster Nt Yes Yes Yes Wholly Pithecophaga jefferyi Critically endangered Yes Yes Yes Wholly Ictinaetus malayensis Nt No Yes No Mainly Aquila pomarincP Nt No No No Mainly Aquila rapaP Nt No No No Mainly Hieraaetus JasciatuP Nt No No No Mainly Hieraaetus pennatuP Nt No No No Mainly Hieraaetus kienerii Nt No Yes No Mainly Spizaetus cirrhatus Nt No No No Mainly Spizaetus bartelsi Endangered Yes Yes Yes Wholly Spizaetus lanceolatus Near threatened Yes Yes Yes Wholly 18 Bildstein et al. Vol. 32, No. 1 Appendix 1. Continued. Species IUCN Conservation Status Endemic Forest Dependent Island Dependent Tropical Distribution Spizaetus philippensis Vulnerable Yes Yes Yes Wholly Spizaetus alboniger Nt No Yes No Wholly Spizaetus nanus Vulnerable No Yes No Wholly Polihierax insignis Near threatened No No No Wholly Microhierax caerulescens Nt No No No Mainly Microhierax latifrons Near threatened Yes No Yes Wholly Microhierax erythrogenys Nt Yes No Yes Wholly Falco naumannP Vulnerable No No No Mainly Falco araea Vulnerable Yes No Yes Wholly Falco moluccensis Nt Yes No Yes Wholly Falco chicquercP Near threatened No No No Mainly Falco severuA Nt No Yes No Mainly Falco jugger Nt No No No Mainly Australotropical Pandion haliaetuF Nt No No No Mainly Aviceda subcristatah Nt No No Yes Mainly Henicopernis longicauda Nt Yes Yes Yes Wholly Henicopernis infuscatus Near threatened Yes Yes Yes Wholly Macheirhamphus alcinu.P Nt No Yes No Mainly Elanus caeruleu.P Nt No No No Mainly Lophoictinia isura Vulnerable Yes No No Mainly Hamirostra melanosternon Nt Yes No No Mainly Milvus migrant 0 Nt No No No Mainly Haliastur indus h Nt No No No Mainly Haliaeetus leucogasteft Nt No No No Mainly Haliaeetus sanfordi Vulnerable Yes Yes Yes Wholly Butastur vndicuA' Nt No Yes No Mainly Accipiter soloensi.P Nt No Yes No Mainly Accipiter fasciatuP Nt No No No Mainly Accipiter mwaehollandiaA Nt No Yes No Mainly Accipiter melanochlamys Nt Yes Yes Yes Wholly Accipiter albogularis Nt Yes Yes Yes Wfiiolly Accipiter rufitorques Nt Yes No Yes Wholly Accipiter haplochrous Nt Yes Yes Yes WTiolly Accipiter henicogrammus Nt Yes Yes Yes Wfiiolly Accipiter luteoschistaceus Near threatened Yes Yes Yes Wholly Accipiter imitator Endangered Yes Yes Yes Wholly Accipiter poliocephalus Nt Yes No Yes Wholly Accipiter princeps Near threatened Yes Yes Yes Wholly Accipiter brachyurus Vulnerable Yes Yes Yes Wholly Accipiter erythraucherP Nt Yes Yes Yes Wholly Accipiter meyerianus Nt No Yes Yes Wholly Accipiter buergersi Data deficient Yes Yes Yes Wholly Accipiter radiatus Endangered Yes No No Mainly Accipiter doriae Near threatened Yes Yes Yes WTiolly Buteo solitarius Near threatened Yes No Yes Wholly Harpyopsis novaeguineae Vulnerable Yes Yes Yes Wholly Aquila gurneyi Near threatened Yes Yes Yes Wfiiolly Falco several Nt No Yes No Mainly a Not threatened. b Occurs in more than one tropical region. f. Raptor Res. 32(l):19-27 © 1998 The Raptor Research Foundation, Inc. CONSERVATION STATUS OF BIRDS OF PREY IN THE SOUTH AMERICAN TROPICS Richard O. Bierregaard, Jr. Department of Biology, University of North Carolina at Charlotte, 9201 University City Blvd., Charlotte, NC 28223-9201 U.S.A. Abstract. — The ICBP/IUCN Red Data Book lists as Threatened or Endangered five of the 82 species of Falconiformes and one of the 32 Strigiformes that breed in tropical South America. The White- collared Kite ( Leucopternis farbesi), an endemic of the Atlantic coastal forests of Brazil, can be added to this list. Because vast areas of South America, most notably its tropical forests, have been inadequately surveyed and few studies have been carried out on raptor species or assemblages, our knowledge of the distribution and natural history of many raptor species, especially owls and forest-dwelling Falconifor- mes, is rudimentary at best. Most needed for conservation purposes is information on population den- sities, distribution, and reproductive biology that will tell us how large populations are and how fast they can reproduce. For most species, these data are anecdotal or nonexistent. Although the majority of South America’s tropical raptors appear not to be globally threatened at present, more information is needed to confirm current assessments, recognize when species become threatened, and to move quickly and efficiently to address such threats. Key WORDS: Falconiformes ; Strigiformes ; endangered species ; habitat loss ; South America ; tropical forest, conser- vation. Estado de conservacion de las aves rapaces en los tropicos de America del Sur Resumen. — El Libro Rojo de la ICBP/IUCN incluye como amenazadas o en peligro a cinco de las 82 especies de Falconiformes y una de las 32 Strigiformes que se reproducen en los tropicos de America del Sur. Leucopternis farbesi, endemica de los bosques costeros del Atlantico en Brasil puede ser adicionada a esta lista. Vastas areas de Suramerica, mas especificamente los bosques tropicales han sido inadecu- adamente investigados y pocos estudios han sido realizados sobre las especies de aves rapaces o sobre sus ensamblajes, nuestro conocimiento de la distribucion e historia natural de muchas especies de aves rapaces, especialmente buhos y Falconiformes de bosque, es rudimentario. Para propositos de conser- vacion se requiere de informacion relacionada con densidades poblacionales, distribucion, biologia reproductiva, que pueda decirnos de que tamano son las poblaciones y que tan rapido se pueden reproducir. Para la mayoria de las especies estos datos son anecdoticos e inexistentes. A pesar de que la mayoria de las aves rapaces de America del Sur no estan globalmente amenazadas en el presente, se requiere de mas informacion para confirmar las evaluaciones actuales y poder reconocer cuando una especie se encuentra en peligro y poder actuar rapidamente para en forma eficiente abordar las amen- azas. [Traduccion de Cesar Marquez] The South American continent is renowned for its avian diversity; roughly 32% (3100 species) of the world’s avifauna is found there, on only 12% of the world’s land surface. Birds of prey are no exception. Eighty-four Falconiformes and 32 Stri- giformes are resident in South America (following the taxonomy in Hume [1991] and del Hoyo et al. [1994]). These totals represent 28% and 21%, re- spectively, of the world’s species in these two or- ders. Most (82) of the Falconiformes (Appendix 1) and all of the owls (Appendix 2) are tropical or are found primarily south of the tropics but have ranges that extend into the tropics (north of 30°S) . Not coincidentally, South America is also re- nowned for its Amazonian rainforests, home to many of the continent’s remarkable avifauna. Among the raptors, 52 Falconiformes and 25 owls are associated with some sort of tropical forest, from the lowlands to high-altitude cloud forests. Tropical forests around the world are being cleared at alarming rates (Whitmore 1997) , with an inevitable loss of primary habitat and fragmenta- 19 20 Bierregaard Vol. 32, No. 1 tion of what remains. These landscape-wide changes have profound implications for the con- servation of biological diversity and have piqued the attention of biologists from the conservation community (e.g., Laurance and Bierregaard 1997). Although the loss of tropical forests is the most conspicuous threat to tropical species, the conver- sion of wooded or open savanna to intensive agri- culture is also taking its toll by reducing available habitat. Birds of prey, as top-order predators with large home ranges and generally low population densi- ties, may be especially sensitive to such habitat loss and fragmentation. Due to this, certain taxa may be important indicators of habitat integrity. Along with their high visibility and general allure, they are ideal “flagship species” for the conservation of critically endangered habitats. In this paper, I review the aspects of the biology of tropical species that I feel need to be better un- derstood if we are to assess accurately their conser- vation status, and then I briefly summarize the ar- eas where our knowledge is weakest. Within this framework, I discuss the principal threats to raptor species in the South American tropics, review 1 the general trends that are apparent, discuss the status of eight species that seem to merit special atten- tion, and make recommendations for future re- search. Data Needed Effective conservation must be based on a sound understanding of a number of biological factors, principally but not exclusively: distribution and population density, taxonomy, breeding behavior and reproductive rates, and habitat requirements, or, conversely, sensitivity to habitat modification. Information on migratory habits and prey special- ization is important as well. Distribution and pop- ulation density together provide an estimate of population size, which is crucial in monitoring the vulnerability of a species. Although it is not often considered in conser- vation discussions, our understanding of taxonomy at the specific and subspecific level is indeed im- portant. In fact, we face many problems in defining biological and phylogenetic species and in decid- ing how to deal with distinct populations (Zink and McKitrick 1995). These questions have conserva- tion implications because conservation laws focus at only the species level. Among Falconiformes, subspecific differentia- tion has hardly been studied in the South Ameri- can tropics. Including subspecies, the 82 Falconi- form species in South America represent only 128 strictly South American taxa. Most species are monotypic and four species alone account for 22 of these taxa (Appendix 1). The constraint of only considering South American forms is clearly arti- ficial since many of the species occur in Central America, Mexico, and the Caribbean, where many are represented by various distinct populations. In- cluding Central American, Mexican, and Caribbe- an populations, the number of taxa represented by the 82 species climbs from 128 (1.6 taxa/species) to 170 (2.1 taxa/species) (Appendix 1). Hence, the much smaller Central American region contains a great deal of biological diversity and is of great con- servation concern, particularly given the greater annual rate of deforestation there when compared to tropical South America (Whitmore 1997) . Data on reproductive biology such as behavior, clutch size, frequency of nesting attempts, survi- vorship of nestlings, and recruitment into the adult population can provide an estimate of a species’ potential rate of population growth. Combined with estimates of population size or density, these data are important in assessing the potential resil- ience of a population. Finally, information on each species’ habitat re- quirements will also help us to assess threats to its persistence. Not all species that are typically found in any given habitat are behaviorally or ecologically restricted to it. A number of North American rap- tors have shown great behavioral plasticity and are nesting in suburban (e.g., Merlins [Falco columbar- ium, Sodhi et al. 1991], Swainson’s Hawks [Buteo swainsoni ; England et al. 1995], Cooper’s Hawks [Accipiter cooperii; Rosenfield et al. 1995]) or urban (Peregrine Falcon [Falco peregrinus\) settings. In Central America, a number of raptors nest in both primary forest as well as more disturbed landscape mosaics of agriculture, second growth, and pri- mary forest (Whitacre pers. comm.). Data in Hand A recent review (Bierregaard 1995) highlighted the gaps in our knowledge of the natural history of the Falconiformes in Meso- and South America. Overall, the species we know the least about are the small residents of primary forest, especially those in the genera Accipiter, Leucopternis, and Mi- crastur. Nests have not been described for 16 spe- cies (Table 1 ) , and fewer than five nests have been March 1998 Raptors in South American Tropics 21 Table 1. South American Falconiformes with very poorly known natural histories. Nest Undescribed Unknown Breeding Biology Scant or No Prey DATA a Leptodon forbesi X X 0 Harpagus diodon X + Circus buffoni + Accipiter poliogaster X X 0 Accipiter superciliosus + Accipiter collaris X X 0 Accipiter ventralis X + Accipiter erythronemius X X 0 Leucopternis schistacea X X + Leucopternis plumbea X X + Leucopternis princeps X X + Leucopternis melanops X X ~h Leucopternis kuhli X X + Leucopternis lacernulata X X + Leucopternis polionota X X + Buteogallus subtilis + Harpyhaliaetus solitarius + Harpyhaliaetus coronatus + Buteo albigula X + Buteo ventralis X + Phalcoboenus carunculatus X Phalcoboenus megalopterus X Phalocoboenus albogularis X + Milvago chimachima X Micrastur plumbeus X X + Micrastur gilvicollis X + Micrastur mirandollei X X + Micrastur buckleyi X 0 Spiziapteryx circumcinctus + a 0 = no prey data available, + = anecdotal data only. described for an additional 12 species. Breeding behavior is unknown for 20 species and is only anecdotally known for an additional 18 species (Ta- ble 1). No prey data have been reported for five species and only anecdotal data are available for an additional 21 species (Table 1). Why do we know so little about so many species? Most of the poorly-known species are small, secre- tive inhabitants of dense, continuous forest, where observation is extremely difficult. Many, such as the forest falcons of the genus Micrastur, are cavity nesters and rarely seen, and field identification of many species is not always easy (Howell and Whit- taker 1995). Given all these difficulties, it is not surprising that with the notable exception of Thiollay’s work in French Guiana (e.g., Thiollay 1989b) there have been no intensive studies of more than a handful of raptor species in tropical South American for- ests. The majority of what we know about most rap- tors is derived from data that were collected inci- dentally during other studies. The discovery of 38 nests of the Barred Forest-falcon ( Micrastur ruficol- lis; Thorstrom et al. 1991) stands out as one of the more remarkable studies. Threatening Factors The loss of primary, native habitats is the most serious threat to the flora and fauna of the world, and raptors in the tropics are no exception. The clearing of tropical rainforests occurred globally at an average rate of 0.81% per annum over the 1980s and shows no sign of abating (FAO 1993, cited by Whitmore 1997). Annual deforestation in Central America and the Caribbean (0.97%) was substan- tially above, while that in tropical South America 22 Bierregaard Vol. 32, No. 1 (0.71%) was slightly below the global decade av- erage (FAO 1993, cited by Whitmore 1997). The tropical forests of South America can be coarsely divided into four regions: the lowlands and foothills west of the Andes, which are biogeo- graphically an extension of the Central American forests (or vice versa; Gentry 1990); the higher el- evation forests of the Andean Cordillera; the Am- azon basin; and the Atlantic forests of Brazil, which extend south through parts of Paraguay and Uru- guay into northern Argentina (Misiones) . Deforestation occurs across these four regions with variable causes, rates, and extents. It is occur- ring most rapidly in northwestern Ecuador and Co- lombia, across a broad front along the northern and southern fringes of the Amazon basin, and throughout the Atlantic coastal forests of Brazil ap- proximately 500 km southeast of Amazonia. Cur- rently, the extent of deforestation runs from roughly 90% in the Atlantic forests (where defor- estation began over 300 yr ago) and Ecuador’s Pa- cific lowlands and foothill forests below 900 m (Dodson and Gentry 1991) to roughly 6% in Ama- zonia as of 1988, with 15% of the region affected by deforestation, isolation, and edge effects (Skole and Tucker 1993). While the conservation of tropical forest is cru- cial, the conversion of open country such as the llanos and cerrado to cattle grazing and intensive agriculture is also vital due to the detrimental ef- fects on a broad range of different raptors. Partic- ularly egregious are the large-scale soybean and sugar cane plantations of eastern and southern Brazil. Compounding the problems associated with implanting vast expanses of monoculture crops is the indiscriminant use of pesticides. Alvarez-Lopez and Rattan (1995) reviewed the status of 14 species of resident, diurnal birds of prey in the Cauca Valley of Colombia. Several spe- cies have declined as expected due to reduction in forest or wetland habitats, but three open-country species unexpectedly declined as well. This may have been due to the effects of heavy' cattle grazing and/or pesticide use. Hunting by humans is certainly a problem for larger species of raptors. The frequency with which radio-tagged Harpy Eagles ( Harpia harpyja ) have been shot is an indication of this (E. Alvarez pers, comm.). Thiollay (1989a) has suggested that many forest species may be sensitive to what would be considered mild hunting pressure, and already re- duced in the large tracts of unsurveyed rainforest where they have been hunted by gold miners, rubber tappers, and indigenous peoples for hun- dreds of years (Redford 1992). Overall Trends Because so little of lowland Amazonia has been clearcut and most raptor species are broadly dis- tributed across the region, there are no species in great peril in this area. In contrast, some species inhabiting the slopes of the Andean Cordillera and Brazil’s Atlantic coastal forests are more vulnerable because they have more limited ranges and these ecosystems have been extensively clearcut. Throughout tropical South America, we can be fairly confident that 53 species are not globally threatened at present. Some species are even in- creasing their range as forests are felled and con- verted to a mosaic of forest fragments, agricultural land and second growth. For example, the Black- chested Buzzard-eagle ( Geranoaetus melonoleucus ; Teixeira et al. 1988) and White-tailed Hawk ( Buteo albicaudatus ; Pacheco and Whitney 1995) are mov- ing northeast into the decimated Atlantic forests of Brazil. Del Hoyo et al. (1994) considered 11 species to be Near Threatened (Appendix 1). Three of these are far too poorly known to make an assessment of their status, and the remaining eight are included primarily because they are large (eg., Harpy and Crested Eagles [Morphnus guianensis ] ) , rare (Black Solitary Eagle [ Harpyhaliaelus solitarius] ) , or have restricted ranges (Mantled Hawk [ Leucopternis po- lionota ] ) . There are insufficient data to adequately assess the status of at least 14 species. Of these, three are considered Near Threatened by del Hoyo et al. (1994) and 11 are probably Not Threatened (Ap- pendix 1). As a group, owls are probably less vulnerable than diurnal birds of prey because there are no very large species with large home ranges, and their nocturnal habits make them much less vul- nerable to hunting. As with the Falconiformes, however, special attention should be paid to spe- cies of owls in the Andes and Pacific slope where endemic species with very restricted ranges occur, as well as those in the Atlantic coastal forests. Three species of owls (Black-capped Screech Owl [Otus atricapillus] , Rusty-barred Owl [Strix hy- lophila ] and Tawny-b r owe d Owl [ Pulsatrix koeniswal- diana ] ) , which are endemic to southern Brazil and portions of its Atlantic coastal forest, should be March 1998 Raptors in South American Tropics 23 watched simply because of the scale of habitat loss in this region. We can presume the Black-capped Screech Owl has small enough home range re- quirements that it should be able to hold out in relatively small remnant forest reserves, while the other two species are too poorly known to specu- late on their situation. Species Of Special Concern The Andean Condor ( Vultur gryphus) is Threat- ened over most of its range (Collar et al. 1992). Although it is still common in some areas, it is very rare in the northern Andes, where reintroduction programs are underway. The species is very poorly known biologically, but apparently has very low re- productive rates and consequently is very suscep- tible to persecution. The recently rediscovered White-collared Kite {Leucopternis forbesi; Teixeira et al. 1987), consid- ered a valid species by del Hoyo et al. (1994), may be the most endangered raptor in South America. While it is not uncommon in its range, the species is restricted to the extreme northeast of the Atlan- tic coastal forests of Brazil, an area undergoing devastating deforestation. The White-necked Hawk ( Leucopternis lacernula- ta) , another endemic to Atlantic forests of eastern Brazil, is also considered vulnerable/rare, with its population only partly protected. A recent report, however, provided interesting observations on hunting behavior, confirming that the species is an insect specialist that follows monkeys, army ants, and even lawn mowers to capture flushed insects (Martuscelli 1996). Such adaptability to the pres- ence of humans in the environment suggests that the species might persist as long as some habitat remains. The Gray-backed Hawk ( Leucopternis occidentalis ) is considered seriously endangered and in need of urgent action. It is endemic west of the Andes in Ecuador and adjacent northwest Peru, where de- forestation is proceeding rapidly, and it maintains sizeable populations at only two sites. Recent stud- ies by Vargas (1995) have substantially increased our knowledge and provided specific recommen- dations for protecting habitats for the species. The Crowned Solitary Eagle ( Harpy haliaetus co- ronatus) is mostly a bird of subtropical to temperate regions of South America, but it does extend into tropical zones in Brazil and Bolivia. Although it has been recorded over a broad geographic range, it apparently occurs at extremely low densities. Be- cause so little is known about possible threats to the species, it is considered only to be Vulnerable and is largely unprotected. It is in need of study. The Lesser Collared Forest-falcon ( Micrastur buckleyi ) is a very poorly known species recorded only from a small area in western Amazonia. It is considered rare, but its secretive habits and simi- larity to the sympatric Collared Forest-falcon ( M semitorquatus ) suggest that the bird may be more widespread. Collar et al. (1992) listed the species as insufficiently known and in need of further pro- tection, concluding, however, that it will probably prove relatively widespread and secure. I feel that it is a species for which we have insufficient data but which is probably not threatened. A western Brazilian record for the species has been retracted; the specimens on which this range extension was based are under study and may represent a new taxon (A. Whittaker pers. comm.). The Plumbeous Forest-falcon ( Micrastur plum- beus), endemic to the western slope of the Andes in Colombia and Ecuador, is Vulnerable, largely unprotected, and is in need of study. The species’ biology is practically unknown, but a study is un- derway in the Rio Nambi Nature Reserve in south- western Colombia with encouraging preliminary results (P. Salaman pers. comm.). The Long-whiskered Owl ( Xenoglaux loweryi) was recently discovered and described at only two lo- calities on the east slope of the Andean Cordillera. It is considered insufficiendy known and in need of further protection (Collar et al. 1992). Restricted Taxa A number of subspecies that are obligate forest dwellers with ranges occurring in areas undergoing rapid deforestation are worthy of careful attention. In Central America and the western slope of the Andes in Colombia and Ecuador, they are the Or- nate Hawk-eagle ( Spizaetus ornatus vicarius), Barred Forest-falcon ( Micrastur ruficollis interstes) , and Col- lared Forest-falcon ( Micrastur semitorquatus naso). In the forests of eastern Brazil and into northern Argentina, there is also a race of the Black Hawk- eagle ( Spizaetus tyrannus tyrannus ). A race of the Barred Forest-falcon (M r. olrogi) occurs only in the dry forests of northwest Argentina. New Research Since my earlier review (Bierregaard 1995), stud- ies have documented range expansions of several species. Black Hawk-eagles and Ornate Hawk-ea- 24 Bierregaard Vol. 32, No. 1 gles, for example, have been recorded in new parts of Paraguay (Williams 1995) and in the third larg- est tract of forest in Brazil’s Atlantic forest in Santa Catarina (Serra do Tabuleiro) where Harpy Eagles were also reported as recently as 1989 (Albuquer- que 1995). The Crested Eagle has been reported in Beni, Bolivia (Pearman 1994) and the Black- and-white Hawk-eagle ( Spizastur melanoleucus) was reported in a new area in Paraguay (Lowen et al. 1995). The Near Threatened Mantled Hawk has been discovered in an important tract of forest in southeastern Brazil (Forrester 1993, cited by Gon- zaga et al. 1995). In Venezuela, 30 breeding pairs of Harpy Eagles have been located (Anonymous 1996, and E. Al- varez pers. comm.). Preliminary results indicate that nests are much closer than would have been predicted by previous estimates of density and home range (Thiollay 1989a). Important data are being collected on local movement of juveniles in- strumented with satellite radiotransmitters. Two important studies on the Gray-backed Hawk and Plumbeous Forest-falcon (Vargas 1995, and P. Sal- aman pers. comm.), species whose biology was pre- viously virtually unknown (Bierregaard 1995), are providing data needed to monitor tropical raptors and design effective conservation action plans. Future Directions Almost any information about any Neotropical raptor, especially forest species, is useful. Studies of endemic species or subspecies in the Atlantic for- ests of Brazil are certainly of a very high priority. Similarly, raptor research in the Andes or Choco region of Colombia and adjacent Ecuador could provide a significant amount of information. In the central Amazonian lowlands, studies of any species in the genera Leucopternis, Acdpiter, or Micrastur are important. Literature Cited Albuquerque, J.L.B. 1995. Observations of rare raptors in southern Atlantic rainforest of Brazil. J. Field Orni- thol. 66:363-369. Alvarez-Lopez, H. and G.H. Kattan. 1995. Notes on the conservation status of resident diurnal raptors of the middle Cauca Valley, Colombia. Bird Conserv. Int. 5: 341-348. Anonymous. 1996. Harpy Eagle conservation program. The Peregrine Fund 1995 Annual Report. The Pere- grine Fund, Inc., Boise, ID U.S.A. Bierregaard, R.O., Jr. 1995. The biology and conser- vation status of Central and South American Falcon- iformes: a survey of current knowledge. Bird Conserv. Int. 5:325-340. Collar, N.J., L.P. Gonzaga, N. Krabbe, A. Madrono Nie- to, L.G. Naranjo, T.A. Parker III, and D.C. Wege. 1992. Threatened birds of the Americas. Smithsonian Inst. Press and International Council for Bird Pres- ervation, Cambridge, U.K. and Washington, DC U.S.A. del Hoyo, J., A. Elliot, and A. Sargatal [Eds.]. 1994. Handbook of birds of the world. Vol. II. New world vultures to guineafowl. Lynx Edicions, Barcelona, Spain. Dodson, C.H. and A.H. Gentry. 1991. Biological extinc- tions in western Ecuador. Ann. MO Bot. Gard. 78:273- 295. England, A.S., J.A. Estep and W.R. Holt. 1995. Nest- site selection and reproductive performance of urban- nesting Swainson’s Hawks in the Central Valley of Cal- ifornia. J. Raptor Res. 29:179-186. FAO. 1993. Forest resources assessment 1990. Tropical countries. FAO Forestry Paper 112. Forrester, B.C. 1993. Birding Brazil: a check-list and site guide. John Geddes, Irvine, CA U.S.A. Gentry, A.H. [Ed.]. 1990. Four neotropical rainforests. Yale Univ. Press, New Haven, CT U.S.A, Gonzaga, L.P, J.F. Pacheco, C. Bauer and G.D.A. Cas- TIGLIONI. 1995. An avifaunal survey of the vanishing montane Adantic forest of southern Bahia, Brazil. Bird Conserv. Int. 5:279-290. Howell, S.N.G. and A. Whittaker. 1995. Field identi- fication of Orange-breasted and Bat Falcons. Cotinga 4:36-43. Hume, R. 1991. Owls of the world. Running Press, Phil- adelphia, PA U.S.A. Laurance, W.F. and R.O. Bierregaard, Jr. [Eds.]. 1997 Tropical forest remnants: the ecology, management, and conservation of fragmented communities. Univ. Chicago Press, Chicago, IL U.S.A. Lowen, J.C., R.P. Clay, T.M. Brooks, E.Z. Esquivel, L. Bartrina, R. Barnes, S.H.M. Butchart and N.I. Etcheverry. 1995. Bird conservation in the Para- guayan Atlantic forest. Cotinga 4:58-64. Martuscelli, P. 1996. Hunting behaviour of the Man- tied Hawk Leucopternis polionota and the White-necked Hawk L. lacernulata in southeastern Brazil. Bull. Brit Ornithol. Club 116:114-1 16. Pacheco, J.F. and B.M. Whitney. 1995. Range exten- sions for some birds in northeastern Brazil. Bull. Brit. Ornithol. Club 1 15:157-163. Pearman, M. 1994. Neotropical notebook. Cotinga 1:26- 29. Redford, K. 1992. The empty forest. BioScience 42:412- 422. Rosenfield, R.N., J. Bielefeldt, J.L. Affeldt and D.J. Beckman. 1995. Nesting density, nest area reoccupan- cy, and monitoring implications for Cooper’s Hawks in Wisconsin./. Raptor Res. 29:1-4. March 1998 Raptors in South American Tropics 25 Skole, D.L. and C.J. Tucker. 1993. Tropical deforesta- tion and habitat fragmentation in the Amazon: satel- lite data from 1978 to 1988. Science 260:1905-1910. Sodhi, N.S., I.G. Warkentin and L.W. Oliphant. 1991. Hunting techniques and success rates of urban Mer- lins ( Falco columbarius) . J. Raptor Res. 25:127-131. Teixeira, D.M., J.B. Nacinovic and F.B. Pontual. 1987. Sobre a redescoberta de Leptodon forbesi (Swann, 1922) no nordeste do Brasil. Abstracts XIV Congresso Brasileiro de Zoologia in Jutz de Fora, Brasil 148. , J.B. Nacinovic and G. Luigi. 1988. Notes on some birds of northeastern Brazil (3). Bull. Brit. Or- nithol. Club 108:75—79. Thiollay, J.-M. 1989a. Area requirements for the con- servation of rainforest raptors and game birds in French Guiana. Conserv. Biol. 3:128-137. . 1989b. Censusing of diurnal raptors in a pri- mary rainforest: comparative methods and species de- tectability. J. Raptor Res. 23:72-84. Thorstrom, R.K., A.M. QuixchAn and G.M. Morales. 1991. Breeding biology of the Barred Forest Falcon (Micrastur ruficollis) . 1991. Pages 121-126 in D.F. Whi- tacre, W.A. Burnham and J.P. Jenny [Eds.], Maya Pro- ject: using raptors and other fauna as environmental indicators for design and management of protected areas and for building local capacity for conservation in Latin America. The Peregrine Fund, Inc., Boise, ID U.S.A. Vargas, H. 1995. Food habits, breeding biology, and sta- tus of the Gray-backed Hawk ( Leucopternis occidentahs ) in western Ecuador. M.S. thesis, Boise State Univ., Boi- se, ID U.S.A. Whitmore, T.C. 1997. Tropical forest disturbance, dis- appearance, and species loss. In W.F. Laurance and R.O. Bierregaard, Jr. [Eds.], Tropical forest remnants, ecology, management, and conservation of fragment- ed communities. Univ. Chicago Press, Chicago, IL U.S.A. Williams, R. 1995. Neotropical notebook. Cotinga 4:65— 68 . Zink, R.M. and M.C. McKitrick. 1995. The debate over species concepts and its implications for ornithology. Auk 112:701-719. Received 16 October 1996; accepted 8 October 1997 26 Bierregaard Vol. 32, No. 1 Appendix 1. Number of Falconiform taxa in Meso- Appendix 1. Continued. America and Tropical South America. Meso- Trop- Meso- Trop- World- Amer- ICAL World- Amer- ICAL wide ICAN S.A. wide ICAN S.A. Species Taxa Taxa Taxa Species Taxa Taxa Taxa Buteo nitidus 4 1 3 CATHARTIDAE Buteo magnirostris 12 6 6 Cathartes aura 4 1 2 Buteo leucorrhous 1 1 Cathartes burrovianus 2 2 2 Buteo brachyurus 2 1 1 Cathartes melambrotus 1 1 Buteo albigula 1 1 Coragyps atratus 3 1 2 Buteo albicaudatus 3 3 Sarcoramphus papa 1 1 Buteo galapagoensis 1 1 Vultur gryphus 1 1 Buteo polyosoma 2 1 ACCIPITRIDAE Buteo poedlochrous 1 ' 1 Buteo albonotatus 1 1 Leptodon cayanensis 2 2 Morphnus guianensis 1 1 Leptodon forbesi 1 1 Harpia harpyja 1 1 Chondrohierax uncinatus 3 2 1 Spizastur melanoleucus 1 1 Elanoides forficatus 2 1 1 Spizaetus tyrannus 2 2 Gampsonyx swainsonii 3 3 Spizaetus ornatus 2 2 Elanus leucurus 2 1 1 Oroaetus isidori 1 1 Rostrhamus sociabilis 3 2 1 Rostrhamus hamatus 1 1 FALCONIDAE Harpagus bidentatus 2 2 Daptrius ater 1 1 Harpagus diodon 1 1 Daptrius americanus 1 1 Ictinia plumbea 1 1 Phalcoboenus carunculatus 1 1 Circus buffoni 1 1 Phalcoboenus megalopterus 1 1 Circus cinereus 1 1 Caracara ( Polyborus ) plancus 4 2 2 Accipiter poliogaster 1 1 Milvago chimachima 2 2 Accipiter superciliosus 2 2 Milvago chimango 2 1 Accipiter collaris 1 1 Herpetotheres cachinnans 3 1 2 Accipiter ventralis 1 1 Micrastur ruficollis 6 1 5 Accipiter erythronemius 1 1 Micrastur plumbeus 1 1 Accipiter bicolor 4 1 3 Micrastur gilvicollis 1 1 Geranospiza caerulescens 6 2 4 Micrastur mirandollei 1 1 Leucopternis plumbea 1 1 Micrastur semitorquatus 2 2 Leucopternis schistacea 1 1 Micrastur buckleyi 1 1 Leucopternis princeps 1 1 Spiziapteryx circumcinctus 1 1 Leucopternis melanops 1 1 Falco sparverius 17 8 7 Leucopternis kuhli 1 1 Falco femoralis 3 1 2 Leucopternis lacernulata 1 1 Falco rufigularis 3 3 Leucopternis semiplumbea 1 1 Falco deiroleucus 1 1 Leucopternis albicollis 4 1 3 Falco peregrinus 19 1 1 Leucopternis occidentalis 1 1 82 species, total taxa: 190 42 128 Leucopternis polionota 1 1 Taxa/ species Buteogallus aequinoctialis 1 1 (tropical S.A. only) 1.6 Buteogallus anthracinus 3 2 1 Taxa/ species Buteogallus subtilis 3 2 1 (tropical M-A. & S.A.) 2.1 Buteogallus urubitinga 2 1 1 Buteogallus meridionalis 1 1 Parabuteo unicinctus 2 2 Busarellus nigricollis 2 2 Geranoaetus melanoleucus 2 2 Harpyhaliaetus solitarius 2 1 1 Harpyhaliaetus coronatus 1 1 March 1998 Raptors in South American Tropics Appendix 2. Number of Strigiformes in Meso-America and Tropical South America. Meso- Trop- World- Amer- ICAL wide I CAN S.A. Species Tax a Taxa Taxa TYTONIDAE Tyto alba 35 9 5 STRIGIDAE Otus albogularis 3 3 Otus atricapillus 3 3 Otus choliba 17 16 Otus colombianus 1 1 Otus guatemalae 17 10 7 Otus ingens 3 3 Otus koepckeae 1 1 Otus marshalli 1 1 Otus petersoni 1 1 Otus roboratus 1 1 Otus watsonii 6 6 Lophostrix cristata 3 2 1 Bubo virginianus 17 3 6 Pulsatrix koeniswaldiana 1 1 Pulsatrix melanota 2 2 Pulsatrix perspicillata 6 2 4 Glaucidium brasilianum 12 2 9 Glaucidium hardyi 1 1 Glaucidium jardinii 2 1 1 Glaucidium minutissimum 8 7 1 Glaucidium peruanum 1 1 Xenoglaux louteryi 1 1 Speotyto cunicularia 19 5 12 Ciccaba albitarsus 2 2 Ciccaba huhula 1 1 Ciccaba nigrolineata 1 1 Ciccaba virgata 8 3 5 Strix hylophila 1 1 Strix rufipes 3 1 Rhinopteryx clamator 3 3 Asio flammeus 10 2 4 Asio stygius 6 4 2 Aegolius harrisii 3 3 34 species, total taxa: 200 53 108 Taxa/ species (tropical S.A. only) 3.2 Taxa/ species (tropical M-A. and S.A.) 4.7 J. Raptor Res. 32(l):28-39 © 1998 The Raptor Research Foundation, Inc. RAPTORS IN THE EAST AFRICAN TROPICS AND WESTERN INDIAN OCEAN ISLANDS: STATE OF ECOLOGICAL KNOWLEDGE AND CONSERVATION STATUS Munir Virani 1 and Richard T. Watson The Peregrine Fund, Inc., 566 West Flying Hawk Lane, Boise, ID 83709 U.S.A. Abstract. — From our review of articles published on diurnal and nocturnal birds of prey occurring in Africa and the western Indian Ocean islands, we found most of the information on their breeding biology comes from subtropical southern Africa. The number of published papers from the east African tropics declined after 1980 while those from subtropical southern Africa increased. Based on our Knowl- edge Rating Scale (KRS), only 6.3% of breeding raptors in the east African tropics and 13.6% of the raptors of the Indian Ocean islands can be considered Well Known, while the majority, 60.8% in main- land east Africa and 72.7% in the Indian Ocean islands, are rated Unknown. Human-caused habitat alteration resulting from overgrazing by livestock and impacts of cultivation are the main threats facing raptors in the east African tropics, while clearing of forests through slash-and-burn methods is most important in the Indian Ocean islands. We describe conservation recommendations, list priority species for study, and list areas of ecological understanding that need to be improved. Key Words: Conservation ; east Africa ; ecology, western Indian Ocean ; islands', priorities', raptors; research. Aves rapaces en los tropicos del este de Africa y en islas al oeste del Oceano Indico: estado del cono- cimiento ecologico y de su conservacion Resumen. — De nuestra recopilacion de articulos publicados sobre aves rapaces diurnas y nocturnas que se encuentran en Africa y en las islas al oeste del Oceano Indico, encontramos que la mayoria de la informacion sobre aves rapaces residentes se origina en la region subtropical del sur de Africa. El numero de publicaciones provenientes de los tropicos del este de Africa, declino despues de 1980, mientras que aquellos del subtropico del sur del continente aumentaron. Con base en nuestra escala de valoracion del conocimiento solo el 6.3% de las especies residentes en los tropicos del este del Africa y el 13.6% de las aves rapaces de islas del Oceano Indico pueden ser consideradas como Bien Conocidas, mientras que la mayoria 60.8% del este de Africa y 72.2% de las islas al oeste del Oceano Indico fueron clasificadas como Desconocidas. La alteracion de habitats de origen antropico como resultado del so- brepastoreo y los impactos de la agricultura son las principales amenazas para las aves rapaces de los tropicos del este de Africa, mientras que la deforestation a partir de la quema y tala de bosques es la amenaza mas importante en las islas del Oceano Indico. Describimos recomendaciones de conservacion, identificamos las especies prioritarias para estudiar e identificamos las areas de entendimiento ecologico que deben ser mejoradas. [Traduction de Cesar Marquez] Raptors are an important tool to focus conser- vation strategies locally, regionally, and globally (Watson 1991). They can be used as “umbrella spe- cies” because their large home ranges and low nesting densities necessitate that any protected ar- eas encompassing viable populations or complete communities protect sufficient habitat and popu- lations of most, if not all, other species in the food 1 Also National Museums of Kenya, Nairobi and Leicester University, U.K. web below them (Thiollay 1992). Because of their top positions in terrestrial and aquatic food webs, raptors can be used as indicators of worldwide pol- lution by pesticides (Newton 1979). Changes in raptor distribution or abundance can serve as a measure of our impact on landscapes, even in re- mote areas (Reichholf 1974). Finally, raptors are popular and charismatic and several species have become significant “flagships” for increasing pub- lic interest and support of conservation programs (Burnham et al. 1992, Thiollay 1992). This paper combines our collective knowledge 28 March 1998 East African and Western Indian Ocean Raptors 29 of raptors in the east African tropics and western Indian Ocean islands as part of a worldwide review of their conservation and ecology. We chose to combine these two regions in one paper to facili- tate comparison of island with continental situa- tions in adjacent geographic areas. The east Afri- can tropics (including all countries between the Tropics of Cancer and Capricorn and east of about 20°E, Zanzibar and Pemba islands) and the western Indian Ocean islands (comprising Madagascar, Mauritius, Seychelles, Reunion, and Comores) sup- port roughly 23% of the world’s avifauna accord- ing to species listed in Martin (1987), Langrand (1990), Perlo (1995), and Zimmerman et al. (1996). These areas contain a broad spectrum of habitats, from montane through lowland, dry, moist, and riparian forests, to open woodland, sa- vanna, scrubland, desert, marsh, mangroves, and others. The east African tropics support 82 species of diurnal birds of prey and 23 nocturnal owls (Britton 1980, Brown et al. 1982, Fry et al. 1988). Three-quarters of these raptors, or roughly 20% of the world’s raptor species, breed within this area while the rest use the region as a migratory path- way. The Indian Ocean islands support 22 endemic raptor species (15 diurnal birds of prey and seven nocturnal owls), five other breeding species, and two Palearctic migrants. Although both these areas boast a tremendous diversity of birds that attracts the attention of bird watchers worldwide, their rap- tors are still poorly understood in many respects, ranging from basic biology to the factors affecting distribution and abundance. Conserving raptors, or using them as a tool to achieve broader conser- vation goals, is impossible without a sound under- standing of their ecological requirements. In this paper, we first give an overview of the state of ecological knowledge and conservation sta- tus of breeding raptors in the east African tropics and the western Indian Ocean islands, based on 546 published papers and recent lists of species in jeopardy. Second, we outline the main conserva- tion issues, contrast threats to raptors on the main- land with those on islands, summarize information on causes of threats, and cite particular examples of problems. Lastly, we discuss priorities for further studies and focus of conservation efforts. Methods We reviewed the literature in peer-reviewed journals on 105 species in Africa, 79 of which breed in tropical east Africa, and 22 that are endemic to the Indian Ocean islands. The review was divided into pre-1980 and post- 1980 material. Pre-1980 titles were obtained from the bib- liography of Brown et al. (1982) and Fry et al. (1988). Post-1980 titles were obtained from a keyword search from 1980 through the first quarter of 1996 on the RIOS- IS (Biological Abstracts Inc.) and RRTAC (Raptor Re- search and Technical Assistance Center, Boise, ID U.S.A ) computer databases. We assumed that the material ob- tained represented a significant proportion of all articles published in peer-reviewed journals, and that these sources accurately reflect ecological knowledge based on scientific studies. Ad-hoc accounts published in newslet- ters or magazines can provide additional knowledge but they were not included in this review as they are difficult to find and their significance to the understanding of a species’ ecology is difficult to assess. Based on a similar review of the status and knowledge of Central and South American raptors (Bierregaard 1995), we assigned scores to each species from 1 to 5 from our Knowledge Rating Scale (KRS) where 1 = an- ecdotal, speculative or unstudied; 2=1 nest or 1 year low sample size studies; 3 = multi-year, local area studies, <5 pairs; 4 = multi-year, >5 pairs; and 5 = >10 pairs, >10 years, or regional studies. This was repeated for four subject areas, factors affecting distribution and abun- dance, population trends, breeding biology, and feeding ecology. Scores for each subject were summed for each species (min. 4, max. 20), and species ranked by score. Species were then grouped into four categories of knowl- edge based on their scores: Unknown (score 4—6), Little Known (score 7-10), Known (score 11-13), and Well Known (score 14 and above). Although these categories were subjective, they simplified our review by separating species into roughly common levels of knowledge. To compare between subject areas, we calculated a “study index” (first line of Tables 1 and 2) by summing the KRS for all species within each subject, and converting the total to a percentage of the maximum possible if all spe- cies had KRS of 5. Species’ total scores were calculated by summing KRS for all species in all subject areas and expressed as a percentage of the maximum possible if all species had KRS of 5. The conservation status of each species was compiled from two sources (Collar et al 1994, Bennun and Njoroge 1996). Results and Discussion State of Ecological Knowledge and Conservation Status. We found 251 and 295 articles in our pre- 1980 and post-1980 literature searches, respective- ly. During the pre-1980 era, raptor research origi- nated almost equally from countries in the east Af- rican tropics (36%) and subtropical southern Af- rica (39%), with 14% originating from tropical west Africa and 12% from north and central Africa. From 1980 through the first quarter of 1996, the proportion of publications originating from sub- tropical southern Africa increased to 55% and de- creased to 25% from countries in the east African tropics, while they increased from the western In- dian Ocean islands (10%) and decreased from tropical west and north Africa (each 5%). The 30 Virani and Watson Vol. 32, No. 1 Table 1. Knowledge rating scores (KRS) for distribution and abundance, population biology, breeding biology, and feeding ecology of 79 raptor species breeding in the east African tropics. Species are listed in order of increasing KRS. KRS scores: 1 = anecdotal, speculative or unstudied; 2 = single nest and single year low sample size studies; 3 = multi-year local area studies; 4 = multi-year and >10 pairs; 5 = >10 pairs, >10 years or regional studies. Conser- vation status for species in the east African region (Bennun and Njoroge 1996): EN = Endangered, NT = Near Threatened, VU = Vulnerable, RR = Regional Responsibility. Global conservation status categories (Collar et al. 1994): EN = Endangered, NT = Near Threatened, VU = Vulnerable. Common Name Scientific Name Dis- TRIB. AND Abund. Pop- Breed- Feed- ULAT. INC ING Bio- Biol- Ecol- LOGY OGY OGY Proportion of Studies in Each Subject Area Species Total Conservation Status 11% 9% 26% 20% 13% Regional Global Buzzard, Mountain Buteo oreophilus 1 1 1 1 4 NT Buzzard, Red-necked Buteo augur alis 1 1 1 1 4 Eagle, Congo Serpent Dryotriorchis spectabilis 1 1 1 1 4 Snake-eagle, Southern Circaetus fasciolatus 1 1 1 1 4 NT NT Banded Falcon, Pygmy Polihierax semitorquatus 1 1 1 1 4 Hawk-eagle, Ayre’s Hieraaetus ayresii 1 1 1 1 4 VU Hawk-eagle, Cassin’s Spizaetus africanus 1 1 1 1 4 Hawk, Long-tailed Urotriorchis macrourus 1 1 1 1 4 Kestrel, Fox Falco alopex 1 1 1 1 4 Kestrel, Grey Falco ardosiaceus 1 1 1 1 4 Kite, African Swallow- Elanus ( Chelictinia ) riocourii 1 1 1 1 4 tailed Kite, Black Milvus migrans 1 1 1 1 4 Owlet, Albertine Glaucidium albertinum 1 1 1 1 4 VU-RR VU Owlet, Chestnut Barred Glaucidium capense caste- 1 1 1 1 4 neum Owlet, Etchecopar’s Glaucidium capense etcheco- 1 1 1 1 4 pari Owlet, Red-chested Glaucidium tephronotum 1 1 1 1 4 VU Owl, Fraser’s Eagle Bubo poensis 1 1 1 1 4 Owl, Congo-bay Phodilus prigoginei 1 1 1 1 4 VU Owl, Pemba Scops- Otus pembaensis 1 1 1 1 4 VU-RR NT Owl, Usambara Bubo vosseleri 1 1 1 1 4 VU-RR VU Secretary-bird Sagittarius serpentarius 1 1 1 1 4 Sparrowhawk, Black Accipiter melanoleucus 1 1 1 1 4 Buzzard, Augur Buteo augur 2 1 1 1 5 Buzzard, Lizard Kaupifalco monogrammicus 1 1 2 1 5 Snake-eagle, Brown Circaetus cinereus 1 1 2 1 5 NT Eagle, Western Banded Circaetus cinerascens 1 1 2 1 5 VU Hawk, African Cuckoo Aviceda cuculoides 1 1 2 1 5 NT Falcon, Taita Falco fasciinucha 1 1 2 1 5 VU VU Goshawk, Gabar Micronisus ( Melierax ) gabar 1 1 2 1 5 Goshawk, Little Banded Accipiter badius 1 1 2 1 5 (Shikra) Kestrel, Dickinson’s Falco dickinsoni 1 1 2 1 5 Owl, Abyssinian Long- Asio abyssinicus 1 1 1 2 5 eared Owl, Pearl-spotted Glaucidium perlatum 1 1 1 2 5 Owl, Pel’s Fishing Scotopelia peli 1 1 2 1 5 VU March 1998 East African and Western Indian Ocean Raptors 31 Table 1. Continued. Dis- TRIB. AND Abund. Pop- Breed- Feed- ULAT. ING ING Bio- Biol- Ecol- LOGY OGY OGY Proportion of Studies in Each Subject Area Species Total Conservation Status Common Name Scientific Name 11% 9% 26% 20% 13% Regional Global Owl, Sokoke Scops- Otus ireneae 1 2 1 1 5 VU-RR VU Owl, Spotted Eagle- Bubo africanus 1 1 1 2 5 Owl, African Wood- Ciccaba woodfordii 1 1 2 1 5 Sparrowhawk, Little Accipiter minullus 1 1 2 1 5 Vulture, Palm-nut Gypohierax angolensis 1 1 1 2 5 Vulture, White-headed Trigonoceps occipitalis 1 1 2 1 5 VU Snake-eagle, Black- Circaetus pectoralis 2 1 2 1 6 chested Hobby, African Falco cuvieri 1 1 2 2 6 Hawk, Bat Macheiramphus alcinus 1 1 2 2 6 NT Owl, Cape Eagle- Bubo capensis 2 1 1 2 6 VU Owl, White-faced Scops- Otus leucotis 1 1 2 2 6 Sparrowhawk, Ovambo Accipiter ovampensis 1 1 3 1 6 NT Vulture, Egyptian Neophron percnopterus 2 2 1 1 6 NT Vulture, African White- Gyps africanus 1 1 2 2 6 NT backed Hawk-eagle, African Hieraaetus spilogaster 1 1 4 1 7 Eagle, Long-crested Lophaetus occipitalis 2 1 2 2 7 Owl, Barred Glaucidium capense 4 1 1 1 7 Vulture, Hooded Necrosyrtes monachus 1 1 4 1 7 Vulture, Ruppell’s Griffon Gyps rueppellii 4 1 1 1 7 NT Falcon, Peregrine Falco peregrinus 1 1 3 3 8 Goshawk, Dark Chanting- Melierax metabates 1 1 3 3 8 Goshawk, Pale Chanting- Melierax canorus 1 1 3 3 8 Harrier-hawk, African Polyboroides typus 1 1 3 3 8 Falcon, Lanner Falco biarmicus 1 2 3 3 9 Owl, Giant Eagle Bubo lacteus 4 1 2 2 9 Falcon, Red-necked Falco chicquera 1 3 3 3 10 NT Goshawk, African Accipiter tachiro 1 3 3 3 10 Kestrel, Rock (Common) Falco tinnunculus 2 2 3 3 10 Kite, Black-winged Elanus caeruleus 1 1 4 4 10 Owl, Barn Tyto alba affinus 1 1 4 4 10 Owl, African Grass Tyto capensis 1 1 4 4 10 VU Owl, Scops Eurasian- Otus scops 1 1 4 4 10 Eagle, African Crowned Stephanoaetus coronatus 3 3 3 3 12 VU Eagle, Tawny Aquila rapax 3 3 3 3 12 Eagle, Wahlberg’s Aquila wahlbergi 2 2 4 4 12 Harrier, African Marsh Circus ranivorus 1 3 4 4 12 NT Sparrowhawk, Red- Accipiter rufiventris 3 3 3 3 12 NT breasted Eagle, Martial Polemaetus bellicosus 3 3 3 4 13 VU Kestrel, Greater Falco rupicoloides 3 3 4 3 13 NT Owl, African Marsh Asio capensis 4 1 4 4 13 NT Vulture, Lappet-faced Torgos tracheliotus 4 4 3 3 14 NT Bateleur Terathopius ecaudatus 4 4 4 4 16 Vulture, Bearded Gypaetus barbatus 4 4 4 4 16 EN 32 Virani and Watson Vol. 32, No. 1 Table 1. Continued. Dis- Pop- Breed- Feed- TRIB. ULAT. ING ING AND Bio- Biol- Ecol- Species Abund. LOGY OGY OGY Total Proportion of Studies Conservation in Each Subject Area Status Common Name Scientific Name 11% 9% 26% 20% 13% Regional Global Eagle, African Fish- Haliaeetus vocifer 4 4 5 4 17 Eagle, Black (Verreaux’s) Aquila verreauxii 4 4 5 5 18 Table 2. Knowledge rating scores (KRS) for distribution and abundance, population biology, breeding biology and feeding ecology of 22 endemic raptors of the western Indian Ocean islands. Species are listed in order of increasing KRS. KRS scores: 1 = anecdotal, speculative or unstudied; 2 = single nest and single year low sample size studies; 3 = multi-year local area studies; 4 = multi-year and >10 pairs; 5 = >10 pairs, >10 years or regional studies. Global conservation status (Collar et al. 1994): CR = Critical; EN = Endangered; VU = Vulnerable; NT = Near-threatened Distrib. AND POPULAT. BREEDING FEEDING SPECIES Abund. Biology Biology Biology Total Proportion of Studies in Each Subject Area CONSER- . VATION Status Common Name Scientific Name 18% 10% 16% 6% 15% Global Falcon, Madagascar Cuck- oo- Aviceda madagascariensis 1 1 1 1 4 Goshawk, Henst’s Accipiter henstii 1 1 1 1 4 NT Harrier-hawk, Madagascar Polyboroides radiatus 1 1 1 1 4 Harrier, Reunion Circus maillardi 1 1 1 1 4 NT Kestrel, Aldabra Falco newtoni aldabranus 1 1 1 1 4 Kestrel, Madagascar Falco newtoni 1 1 1 1 4 Owl, Grand Comoro Scops- Otus pauliani 1 1 1 1 4 CR Owl, Malagasy Scops- Otus rutilus 1 1 1 1 4 Owl, Seychelles Scops- Otus insularis 1 1 1 1 4 CR Owl, White-browed Ninox superciliaris 1 1 1 1 4 Sparrowhawk, Anjouan Accipiter francesii pusillus 1 1 1 1 4 Sparrowhawk, Frances’s Accipiter francesii 1 1 1 1 4 Sparrowhawk, Madagascar Accipiter madagascariensis 1 1 1 1 4 NT Serpent-eagle, Madagascar Eutriorchis astur 2 1 1 1 5 CR Owl, Anjouan Scops Otus rutilus capnodes 2 1 1 1 5 CR Owl, Madagascar Long- eared Asia madagascariensis 1 1 1 2 5 Kestrel, Banded Falco zoniiientris 2 1 2 2 7 Owl, Madagascar Red Tyto soumagnei 2 1 2 2 7 EN Buzzard, Madagascar Buteo brachypterus 3 1 4 4 12 Kestrel, Seychelles Falco araea 4 4 4 4 16 VU Fish-eagle, Madagascar Haliaeetus vociferoides 5 4 4 4 17 CR Kestrel, Mauritius Falco punctatus 5 5 5 4 19 EN March 1998 East African and Western Indian Ocean Raptors 33 post-1980 shift in the proportion of studies from east Africa to southern Africa may be partly due to the death in 1980 of Leslie Brown who contributed almost a quarter of all pre-1980 articles from east Africa. Of the four subject areas reviewed, breeding bi- ology (26%) was best studied in the African rap- tors, followed in descending order by feeding ecol- ogy, factors affecting distribution and abundance, and population biology (Table 1). Indian Ocean island species were mostly less well-studied than continental species, but scored higher in factors affecting distribution and abundance (18%), fol- lowed in descending order by breeding biology, population biology, and feeding ecology (Table 2). There was little difference in species’ total scores between continental species (13%, Table 1) and Indian Ocean island species (15%, Table 2), both of which indicate our knowledge is substantially in- complete. Breeding biology may be the most often studied aspect of raptor ecology on mainland Af- rica because there are many large species, often inhabiting open landscapes, that build large con- spicuous nests which are used year after year, thus facilitating observations on nesting behavior. The relatively small sizes of the Indian Ocean islands perhaps enable a better and more accurate survey of distribution and abundance of island raptors. The fact that many Indian Ocean island species are Threatened may also have contributed to an inter- est in distribution and abundance studies. Popu- lation trends and demographic parameters were least studied in both regions (Tables 1 and 2) pos- sibly being the most difficult kind of studies to complete. Monitoring and understanding popula- tion changes is probably the most important kind of knowledge needed for species that are in jeop- ardy. Of 79 species that breed in the east African trop- ics, five (6.3%) scored a total KRS >14 and could be said to be Well Known, eight (10.1%) scored from 11-13 and might be considered Known, 18 (22.8%) were rated Unknown in at least two sub- ject areas, and may be considered Little Known, while the majority' 48 (60.8%) were rated Un- known in all four subject areas (Table 1). Well Known species were large and charismatic, or lo- cally threatened such as the Lappet-faced Vulture ( Tm'gos tracheliotus), Bearded Vulture ( Gypaetus bar - batus ), Bateleur ( Terathopius ecaudatus ), African Fish-eagle ( Haliaeetus vocifer ) , and Black Eagle (Aq- uila verreauxit) (Table 1). Almost 40% of all the breeding raptors in Africa have been placed in a recently compiled east Af- rican list of birds in jeopardy (Table 1, Bennun and Njoroge 1996). This regional list, comprising spe- cies found in Kenya, Uganda, Tanzania, Rwanda, and Burundi, was compiled to give more detailed resolution of conservation priorities in these coun- tries and to set a regional agenda for research, monitoring, and conservation. Of the 30 breeding raptors listed in jeopardy in the east African re- gion, one species is listed as Endangered, 14 as Vul- nerable, and 15 as Near Threatened (Table 1). In contrast, only seven of the 79 breeding African spe- cies were listed as globally Threatened (Table 1, Collar et al. 1994). This substantial difference is because many raptors have continental ranges which make their global (but not necessarily re- gional) status more secure. There may also be a regional awareness that raptor populations are de- clining because once-common species are not seen as frequently as they were within memory, but data are lacking. It may also reflect an inherent prob- lem of categorizing a species’ status based on stan- dard criteria for which data are almost invariably lacking, especially on a global scale. Species considered globally Vulnerable or Near Threatened that rated as Unknown in our KRS and, therefore, should be targeted for study in- cluded Southern Banded Snake-eagle ( Circaetus fas- ciolatus ), Albertine Owlet ( Glaucidium albertinum), Congo Bay-owl ( Phodilus prigoginei ) , Pemba Scops- owl ( Otus pembaensis), Usambara Eagle-owl ( Bubo vosseleri ) , and Taita Falcon ( Falco fasciinucha ) (Ta- ble 1). The Sokoke Scops-owl ( Otus ireneae), previ- ously listed as Endangered (Collar and Stuart 1985) but now listed as Vulnerable (Collar et al. 1994, Bennun and Njoroge 1996), was rated Un- known, but a recent study has increased our knowl- edge of this bird to at least Known in all four sub- jects (Virani 1995a). Of 22 endemic Indian Ocean island species, only three (14%) rated as Well Known (Table 2). The majority of species (83.6%) were rated Little Known or Unknown (Species Total <10, Table 2). The Endangered Mauritius Kestrel ( Falco puncta- tus ) and the Vulnerable Seychelles Kestrel ( Falco araea) were considered Well Known. The Endan- gered Madagascar Fish-eagle ( Haliaeetus vocifero- ides ) was rated Known but recent studies have raised our knowledge level to Well Known (Watson et al. 1996, Rafanomezantsoa 1997, Watson 1997, Watson et al. 1997). Half of the endemic raptors 34 Vi RANT AND WATSON Vol, 32, No. 1 of the Indian Ocean islands were listed at some level of global jeopardy (Collar et al. 1994). Of these, five were listed as Critically Endangered, two as Endangered, one as Vulnerable, and three as Near Threatened (Table 2). Among the Unknown species that should be tar- geted for study were the Endangered Grand Com- oro Scops-owl ( Otus pauliani ) , Seychelles Scops-owl ( Otus insularis) , Anjouan Scops-owl ( Otus capnodes ) , and Anjouan Sparrowhawk ( Accipiter francesii pusil- lus ) (Table 2). Two notable recent rediscoveries, the Madagascar Serpent-eagle ( Eutriorchis astur) and Madagascar Red Owl (Tyto soumagnei ), have changed the species’ status from possibly extinct to extant (Halleax and Goodman 1994, Thorstrom et al. 1995); they were listed as Critically Endangered and Endangered, respectively (Collar et al. 1994). Studies on the Madagascar Red Owl in 1994-95 provided first data on breeding and diet (Thor- strom et al. 1997, Thorstrom and Rene de Roland 1997) so that the species could be rated as Known m two subject areas. Threats to Raptors. The main threats facing rap- tors in the east African tropics can be summarized as rapidly changing land-use patterns resulting from an increase in human population size (Sorley and Andersen 1994). The precise impacts of changing land uses on breeding and migrant rap- tors are species specific. In general, anthropogenic habitat alteration can influence raptor distribution and abundance through direct changes in habitat characteristics, such as availability of suitable nest- ing and perching sites (Enderson 1964, Marion and Ryder 1975, Stahlecker 1978, Janes 1984), or indirectly through changes in prey abundance and availability (Baker and Brooks 1981). In east Africa, we believe that four main factors affect raptor den- sity and diversity as a result of changes in habitat that accompany human land-use pressures: (1) overgrazing by domestic livestock resulting in the alteration of original vegetative cover that affects prey abundance and distribution (Sorley and An- derson 1994); (2) impact of cultivation that com- pletely replaces native vegetation and involves the use of pesticides and fertilizers potentially toxic to raptors and other vertebrates (Sorley and Ander- son 1994); (3) human hunting pressure that di- minishes prey populations (Sorley and Anderson 1994); and (4) direct human persecution (Reran 1981) with resulting changes in behavior (Knight et al. 1989), species abundance and diversity (Craighead and Mindell 1981), and indirect per- secution through secondary poisoning as in Kenya (Thomsett pers. comm.). Threats to raptors on the Indian Ocean islands are similarly related to increasing human popula- tions but they tend to be more extreme because of the limited area of islands, limited potential for dis- persion of both humans and raptors, and high de- gree of endemism among island raptor species. Threats to raptors depend on the human coloni- zation history, the faunal community, and the set of conservation problems that are unique to each island. For these reasons we will describe the con- servation issues in some detail for each island sep- arately. Madagascar, the largest of the Indian Ocean is- lands (587 000 km 2 ), supports 22 species of diurnal and nocturnal birds of prey. All except two of these breed on Madagascar. Eleven species, 10 of which are found in forest habitats and one in wetlands and coastal areas, are endemic to the island. Sub- fossil remains indicate that two Aquila species ex- isted on Madagascar during the Quaternary (Goodman and Rakotozafy 1995) and a Stephan- oaetus species has been identified from the Holo- cene (Goodman 1994), but neither genus now sur- vives on Madagascar. The island was relatively re- cently colonized by humans who are known to have been present since 1890 ± 90 B.R (MacPhee and Burney 1991). With a current human popu- lation of about 14 million, a population growth rate of 3.1% per year and per capita income around $230 per year (World Bank 1992), the ma- jor threats to raptors, their habitats, and most oth- er biota are driven by the subsistence needs of the human population. The greatest single threat to extant endemic raptors is from human induced habitat modification that renders forest or wetland habitat unsuitable for them. Clearing of eastern rainforest by slash-and-burn agriculture was esti- mated to occur at an average rate of 1 1 1 000 ha/ year from 1950-85 when 3.8 million ha (34% of original forest cover) remained (Green and Suss- man 1990). Most of the western dry deciduous for- ests have been replaced by relatively sterile savan- na, but deforestation continues in the west when deliberate grassland fires escape into forest areas. Loss of wetlands due to drainage and modification for production of rice continues and protection of remaining wetlands is unpopular because of the need to put new areas into production. Only one significant wetland exists within a protected area March 1998 East African and Western Indian Ocean Raptors 35 (Lake Tsimanampetsotsa, Langrand and Goodman 1995). Habitat conservation is the single most impor- tant priority in Madagascar, with greater emphasis on wetlands than has been achieved to date. Man- agement of wild populations of certain species, like the Madagascar Fish-eagle (Watson 1997), may also be needed in the near future, and captive breeding may be required not long after that. In the long term, however, sustainable agriculture and agro- forestry to provide local inhabitants with needed food, fuel and fiber, accompanied by a reduction of population growth, are among the prerequisites for effective conservation. Addressing social and economic needs of local peoples are needed if con- servation is to succeed. For example, ways to en- hance local traditional rules and taboos are being explored to achieve conservation of habitats in the absence of protected areas (Razandrizanakanirina and Watson 1997). Besides habitat loss, raptors are persecuted, es- pecially in the west where we know nestlings of Madagascar Fish-eagle have been taken for food or as pets and adults have been killed. Given the low population size and low productivity of this species (Watson et al. 1997), the loss of just a few nestlings and adults each year has a significant impact on the species’ survival (Watson 1997). Much perse- cution occurs out of ignorance about the value and rarity of species so increased public awareness is urgent. The Seychelles comprise three large islands, the largest of which is Mahe (145 km^) , and numerous others. It supports two endemic raptors, the forest dwelling Seychelles Kestrel with a world population of about 420 pairs on Mahe, Silhouette and through reintroduction on Praslin, as well as some satellite islands (Watson 1989), and Seychelles Scops-owl with a population of perhaps 80 pairs all in highland forests on Mahe. Threats to the kestrel were loss of suitable habitat on Praslin in the 19th and early 20th centuries, and human persecution. Public awareness, enforcement of bird protection laws, and banning of private ownership of firearms has inhibited illegal shooting (Watson 1981), while habitat recovery on Praslin has allowed the reintro- duction of the species to this island. Much of the present day highland forest habitat (secondary growth) of the Seychelles Scops-owl is incorporated in the Morne Seychellois National Park. A better understanding of the species’ food and nesting re- quirements is needed to assess the effects of log- ging outside the protected area. Monitoring of small island populations is clearly desirable be- cause of their susceptibility to extinction from ran- dom catastrophic events as well as human induced change. Four volcanic islands form the Comoro archi- pelago (Ngazidja or Grand Comoro, Ndzuani or Anjouan, Mwali or Moheli, and Maore or Mayotte) . Each island supports its own, mainly forest dwell- ing avifauna (Louette and Stevens 1992). Humans arrived perhaps 1000 years ago and deforestation on lowland coastal areas is complete. Most endem- ic species remain only in higher altitude forest and tree-heath on top of Mt. Karthala on Ngazidja and on Mwali. Little to no forest remains, even at high altitudes, on Ndzuani and Maore. The only re- maining endemic raptor species are the Grand Co- mores Scops-owl (Karthala Scops-owl) and An- jouan Scops-owl. The latter was rediscovered on Ndzuani in 1992 having not been recorded with certainty since about 1886 (Safford 1993). It is found only in a few remaining patches of upland forest (above 800 m) where the population is es- timated as probably 100-200 pairs. The species is endangered by forest clearance and capture for food by people. Safford (1993) recommended an investigation of the feasibility and consequences of the species’ introduction to neighboring island of Mwali, as this may be the species’ only hope for survival. The Grand Comores Scops-owl is con- fined to forest (virgin through degraded and small patches) between 1000-1900 m on Mt. Karthala on Ngazidja (Herremans et al. 1991). A 1989 study es- timated that there was perhaps more than 1000 pairs on the island assuming 10 000 ha of suitable habitat and a territory size of about 5 ha for each owl pair. Although encouraging, the data are ten- uous and there remains the long-term threat of habitat loss through forest fragmentation from fires, logging, and clearing for pasture. The spread of the introduced Indian Myna ( Acridotheres tristis ) as forest is opened is also a threat to endemic spe- cies from competition and other interspecific in- teractions. Various subspecies of Frances’s Sparrowhawk (A. francesii) are found on the Mascarene islands. A. francesii francesii occurs on Madagascar, but on Co- mores it is represented by three subspecies, none of which are present on Mwali. A. francesii griveaudi is found on Ngazidja but is not common (del Hoyo et al. 1994), A. francesii pusillus on Ndzuani was once common but is now probably close to extinc- 36 Virani and Watson Vol. 32, No. 1 tion due to hunting (Louette and Stevens 1992), and A. francesii brutus on Maore is common on the moist west side (del Hoyo et al. 1994). Rainforest protection wherever it occurs is the highest con- servation priority on the Comores. Remaining for- est is modified by firewood collecting, timber ex- traction, understory clearing for vegetables and crops, and cattle grazing. The creation of a forest reserve on Mt. Karthala would protect habitat for four endemic birds (Louette and Stevens 1992) and similar habitat protection may be feasible on Mwali where the human population is still low. There is little hope for establishing a reserve on Ndzuani due to population pressure. Control of hunting which may affect raptors, control of ani- mal introduction, and public education are also high priorities (Louette and Stevens 1992), as are investigations into introduction of the Anjouan Scops-owl to Moheli and protection of specific nest sites on Ndzuani. The island of Mauritius (1843 km 2 ) was covered with native forest when Europeans first began to settle on it near the end of the 17th century (Vaughan and Wiehe 1937), but destruction for lumber, firewood, and agricultural land was mas- sive. Habitat loss, hunting, and introduced species probably contributed to the demise of the Mauri- tian Owl ( Scops commersoni) along with the Dodo (Rap has cucullatus) and other fauna during the 18th century (Diamond 1987). Today only small, highly degraded vestiges of forest remain on rug- ged uplands, principally Black River Gorges (Cheke 1987, Jones and Owadally 1988). Loss of native forest, use of organochlorine pesticides, and predation by introduced species undoubtedly con- tributed to the decline of the Mauritius Kestrel which, in 1974, was reduced to just four wild birds confined to about 4000 ha of native forest (Cade and Jones 1993). Captive breeding and manage- ment in the wild has returned the population size to 56-68 breeding pairs and 222-286 individuals (end of 1993-94 breeding season) and there is rea- son to hope that the population may rise to 200- 250 pairs occupying degraded or even exotic for- est-like habitats (Jones et al. 1994). Reunion is a volcanic island of 2510 km 2 that lies 800 km east of Madagascar and 200 km west of Mauritius. Colonized and still run as a Department by the French, some endemic fauna are known to have become extinct from hunting and the effects of introduced species. The history of human de- struction of native vegetation parallels that of Mau- ritius, but was somewhat constrained by the very rugged topography. A proportionately larger area of native forest remains uncut, but nearly all of it is over 500 m and most over 1000 m elevation (Di- amond 1987), Currently 35% of the land surface is forest and woodland, 26% agricultural, and the remaining is listed as “other.” The only raptor known to breed is the Reunion Harrier ( Circus maillardi ) that is also found on Madagascar and Co- mores. Today, the species persists in exotic vege- tation. It suffered a severe post-war decline but has increased since banning DDT. Introduction of spe- cial protection in 1966 to reduce persecution seems ineffective (Thiollay pers. comm.). There is compelling evidence of a yet undescribed Scops Owl in the forests (Bretagnolle and Attie 1996) that needs investigation. Conservation and Study Recommendations. Conservation needs of island fauna and flora are generally more urgent than continental species, except where limited distributions on continents mimic the island situation (e.g., Sokoke Scops-owl, Virani 1995a, 1995b). Conservation needs for rap- tors on the Indian Ocean islands are variable de- pending on the island, but they have the following priorities in common: conservation of native hab- itats, prevention of exotic species introduction and control of existing introduced exotics, and preven- tion of direct exploitation, persecution, and inter- ference. Conservation needs for raptors in the east African tropics include protection of habitat es- pecially where species’ distributions are limited to small habitat patches, such as the Sokoke Scops- owl, control of environmental contaminants, and control of persecution through public education and awareness. In particular, because raptors typi- cally need very large ranges in which to forage and survive, current national parks may not be suffi- cient to maintain viable raptor populations. It is important, therefore, to develop a “living with wildlife” ethic among people that will allow coex- istence of raptors on land shared with people. Where conservation organizations and individual expertise are absent or poorly developed, such as in Madagascar and Comores, we recommend de- velopment of local capacity through training at technician through doctoral level, technical advis- ing, and provision of material and financial re- sources (Watson and Lewis 1994). For raptors rated as Little Known and at least Threatened (Tables 1 and 2), priority should be given to studies that collect the information need- March 1998 East African and Western Indian Ocean Raptors 37 ed to determine the best coarse of action to re- duce the probability of the species’ extinction. Such studies include, but are not limited to, the following: (1) measuring the species’ population size and distribution (e.g., Sokoke Scops-owl [Vir- ani 1995a, 1995b]) to include genetic distribution if subpopulations exist that are small and disjunct; (2) understanding the species’ nesting and forag- ing habitat needs (e.g., Madagascar Fish-eagle [Berkelman 1997] and Madagascar Red Owl); (3) monitoring population trends, productivity, surviv- al to maturity and adult survival (e.g., Madagascar Fish-eagle [Watson et al. 1997]); (4) understanding breeding and other behaviors that may affect the species’ ability to recover from low population size (e.g., Madagascar Fish-eagle [Watson et al. 1997]); (5) understanding movements including migration and natal dispersal (e.g., Madagascar Fish-eagle [Rafanomezantsoa 1997]); and (6) understanding the effects of human persecution and other an- thropogenic effects (e.g., Madagascar Fish-eagle [Watson 1997]. In addition to study of Little Known Threatened species, in a few cases that are Known, such as the Madagascar Fish-eagle, Grand Comores Scops-owl, or Anjouan Scops-owl, we en- dorse urgent protection of key breeding sites, and, as needed, preliminary use of wild population management to increase the population size and reduce the risk of extinction (Louette and Stevens 1992, Safford 1993, Watson et al. 1996, Watson 1997). Wetlands and the water-forest ecotone in Madagascar need focused conservation efforts if they are to survive (Watson 1997). Efforts to pro- tect significant blocks of remaining rainforest in Madagascar (e.g., Projet Masoala 1995, which used the area estimated for a viable population of Mad- agascar Serpent-eagles to help justify the size of protected area) are important for the survival of the Madagascar Serpent-eagle, Madagascar Red Owl, and other forest species and should be en- couraged and duplicated wherever possible. Acknowledgments We thank Alan Kemp, Leon Bennun and Andrew Jen- kins for their constructive review of this paper. Literature Cited Baker, J.A. and R.J. Brooks. 1981. Distribution patterns of raptors in relation to density of meadow voles. Con- dor 83:42-47. Bennun, L.A. and P. Njoroge. 1996. Birds to watch in east Africa: a preliminary Red Data list. Research Rep. 23, Centre for Biodiversity, National Museums of Ke- nya, Nairobi, Kenya. Berkelman, J. 1997. Habitat requirements and foraging ecology of the Madagascar Fish-eagle. Ph.D. disserta- tion, Virginia Polytechnic Institute and State Univ , Blacksburg, VA U.S.A. Bierregaard, R.O., Jr. 1995. The biology and conser- vation status of Central and South American Falcon- iformes: a survey of current knowledge. Bird Conserv Int. 5:325-340. Britton, P.L. [Ed.]. 1980. Birds of east Africa, their hab- itat, status and distribution. East Africa Natural His- tory Society, Nairobi, Kenya. Bretagnolle, V. and C. Attie. 1996. Comments on a possible new species of Scops Owl Otus sp. on Reun- ion. Bull. A.B.C. 3(1) :36. Brown, L.H., E.K. Urban and K. Newman. 1982. The birds of Africa. Vol. I. Academic Press, London, U K Burnham, W.A., D.F. Whitacre and J.P. Jenny. 1992. The Maya Project: use of raptors as tools for conser- vation and ecological monitoring of biological diver- sity. Pages 257-264 in B.-U. Meyburg and R.D. Chan- cellor [Eds.], Raptor conservation today. World Work- ing Group on Birds of Prey, Berlin, Germany. Cade, TJ- and C.J. Jones. 1993. Progress in restoration of the Mauritius Kestrel. Conserv. Biol. 7:169-175. Cheke, A.S. 1987. An ecological history of the Mascare- ne Islands, with particular reference to extinctions and introductions of land vertebrates. Pages 5-89 m A.W. Diamond [Ed.], Studies of Mascarene Island birds. Cambridge Univ. Press, Cambridge, U.K. Collar, N.J. and S.N. Stuart. 1985. Threatened birds of Africa and related islands: the ICBP/IUCN Red Data Book. International Council for Bird Preserva- tion and International Union for Conservation of Na- ture and Natural Resources, Cambridge, U.K. Collar, N.J., M.J. Crosby and A.J. Stattersfield. 1994. Birds to Watch 2. The world list of threatened birds Birdlife Conserv. Ser. No. 4. BirdLife International, Cambridge, U.K. Craighead, F.C., JR- and D.P. Mindell. 1981. Nesting raptors in western Wyoming, 1947 and 1975./. Wildl. Manage. 45:865-872. del Hoyo, J., A. Elliot and J. Sargatal [Eds.]. 1994. Handbook of birds of the world. Vol. 2. New World vultures to guineafowl. Lynx Edicions, Barcelona, Spain. Diamond, A. W. [Ed.]. 1987. Studies of Mascarene Island birds. Cambridge Univ. Press, Cambridge, U.K. Endersqn, J.H. 1964. A study of the Prairie Falcon in the central Rocky Mountain Region. Auk 81:332-352 Fry, C.H., S. Keith and E.K. Urban [Eds.]. 1988. The birds of Africa. Vol. III. Academic Press, London, U K. Green, G.M. and R.W. Sussman. 1990. Deforestation his- tory of the eastern rainforests of Madagascar from sat- ellite images. Science 248:212-215. Goodman, S.M. 1994. Description of a new species of 38 Virani and Watson Vol. 32, No. 1 subfossil eagle from Madagascar: Stephanoaetus (Aves: Falconiformes) from the deposits of Ampasambazim- ba. Proc. Biol. Soc. Wash. 107:421-428. and L.M.A. Rakotozafy. 1995. Evidence for the existence of two species of Aquila on Madagascar dur- ing the Quaternary. Geobios 28:241-246. Halleax, D. and S. Goodman. 1994. Rediscovery of the Madagascar Red Owl Tyto soumagnei (Grandidier 1878). Bird Conserv. Int. 4:305-311. Herremans, M., M. Louette and J. Stevens. 1991. Con- servation status and vocal and morphological descrip- tion of the Grand Comoro Scops Owl Otus pauliani Benson 1960. Bird Conserv. Int. 1:123-133. Janes, S.W. 1984. Influences of territory composition and interspecific competition on Red-tailed Hawk re- productive success. Ecology 65:862-870. Jones, C.G. and A.W. Owad al ly. 1988. The life histories and conservation of the Mauritius Kestrel Falco punc- tatus (Temminck 1823), Pink Pigeon Columba mayeri (Prevost 1843) and Echo Parakeet Psittacula eques (Boddart 1783). Proc. R. Soc. Arts Sci. Mauritius 5:79- 130. , W. Heck, R.E. Lewis, Y. Mungroo, G. Slade and T. Cade. 1994. The restoration of the Mauritius Kes- trel Falco punctatus population. Ibis 137:S173— S180. Keran, D. 1981. The incidence of man-caused and nat- ural mortalities to raptors. Raptor Res. 15:108-112. Knight, R.L., D.E. Anderson, M.J. Bechard and N.V. Marr. 1989. Geographic variation in nest-defence be- haviour of the Red-tailed Hawk Buteo jamaicensis. Ibis 131:22-26. Langrand, O. 1990. Guide to the birds of Madagascar. Yale Univ. Press, New Haven, CT U.S.A. and S. Goodman. 1995. Monitoring Madagas- car’s ecosystems: a look at the past, present, and fu- ture of its wetlands. Pages 203-214 in T.B. Herman, S. Bondrup-Nielson, J.H. Willison and N. Munro [Eds.], Ecosystem monitoring and protected areas. Science and Management of Protected Areas Associ- ation, Nova Scotia, Canada. Louette, M. and J. Stevens. 1992. Conserving the en- demic birds on the Comoro Islands, I: general consid- erations on survival prospects. Bird Conserv. Int. 2:61— 80. MacPhee, R.D.E. and D.A. Burney. 1991. Dating of modified femora of extinct dwarf hippopotamus from southern Madagascar: implications for constraining human colonization and vertebrate extinction events. J. Arch. Sci. 18:695-706. Marion, W.R. and R.A. Ryder. 1975. Perch-site prefer- ence of four diurnal raptors in northeastern Colora- do. Condor 77:350-352. Martin, B.P. 1987. World birds. Guinness Superlatives, Enfield, U.K. Newton, I. 1979. Population ecology of raptors. T. & A.D. Poyser, London, U.K. Perlo, B.V. 1995. Birds of eastern Africa. Harper Col- lins, London, U.K. Projet Masoala. 1995. Proposition des limites du Parc National Masoala. CARE International Madagascar, Wildlife Conservation Society, The Peregrine Fund, Antananarivo, Madagascar. Rafanomezantsoa, S. 1997. Behavior and natal dispersal of fledgling Madagascar Fish-eagles. Pages 403-412 in R.T. Watson [Ed.], Madagascar wetlands conservation project. Progress report III, 1995-1996. The Pere- grine Fund, Boise, ID U.S.A. Razandrizanakanirina, D. and R.T. Watson. 1997. The process of developing the wetland community-based conservation project: 1997-1998. Pages 33-44 in R.T. Watson [Ed.], Madagascar wetlands conservation proj- ect. Progress report III, 1995-1996. The Peregrine Fund, Boise, ID U.S.A, Reichholf, J. 1974. Artenreichtum, Haufigkeit und Div- ersitat derGreifvogel in einigen Gebieten yon Suda- merika. J. Ornithol. 115:381-397. Safford, R.J. 1993. Rediscovery, taxonomy and conser- vation of the Anjouan Scops Owl Otus capnodes (Gur- ney 1889). Bird Conserv. Int. 3:57-74. Sorley, C.S. and D.E. Andersen. 1994. Raptor abun- dance in southcentral Kenya in relation to land-use patterns. Afr. J. Ecol. 32:30-38. Stahlecker, D.W. 1978. Effect of a new transmission line on wintering birds of prey. Condor 80:444-446. Thiollay, J-M. 1992. A world review of tropical forest raptors. Current trends, research objectives and con- servation strategy. Pages 231-239 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation to- day. World Working Group on Birds of Prey, Berlin, Germany. Thorstrom, R., R.T. Watson, B. Damary, F. Toto, M. Baba and V. Baba. 1995. Repeated sightings and first capture of a live Madagascar Serpent-eagle Eutriorchis astur. Bull. B.O.C. 115:40—45. Thorstrom, R. and L. A. Rene de Roland. 1997. First nest record and nesting behaviour of the Madagascar Red Owl ( Tyto soumagnei). Ostrich 68:42-43. Thorstrom, R., J. Hart and R.T. Watson. 1997. New record, ranging behaviour, vocalization and food of the Madagascar Red Owl (Tyto soumagnei). Ibis 139: 477-481. Vaughan, R.E. and P.O. Wiehe, 1937. Studies on the vegetation of Mauritius, I. A preliminary survey of plant communities./. Ecol. 25:289-343. Virani, M.Z.A. 1995a. The ecology of the endangered Sokoke Scops Owl Otus ireneae. M.S. thesis, Univ. Leicester, U.K, . 1995b. Sokoke Scops Owl in Tanzania. Swara: May-June. Watson, J. 1981. Population ecology, food and conser- vation of the Seychelles Kestrel ( Falco araea) on Mahe. Ph.D. dissertation, Univ. Aberdeen, U.K. . 1989. Successful translocation of the endemic March 1998 East African and Western Indian Ocean Raptors 39 Seychelles Kestrel Falco araea to Praslin. Pages 363- 367 in Meyburg B.-U. and R.D. Chancellor, [Eds.], Raptors in the modern world. World Working Group on Birds of Prey, Berlin, Germany. Watson, R.T. 1991. Using birds of prey as an environ- mental conservation tool: The Peregrine Fund’s World Programme. Environ . Conserv. 18:269-270. , 1997. Madagascar Fish-eagle conservation. Pages 45-49 in R.T. Watson [Ed.], Madagascar wetlands con- servation project. Progress report III, 1995-1996. The Peregrine Fund, Boise, ID U.S.A. and R. Lewis. 1994. Raptor studies in Madagas- car’s rainforest. Pages 283—290 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation today. World Working Group on Birds of Prey, Berlin, Ger- many. , S. Thomsett, D. O’Daniel and R. Lewis. 1996. Breeding, growth, development, and management of the Madagascar Fish-eagle ( Haliaeetus vodferoides) J Raptor Res. 30:21-27. , S. Razafindramanana, R. Thorstrom and S. Ra- fanomezantsoa. 1997. Breeding biology, productivi- ty, siblicide and conservation of the Madagascar Fish- eagle. Pages 289-306 in R.T. Watson [Ed.], Madagas- car wetlands conservation project. Progress report III, 1995-1996. The Peregrine Fund, Boise, ID U.S.A. World Bank. 1992. World development report 1992: de- velopment and the environment. Oxford Univ. Press, New York, NY U.S.A. Zimmerman, D.A., D.A. Turner and D-J. Pearsin. 1996. Birds of Kenya and northern Tanzania. Princeton Univ. Press, Princeton, NJ U.S.A. Received 16 October 1996; accepted 12 November 1997 J. Raptor Res. 32(1) :40-55 © 1998 The Raptor Research Foundation, Inc. CURRENT STATUS AND CONSERVATION OF FALCONIFORMES IN TROPICAL ASIA Jean-Marc Thiollay Laboratoire d Ecologie, E.N.S., 46, rue d’Ulm 75230 Paris Cedex 05, France Abstract. — I reviewed the distribution, abundance, habitat selection, and population trends of 69 di- urnal raptors found in 13 subregions from India to Indochina, Sumatra, and Java. At least four species are Endangered or Vulnerable and 19 are at lower risk or have insufficient data to determine their status. Among 49 breeding species, 33 have a world range mostly within South Asia, half with high conservation value scores. Others have small (six species) or substantial (10 species) populations but a much wider distribution. Of all 69 taxa, eight are mature forest-interior species, six are associated with gaps or disturbance, 14 are woodland, edge, or upper canopy dwellers, 14 are associated with grasslands or cultivation, and seven are river or wedand species. Most wintering migrants (20) are nonforest species. There is evidence of largescale population declines on the continent of Asia, as well as insular syndrome involving increased density and wider habitat breadth on islands. The most important threats to diurnal raptors are (1) habitat loss, especially of lowland rainforest to cultivation, (2) habitat degradation, mainly from logging, and (3) forest fragmentation. Few species are successfully adapting to new man-made habitats and none are from the forest-interior guild. Main conservation priorities include surveys and ecological studies, extension and enforcement of protected areas, and sustainable logging practices. Key Words: raptor populations:, South Asia; distribution; conservation. Estado actual y conservation de falconiformes en el Asia tropical Resumen. — Resumi la distribution, abundancia, selection de habitat, y tendencias poblacionales de 69 aves rapaces diurnas de 13 subregiones desde India hasta Indochina, Sumatra, yjava. Por lo menos 4 especies estan amenazadas o son vulnerables, 19 estan en menos riesgo o no existe information sufi- ciente para determinar su estado. Entre las 49 especies, 33 tienen un area de distribution dentro del Sur de Asia, la mitad de estas tienen un valor de conservation alto. Otras tienen poblaciones reducidas (6 especies) o substanciales (10 especies) con distribuciones mas amplias. De todos los 69 taxones, 8 son especies del interior de bosques maduros, 6 estan asociados a zonas abiertas o disturbadas, 14 son especies de bosque, borde o dosel, 14 estan asociados a pastizales o cultivos, y 7 son especies de rfos o humedales. Casi todas las migratorias (20) no son especies de bosque. Existen evidencias sobre la di- minution poblacional de las especies a gran escala en el continente Asiatico, como tambien sobre el sindrome insular que incluye el aumento de densidades y espacios entre habitats en islas. Las amenazas mas importantes para las aves rapaces diurnas son: (1) Perdida de habitat, especialmente de los bosques de lluvia de tierras bajas convertidas a cultivos, (2) degradation del habitat a partir de actividades forestales, y (3) fragmentation del bosque. Pocas especies se estan adaptando exitosamente a los nuevos habitats creados por el hombre. Las prioridades de conservation, incluyen investigation es y estudios ecologicos, apoyo y extension de areas protegidas y practicas forestales sostenibles. [Traduction de Cesar Marquez] Several Asian countries have the densest and fas- test growing human populations in the world, as well as the fastest growing economies and stan- dards of living. As a result, there is dramatic pres- sure on natural habitats, resources, and biodiver- sity of this area. Raptors are good indicators of environmental changes because they are top predators, they in- clude species with a variety of specific habitat re- quirements, prey selection and degrees of toler- ance to disturbance, and their species numbers, at least in some areas, are correlated with the total bird species richness (Thiollay 1997). My aim here is to review the composition, distribution, popula- tion levels, and conservation priorities of diurnal raptors in Southeast Asia in the early 1990s. Most information about the geographical distribution of raptor species is scattered in the literature and 40 March 1998 Status of Raptors in Tropical Asia 41 much less is known about their habitat selection and their actual abundances. These last critical components of their status are rapidly changing and may be different today than they were report- ed in regional syntheses based on information that is decades old. To update and complement pub- lished data, I shall rely primarily on the results of numerous personal raptor surveys and field studies carried out in 27 states or provinces in eight coun- tries during the last 20 yr. Study Area The area under consideration extends from India to Indochina, south of the Himalayas and China, and down to the Greater Sundas, west of the Wallace Line (Fig. 1). This is a faunistically homogeneous, 7 million-km 2 re- gion, that now has a human population of >1.4 billion people. It is divided into three broad biogeographic zones: Indian (India to Burma), Indochinese (Thailand to Vietnam), and Sundan (Malaysia to Java). In terms of its raptors, the area can be divided more precisely into 13 subregions (Table 1). The main vegetation type is a rich, closed-canopy, moist forest with trees approximately 50 m in height (Whitmore 1985). At best, only 42% of the original forest remains (global estimate from Collins et al. [1991], as- suming a constant deforestation rate since the 1980s). This estimate includes evergreen to semi-deciduous, low- land and montane forests that have often already been logged or degraded, where annual rainfall is 2-4 m or more and no month receives under 60 mm of precipi- tation. At higher latitudes, marked dry seasons (^3 months with <60 mm of rainfall) result in drier, more deciduous monsoon forests grading into lower wood- lands or even subdesert scrubland in north western In- dia. Lowland swamp forests are locally extensive in Bor- neo and Sumatra. Mangroves ( Avicennia , Rhizophora) are widespread along low coastlines and estuaries and cover over 4000 km 2 in the Sundarbans of Bangladesh alone Former coastal forests have been replaced by coconut 42 Thiollay Vol. 32, No. 1 Table 1. Total size of study areas, remaining forested areas (1980-88), and existing protected areas in 1000s of km 2 (from Collins et al. 1991). Subregions Land Area Rain Forest Monsoon Forest 1 Mangrove and Swamp Forest Protected Areas 2 Northern India a 1805.9 0.9 98.1 1.2 106.0 Peninsular India 944.6 20.0 18.4 0.1 16.6 Northeastern India b 640.4 46.7 38.8 4.5 9.3 Sri Lanka 64.7 1.4 10.8 7.2 Andamans-Nicobars 6.3 2.6 3.6 0.6 0.6 Burma (Myanmar) 657.7 220.3 88.4 3.0 7.0 Thailand + Cambodia 688.3 127.4 79.2 13.4 66.9 Laos + Vietnam 556.1 134.3 45.3 1.6 8.2 Peninsular Malaysia 0 330.3 64.5 5.2 6.2 Northern Borneo 203.9 113.5 21.7 8.1 Kalimantan 532.1 323.6 73.7 29.7 Sumatra d 472.6 155.3 75.3 45.0 Java + Bali 138.6 12.8 9.2 3.7 Total 7041 1223 382 209 314 1 Includes both lowland and montane forest, and from intact to highly degraded stands. 2 Includes also nonforest areas and low protection status areas such as protection forests or hunting reserves. a North of 20°N, excluding Himalayan and Tibetan ranges. b Includes Bangladesh. ' Includes Peninsular Thailand, Tenasserim and Singapore. d Includes outlying islands. plantations. Upper montane areas above tree line are in- significant in size. Wetlands are mostly associated with rivers and to a less- er extent with marshes, peat swamps, coastal lagoons, many artificial lakes, and vast areas of temporarily inun- dated ricefields. Permanent agriculture is often intensive over huge, densely-inhabited areas. Nevertheless, a sub- stantial density of trees, patches of grassland for catde grazing, and fallow areas with low second growth are re- tained. In more remote and forested areas on slopes, shifting cultivation has resulted in various successional stages of secondary growth. Tree plantations are often extensive. They range from the diversified, forest-like agroforests of Indonesia to the vast and uniform indus- trial plantations of fast growing exotic trees grown for fuel or pulpwood ( Eucalyptus , Gmelina, and Acacia) , lum- ber ( Tectona , Pinus , and Dipterocarpaceae) , fruit (Mango), and especially rubber ( Hevea ) and oil ( Elaeis ). Together with open cultivated woodlands and home gardens, they currently cover much larger areas than natural forests. Urban, industrial, and recreational areas are increasing everywhere with the increasing human population and economic development. Methods Only resident and regular wintering migrant diurnal raptors were considered, while vagrants and transient mi- grants were ignored. I surveyed raptor populations over extensive areas from 1975-95, mostly in Nepal, northern and southern India, Andamans, Sri Lanka, Malaysia, Bor- neo, Sumatra, Java, Laos, and Vietnam. In each region, two sets of methods were used to assess the relative abun- dance and habitat selection of raptors. First, extensive transect counts were made along roads, tracks, and forest trails, either by car with numerous stops or on foot. The second method was an adaptation of the point count method (Thiollay 1989, 1995) where 1-km 2 sample quad- rats, predominantly covered by a single habitat type, were censused for 4 hr from vantage lookouts during morning periods (0900-1200 H) when most raptors were active The results were expressed in mean number of individ- uals, respectively, per km of transect or per 1-km 2 sample plot. Differences between these abundance indices (with- in, but not between species, because of unequal detect- abilities) were tested among habitats and study areas us- ing Kruskal-Wallis one-way AN OVA, followed by a non- parametric multiple pair-wise comparison procedure based on two-tailed Mann-Whitney 6-tests (Sokal and Rohlf 1981). Habitat types, where each individual raptor was record- ed in cumulative random samples, were used to assess specific habitat selection compared to habitat availability using the method of Neu et al. (1974). Thus, species were classified into broad categories of habitat prefer- ence, illustrating their distributional trend along the hab- itat gradient from primary forest to deforested habitats and their ability to withstand increasing deforestation. Random records were also used to measure specific hab- itat niche breadth, (3 = S(p ; 2 ) -1 , where p ; was the pro- portion of birds sighted in the i th habitat category. Two- tailed sign tests were used for interspecifc comparisons of habitat niches. March 1998 Status of Raptors in Tropical Asia 43 Table 2. Component scores of the conservation index attributed to tropical Asian raptors. The total index was the sum of the species rank value in each of three parameters. Conservation Parameters Rank Range Area Main Habitat Population Trend 0 >75% of S. Asia Open and cultivated habitats Stable or increasing 1 50-75% of S. Asia Forest edges or woodlands Probable but undocumented decline 2 25-49% of S. Asia Disturbed or secondary forests Moderate or locally documented decline 3 <25% of S. Asia Primary forest interior Significant and widespread decrease The following habitat categories were defined: WET, all wetlands from marshes, lakes and rivers to swamp for- ests and mangroves; CULT, grasslands and cultivated ar- eas; TREE, open woodlands, orchards, isolated woodlots, and edges; FOR 1, highly degraded forest with dense sec- ond growth and open canopy, as well as tree plantations; FOR 2, secondary forest heavily logged or disturbed and agroforests; FOR 3, high, submature but logged forest with almost continuous canopy; and FOR 4, large tracts of dense, mostly evergreen, undisturbed, primary forest. Most accounts are based on personal observations ei- ther unpublished or in Thiollay (1978, 1983, 1993, 1995, 1996, 1997) and Thiollay and Meyburg (1988), and sup- plemented by information provided by numerous orni- thologists. The following references were used to sum- marize data available for each country not personally sur- veyed and to estimate population trends by comparison with past status: Baker 1928, Delacour andjabouille 1931, Whistler 1941, Deignan 1963, Smythies 1953, 1981, Wil- dash 1968, Henry 1971, King et al. 1975, Medway and Wells 1976, Ali and Ripley 1978, Morris 1986, van Marie and Voous 1988, Lekagul and Round 1991, MacKinnon and Philipps 1993, del Hoyo et al. 1994. A numerical conservation index was assigned to each species whose breeding population was mostly within the study region according to Usher (1986). It was the sum of three 0-3 scores, such that the species with the most limited geographic range and habitat preference, the highest forest use, and the most declining populations (e.g., the most Vulnerable or Threatened species) got the highest ranking (Table 2). All values were conservative estimates to avoid an overemphasis of the threat level. Results Raptor Community Composition and Distribu- tion. In South Asia, 20 species of Falconiformes are nonbreeding migrants and are found only in win- ter. They include 12 widespread species (5 harriers ( Circus spp., Chinese Goshawk [Accipiter soloensis], Japanese Sparrowhawk [A. gularis ] , Grey-faced Buz- zard [ Butastur indicus ], Common Buzzard [Buteo buteo ], Steppe Eagle [ Aquila nipalensis ], Booted Ea- gle [Hieraaetus pennatus ] , and Eurasian Hobby [Fal- co suhbuteo ] ) . Eight species (Eurasian Black Vulture \Aegypius monachus ] , Eurasian Sparrowhawk [Accip- iter nisus\, Long-legged Buzzard [ Buteo rufinus ] , Greater Spotted Eagle [ Aquila clanga] , Eurasian Imperial Eagle [A. heliaca ], Lesser Kestrel [Falco naumanni\ , Merlin [F. columbarius ] , and Saker Fal- con [F. cherrug]) are rare or local in their occur- rence. Among the 49 breeding species, 15 are spread over the three geographical zones, eight are in two zones, 20 are in only one zone, and six are restrict- ed to Andamans or Java (Table 3). The number of resident species steadily decreases from large con- tinental areas (north and northeast India, 35 taxa), the large southern peninsulas (south India to In- dochina, 23-28 taxa), narrow peninsulas and the largest islands (Malaysia and Borneo, 19-20 taxa), to more outlying islands (Sri Lanka to Java, 17-18 taxa) and oceanic archipelagoes (4—6 taxa). The reduction in species richness is related to both land area, degree of isolation, and historical fac- tors. Habitat Selection and Implications. Eight species associated with interior primary moist forests, ex- cept the island endemics Andaman Serpent-eagle ( Spilornis elgini ) and Nicobar Serpent-eagle ( S. klos- si) , are intolerant of logging or forest fragmenta- tion, These species may have no viable populations outside the current extent of undisturbed forest. Crested Goshawk ( Accipiter trivirgatus) and Wal- lace’s Hawk-eagle ( Spizaetus nanus ) are almost ex- clusively found in lowland forest while Jerdon’s Baza (Aviceda jerdoni) and Javan Hawk-eagle ( Spi- zaetus bartelsi ) are found at elevations as high as submontane forest. Blyth’s Hawk-eagle ( Spizaetus alboniger ) and Mountain Hawk-eagle ( S. nipalensis) are restricted to hill and montane forests and they withdraw in winter from upper altitudes to lowland India and Malaysia. Six forest species actually use forest gaps, upper canopy, and edges for hunting and are more tol- erant to disturbance, fragmentation, and logging. They may actually benefit from partial opening of 44 Thiollay Vol. 32, No. 1 Table 3. Distribution of 49 raptor breeding species within South Asia. Distributed in Number of Species S to NE Indta Andamans AND Nicobars Thailand to Vietnam Malaysia Borneo Sumatra Java Examples 4 + + + + + Spilornis cheela, Accipiter virga- tus 7 + + + + Pernis ptilorhynchus, Accipiter tri- virgatus 1 + + Aviceda leuphotes 4 + + + Aviceda jerdoni, Ichthyophaga humilis 1 + + + Falco severus 1 + + + Falco peregrinus 6 + + Accipiter badius, Milvus migrans 11 + Butastur teesa, Falco chicquera 3 + Spilornis elgini , S. klossi 1 + Polihierax insignis 1 + + Butastur liventer 5 -I- Spizaetus alboniger, S. nanus 1 + + Michrohierax fringillarius 3 + Spizaetus bartelsi, Falco moluccen- sis Total 35 8 24 22 18 Andamans-Nicobars (oceanic archipelago) and Java (landbridge island), because of their endemic taxa, have been separated from the larger Indian, Indochinese and Sundan subregions. the forest. Five species (Black Baza [Aviceda leupho- tes ] , Oriental Honey-buzzard [Pernis ptilorhynchus ] , Besra [Accipiter virgatus ], Indian Black Eagle [Icti- naetus malayensis ], and Rufous-bellied Eagle [Hi- eraaetus kieneni] ) have a wide altitudinal range, while one species (Kinabalu Serpent-eagle [Spilor- nis kinabaluensis ]) is restricted to montane forest. Populations of honey-buzzards and bazas are aug- mented in winter when the northern subspecies migrates south. All six species easily fly between separate forest patches. Nineteen species are associated with woodlands, edges, and open or degraded secondary forests. They are not true forest species and tolerate a moderate level of deforestation and readily accept tree plantations. Crested Serpent-eagle ( Spilornis cheela), Shikra ( Accipiter badius), Nicobar Sparrow- hawk (A. butleri), Changeable Hawk-eagle ( Spizaetus cirrhatus), White-rumped Pygmy Falcon ( Polihierax msignis), and Red-necked Falcon ( Falco chicquera ) forage in forests from mid-level to the ground. All species of falconets ( Microhierax spp.) take prey in flight from exposed perches, including emergent trees of closed canopies. Bat Hawks ( Macheiram - phus alcinus) , Oriental Hobbies ( Falco severus) , and Peregrine Falcons ( F. peregrinus) hunt above or around the forest and are more sensitive to prey abundance than to vegetation structure. Wintering migrants (Chinese Goshawk, Japanese Sparrow- hawk, Eurasian Sparrowhawk, Common Buzzard, Booted Eagle \Hieraaetus pennatus ] , and Eurasian Hobby) share the least wooded habitat types. Ten migrant and 14 resident species use open grasslands and cultivated and suburban areas. They are mostly distributed in the drier, less forested northern tropics. They appear to avoid dense for- ests, but only the smaller species (Black-winged Kite [ Elanus caeruleus ], two species of buzzards [Bu- tastur spp.], and three species of falcons), migrants (Grey-faced Buzzard [Butastur indicus ], Long-leg- ged Buzzard, two species of harriers and three spe- cies of falcons), and scavengers (Black Kites [Mil- vus migrans], and six species of vultures) may ac- tually benefit from extensive deforestation, culti- vation, and overgrazing. Eagles (Short-toed Snake-eagle [ Circaetus gallicus\ and three species of Aquila ) usually remain associated with more natu- ral habitats. March 1998 Status of Raptors in Tropical Asia 45 Eleven wetland species use marshes and rice- fields, freshwater lakes, coastal lagoons, rivers and forest streams, or sea coasts. Residents are Brah- miny Kite ( Haliastur indus) , White-bellied Sea-eagle ( Haliaeetus leucogaster ) , Pallas’s Sea-eagle (H. leuco- ryphus), Grey-headed Fish-eagle ( Ichthyophaga ich- thyaetus), Lesser Fish-eagle (/. humilis ) , and Lesser Spotted Eagle ( Aquila pomarina ) . Migrants include Western Marsh-harrier ( Circus aeruginosus ) , Eastern Marsh-harrier (C. spilonotus ), Pied Harrier (C. me- lanoleucos), and Greater Spotted Eagle. The Osprey (Pandion haliaetus) has a small coastal breeding population in Java but is overall a widespread win- tering species. The response of wetland raptors to habitat changes differs among species. Large pi- scivorous birds such as sea and fishing eagles are often adversely affected because of decreasing fish stocks and suitable trees for hunting perches along banks. More terrestrial eagles (e.g., Aquila) may be affected by a lack of undisturbed shallow 7 marshes and humid grasslands, whereas more versatile spe- cies (e.g., species of Haliastur and Circus) may be benefitted because they are much more adaptable to heavy human disturbance of wetlands. Northern migrants, except locally the Booted Eagle (Hieraaetus pennatus), are virtually absent from forests. They make up 32-42% of all raptor species in the other three habitat categories. Conservation Status. The number of Threat- ened or Near Threatened species is alarming. Among the 49 breeding taxa, 22 have a high con- servation index (5-9) and/or are in one of the categories of threat in the IUCN World Red List (Table 4, Collar et al. 1994). There is no extinct species nor is any raptor population currently de- pendent on a captive-breeding program. However, two subspecies of the Lesser Spotted Eagle ( Aquila : pomarina hastata) from India, and Wallace’s Hawk- eagle ( Spizaetus nanus stresemanni) from Nias Is- land, W. Sumatra, seem to be Critically Endan- gered (Thiollay 1996). All eight forest interior species, nearly half of the more tolerant forest or wedand species, and 25% of woodland and open habitat species are consid- ered Rare, Threatened or of Special Concern be- cause of their high conservation index (Table 5). Only two wintering migrants are globally Vulnera- ble (Greater Spotted Eagle) or Near Threatened (Eurasian Black Vulture, Aegypius monachus). Sev- eral other migrants have small and local popula- tions in South Asia, but they are widespread else- where. Population Trends. There are very few accurate data on raptor population dynamics in South Asia or trends in recent decades, yet many species con- sidered common or widespread 50 yr ago in re- gional accounts are now obviously rarer and more local. This is especially true for eagles, both from rainforests (hawk-eagles) and wetlands (fishing ea- gles), but also for many usually abundant species in genera such as Milvus , Haliastur, Elanus, Accipiter, Butastur, and Falco, which are now rare, if not al- most extirpated from large, intensively cultivated areas in Thailand, Vietnam, Malaysia, and Java. The decline is especially striking for vultures ( Gyps spp. and Sarcogyps calvus ) w 7 hich have apparendy disappeared, at least as breeders, from vast regions in Peninsular Malaysia, Thailand, and Indochina where they were once considered common. Even in India, their main stronghold, my roadside counts over a 20-yr period show significant de- clines in some areas. Finally, at least the forest in- terior species which cannot adapt to degraded woodlands or even to fragmented secondary forest, may be decreasing in parallel with deforestation. Specific Regional and Distribution Patterns. An increased density and a wider habitat niche of some species may result from competitive release when fewer competitors coexist, especially on is- lands (Wright 1980, Blondel 1990). Field measure- ments showed such striking examples of habitat shifts and density compensations. For example, Crested Serpent-eagles are widespread everywhere in secondary forests and woodlands in the absence of any congeners. However, on the Andaman ar- chipelago, it is associated only with mangroves, for- est edges or gaps, and deforested areas, whereas the endemic Andaman Serpent-eagle occupies all types of forest interior (Fig. 2), In another exam- ple, Changeable Hawk-eagle is restricted to open woodlands and degraded forests in India at upper elevations, in Malaysia, Borneo, and Sumatra be- cause another Spizaetus always replaces it in dense forests. In Vietnam, lowland India, and the Anda- mans, where there is no forest Spizaetus, Changea- ble Hawk-eagles, are found throughout primary forests and more open secondary stands (Fig. 3). Altitudinal segregation occurs between Wallace’s Hawk-eagle (in the lowlands) and Blyth’s Hawk-ea- gle (at upper elevations) throughout their sympat- ric distribution. The Besra is a common forest rap- tor at any elevation from southwest India to Viet- nam and Java, in sympatry with the larger Crested Goshawk. Conversely, in northern India, Nepal, 46 Thiollay Vol. 32, No. 1 Table 4. Conservation status of breeding raptors in tropical Asia, west of the Wallace Line. The conservation index (see Methods) is given for mostly South Asian species, and not for taxa with extra world range. The IUCN status (criteria in Mace and Stuart 1994) involves only Threatened (= Endangered or Vulnerable) or Near Threatened (low risk) species. Species Conservation Status Species whose world range is mostly within South Asia Jerdon’s Baza (Aviceda jerdoni) 7, low risk Black Baza ( Aviceda leuphotes) 5 Oriental Honey-buzzard ( Pernis ptilorhynchus) 3 Brahminy Kite ( Haliastur indus) 2 White-bellied Sea-eagle ( Haliaeetus leucogaster) 3 Lesser Fish-eagle ( Ichthyophaga humilis ) 6, low risk Grey-headed Fish-eagle ( Ichthyophaga ichthyaetus) 6, low risk Indian White-backed Vulture ( Gyps bengalensis) 4, low risk Long-billed Vulture ( Gyps indicus) 4, low risk Red-headed Vulture ( Sarcogyps calvus ) 7, low risk Crested Serpent-eagle ( Spilornis cheela) 2 Nicobar Serpent-eagle ( Spilornis klossi ) 6, low risk Kinabalu Serpent-eagle ( Spilornis kinabaluensis) 7, data deficient Andaman Serpent-eagle ( Spilornis elgini) 6, low risk Crested Goshawk ( Accipiter trivirgatus) 5 Nicobar Sparrowhawk ( Accipiter butleri ) 7, low risk Besra (Accipiter virgatus) 4 White-eyed Buzzard ( Butastur teesa) 4 Rufous-winged Buzzard ( Butastur liventer ) 6, low risk Indian Black Eagle ( Ictinaetus malayensis ) 4 Rufous-bellied Eagle ( Hieraaetus kienerii) 6, low risk Changeable Hawk-eagle ( Spizaetus cirrhatus) 2 Mountain Hawk-eagle ( Spizaetus nipalensis) 7, low risk Blyth’s Hawk-eagle ( Spizaetus alboniger) 7 Javan Hawk-eagle ( Spizaetus bartelsi ) 9, Endangered Wallace’s Hawk-eagle ( Spizaetus nanus) 9, Endangered White-rumped Pygmy Falcon ( Polihierax insignis) 5, low risk Collared Falconet ( Microhierax caerulescens ) 4 Black-tighted Falconet ( Microhierax fringillarius ) 4 White-fronted Falconet ( Microhierax latifrons ) 5, data deficient Pied Falconet ( Microhierax melanoleucus ) 5, low risk Oriental Hobby (Falco severus ) 4 Laggar Falcon (Falco jugger) 3 Species with small and marginal breeding population in South Asia, ranging mo Osprey (Pandion haliaetus ) stly outside the region Pallas’s Sea Eagle (Haliaeetus leucoryphus) Eurasian Griffon (Gyps fulvus) Short-toed Snake-eagle ( Circaetus gallicus) Vulnerable Lesser-spotted Eagle (Aquila pomarina) Spotted Kestrel (Falco moluccensis ) Endangered Species with a substantial population in South Asia, but with a larger population Bat Hawk (Macheiramphus alcinus) Black-winged Kite (Elanus caeruleus) Black Kite (Milvus migrans) Egyptian Vulture (Neophron percnopterus) Shikra (Accipiter badius ) Tawny Eagle (Aquila rapax) elsewhere (Asia to Africa) March 1998 Status of Raptors in Tropical Asia 47 Table 4. Continued. Species Conservation Status Bonelli’s Eagle ( Hieraaetus fasciatus) Common Kestrel ( Falco tinnunculus) Red-necked Falcon ( Falco chicquera) Peregrine Falcon ( Falco peregrinus) low risk Borneo, Sumatra, and maybe Malaysia, it is con- fined to upper elevations (1000-2000 m) separated from the Crested Goshawk which occurs alone in the lowlands. When all species were pooled, the density index of raptors in mature forests (but not in deforested habitats) was similar in the study areas of southwest India, southern Vietnam and western Sumatra, but it increased significantly in Sri Lanka and the An- daman Islands (+45%, P < 0.001, Table 6). Most taxa exhibited a significant density increase in un- disturbed forests both between small and large is- lands and between islands and continental areas. The two Serpent-eagles in the Andamans reached extremely high densities in their preferred habitat: 2.78 birds/km 2 sample plot for Andaman Serpent- eagles in primary forests and 2.25 birds/km 2 for Crested Serpent-eagles in mangroves vs. 0.71-0.98 birds/km 2 for the latter in continental woodlands. The abundance of Black Bazas ( Aviceda leuphotes) was three times higher in the Andamans than in Vietnam. However, the density of the one species of goshawk and hawk-eagle in Andamans was equaled and even surpassed by the cumulative abundance of all goshawks or hawk-eagles and ea- gle species ( Hieraaetus spp.) in the forests of the four other study areas. High population densities enable birds on relatively small islands to minimize their extinction risk. A higher tolerance to habitat change or disturbance may also allow them to bet- ter withstand human-made habitat degradation and fragmentation which tend to be especially se- vere on islands. Main Threats and Conservation Issues Habitat Loss. Most natural habitats in tropical Asia are decreasing primarily because of defores- tation. Most areas with fertile, deep soils have long histories of cultivation. The few remaining forests are on infertile soils, peat swamps, and steep slopes that support poorer forest types and less complex animal communities (Collins et al. 1991). Lowland rainforests, the richest of them all, are most at risk and are the target of most new setdements. Moist forests were disappearing during the early 1980s at an estimated rate of 1 7 000 km 2 /yr within our study area alone (FAO 1988, Collins et al. 1991) and there is no evidence that this trend has decreased since then. Clearcutting for either shifting or per- manent industrial cultivation is the main cause of deforestation. In monsoon forests, heavy collection of fuel wood is an additional pressure. Montane forests are more secure because of their poor ac- cessibility and suitability for agriculture but they have also lower raptor species richness. In north- ern Borneo and western Sumatra, I recorded 10- 11 species in all primary forests surveyed below 1200 m while at 1200-2200 m, there were only 4— 6 species. The destruction of swamp and riverine forests Table 5. Status, habitat selection and conservation of raptors in South Asia. Number of species in each category among the 69 regular species. Resident Wintering Rare 3 Threatened 15 Mature forest interior species 8 4 8 Gap tolerant forest species 6 2 3 Woodland or forest edge species 14 6 5 5 Dry open habitat species 14 10 8 6 Wetland species 7 4 5 5 3 Rare = low density and/ or patchy distribution in suitable habitats. b Threatened = Endangered to low risk, data deficient and/or conservation value 5-9. 48 Thiollay Vol. 32, No. 1 Figure 2. Comparative distribution of the widespread Crested Serpent-eagle ( Spilornis cheela ) in India, Sumatra, and Andamans and the endemic Andaman Serpent-eagle (S. elgini) in the Andamans. Percentages of random records in each habitat category (see text for definition). B = habitat niche breadth. Values on bars are the selection indices from relative habitat availability. * = statistically significant positive or negative selection (Neu et al. 1974). affects eagles and the drainage of wetlands is det- rimental to harriers and other wintering migrants. Ricefields that replace marshes are much poorer in vertebrate prey, and mangroves are heavily used for timber and shrimp farming. Large mangrove forests in estuaries, such as the Sundarbans (Sarker 1986), harbor dense populations of several raptors that have already been lost from similar areas in southern Vietnam (Morris 1986). Habitat Degradation. Habitat degradation is an insidious, widespread phenomenon that diminish- es the quality of natural habitats. Raptor commu- nities become less complex in these areas and their overall density may also decline. From open wood- lands to undisturbed rainforests in all five study areas, the overall mean species richness per sample plot increased by 162% and the mean abundance index by 176% (Table 4). The main causes of forest degradation are log- ging and shifting cultivation. Both lead to second- ary stands whose general characteristics include lower and more open canopies, a reduced number March 1998 Status of Raptors in Tropical Asia 49 Habitats Figure 3. Comparative distribution of Changeable Hawk-eagle ( Spizaetus cirrhatus ) with a forest congener in India and Sumatra and without competitor in the An- daman Islands. Symbols as in Fig. 2. and diversity of large trees, a denser undergrowth, and a higher frequency of gaps and dense vine tan- gles than primary forest. Effects of logging are se- vere (Burgess 1971). During the first years after logging, the density, diversity, and accessibility of many prey species decrease markedly (Wilson and Johns 1982, Johns 1985, 1986, 1987, Bennett and Dahaban 1995, Hill et al. 1995), although a partial recovery occurs after 10 yr and suitable manage- ment practices may minimize losses (Johns 1989, 1992). The ultimate stages of forest degradation are dense stands of bamboo ( Bambusacae ) or patch- es of high coarse grass ( Imperata cylindrica), both of which prevent forest regeneration and are unsuit- able for most raptors. Road building through forests brings distur- bance, hunting, gaps, and dense edges. Sylviculture leads to decreasing biodiversity and permanently submature stands because some fast-growing spe- cies are favored over hardwoods (Panayotou and Ashton 1992). The widespread exploitation of mi- nor forest products, especially resins and rattans ( Calamus sp.) is an additional cause of disturbance and hunting. Raptor species adapting to man-made habitats naturally occur in open grasslands, woodlands, for- est edges, and gaps. They are generalists and have a wide distribution. Most artificial landscapes are unsuitable for dense forest specialists with indus- trial tree plantations being the poorest. Even the diversified traditional agroforests of Indonesia do not conserve more than half the forest raptor spe- cies and only as long as natural forest remains Table 6. Mean number of raptor/ species and adult birds on sample plots surveyed in five areas of South Asia. SW Ghats South Western Sri Lanka (India) Vietnam Sumatra SW Andamans Total Number of 1-km 2 Sample Plots 156 56 103 18 130 Open habitats Species richness/ 1.71 ± 1.14 1.21 ± 1.05 0.47 ± 0.15 1.00 ± 0.63 0.79 ± 0.17 (CULT + TREE) plot (x ± SD) Individuals/plot 2.37 ± 1.74 2.00 ± 1.46 0.77 ± 1.09 1.17 ± 0.75 1.04 ± 1.12 (x ± SD) Degraded forests, Species richness/ 2.11 ± 0.77 1.77 ± 1.36 1.29 ± 0.77 2.50 ± 0.54 1.77 ± 0.82 agroforests plot (x ± SD) and mangroves Individuals/plot 3.06 ± 1.24 2.77 ± 2.04 1.83 ± 0.15 4.17 ± 1.16 2.77 ± 1.45 (FOR 1 + FOR 2) (x ± SD) Primary or Species richness/ 2.55 ± 0.97 2.45 ± 1.08 2.64 ± 0.87 3.33 ± 1.63 2.60 ± 0.65 litde disturbed plot (x ± SD) forests Individuals/plot 3.65 ± 1.47 3.67 ± 1.68 3.43 ± 1.16 4.33 ± 1.36 5.20 ± 1.51 (FOR 3 + FOR 4) (x ± SD) 50 Thiollay Vol. 32, No. 1 nearby (Thiollay 1995). Many dams, built in agri- cultural areas, are favored by Black and Brahminy Kites, Grey-headed Fish-eagle, and wintering har- riers which also benefit from irrigated and more wooded surroundings. Forest Fragmentation and Insularization. Be- cause of their necessarily large foraging range, low density, and sensitivity to disturbance, forest rap- tors may be severely affected by forest fragmenta- tion, especially forest interior species that are re- luctant to cross large open areas. In lowland forest study areas of India, Sri Lanka, Vietnam, and Su- matra, there were 0.16-0.34 bazas, hawk-eagles, ea- gles ( Hieraaetus spp.) or Besras per km 2 . Assuming that there was at least an extra bird for each ter- ritorial pair, these figures represented an average density of about one pair of each species/ 1000— 1500 ha of primary forest. Two examples illustrate the magnitude of the fragmentation process. In the 1600-km long west- ern Ghats of Peninsular India, only 20 000 km 2 of humid evergreen forest remain, but the largest un- disturbed continuous fragment is the 90 km 2 Silent Valley National Park (Daniels 1996). In Cochinchi- na (23 500 km 2 ), a single 280 km 2 patch of semi- evergreen forest remains (approximately 1% of the province) within the 350 km 2 Nam Cat Tien Na- tional Park. The Indian Black Eagle, formerly re- corded, has not been found there during recent intensive surveys. In the Andaman archipelago, the most extinc- tion-prone raptors on small islands were the rarest and most specialized species, but unexpectedly they were also the smallest ones. Serpent-eagles and hawk-eagles were found on all islands smaller than 1 km 2 . Some pairs range over two or three small islands showing a high tolerance to naturally fragmented forest (Thiollay 1997). Persecution and Pollution. Raptor shooting is common, as well as trapping and nest robbing for pets, trophies, and sales to restaurants. Trade is es- pecially prominent in Indonesia, Thailand, and In- dochina. The lack of hunting in large parts of In- dia results in an obviously higher frequency of rap- tors in densely inhabited areas than anywhere else. Hunting and trapping pressure, however, may have even more severe effects through reduction of prey species, notably for eagles. In large parts of Indo- china, wildlife has been reduced to extremely low levels. Overfishing in streams, lakes, and marshes may also be a limiting factor for piscivorous eagles. There is no monitoring of pesticide levels in Asian raptors nor definite evidence of their effects. However, pesticides are heavily used in most coun- tries (except Laos) , especially Indonesia and Thai- land. Ricefields are routinely sprayed even in re- mote areas. This trend is probably increasing be- cause of the intensification of agriculture and the use of more productive but pest-sensitive new va- rieties. The dramatic disappearance of raptors from several cultivated regions (e.g., Java [van Bal- en et al. 1993], Thailand [Lekagul and Round 1991], and Vietnam [Thiollay pers. obs.]) suggests that pesticides have had a negative effect on rap- tors over the last 20 yr. No comparable declines have been recorded in forested or little-cultivated areas. The disappearance of the once common vul- tures and kites in populated areas, especially Thai- land, Indochina, and Malaysia, can be explained by a lower food availability due to increased sani- tation. This may have also begun to occur in India. Conclusions and Recommendations In summary, tropical Asian raptors may be divid- ed into three broad groups from a conservation point of view. First, true forest species mostly sen- sitive to habitat loss, degradation, and fragmenta- tion, need protected areas of undisturbed forests. The second group includes forest edge or gap and woodland species that are more tolerant to forest disturbance and opening. Their conservation re- quires habitat management, measures to minimize the effects of habitat changes, and provision of their minimum needs in terms of tree cover and prey availability. Most wetland species are also de- pendent on habitat management actions. The last group includes the grassland species which are more adaptable to human activity and are now largely found in cultivated and deforested areas. Their highest management priority is the direct protection from persecution and pollution. Most migrants also fall in this category. Basic knowledge necessary to design appropriate conservation strategies is often lacking. Specific ecological requirements are little known, popula- tion levels and trends are not quantitatively docu- mented, and threats and causes of declines are poorly understood. Also, demographic parameters, minimum viable population sizes, and dispersal abilities have never been investigated in most Asian raptor species. An intensive survey and conserva- tion action plan are currently in progress only for the Javan Hawk-eagle (estimated population 100 March 1998 Status of Raptors in Tropical Asia 51 pairs, van Balen 1995). Assessing the distribution, size, and viability of remaining populations of threatened forest and wetland eagles such as hawk- eagles and fishing eagles is an urgent priority. Only immediate measures could save the Critically En- dangered subspecies of the Wallace’s Hawk-eagle ( Spizaetus nanus stresemanni) from Nias Island or the Lesser Spotted Eagle ( Aquila pomarina hastata ) from Indo-gangetic plains. The extension and enforcement of the current network of protected areas is a high priority. Pro- tected areas cover only 4.5% of Southeast Asia (IUCN 1990) and many of them are heavily dis- turbed and increasingly encroached by people. The highest conservation priority is the lowland rainforest which is fast disappearing everywhere. Forest management policies must promote biodi- versity conservation in production forests, reduce logging damages, maintain a network of unlogged patches, and avoid large plantations of exotic trees. Tree cash crops could be made more hospitable to raptors by increasing their structural and floristic diversity, and traditional agroforests must be en- couraged (Thiollay 1995). Finally, cultivated areas become the most widespread habitat and the main wintering ground for migrants. Therefore, their sound management is a priority for the conserva- tion of raptors in Asia, with an emphasis on the maintenance of trees and fallow patches, and a drastic reduction of hunting pressure and pesticide use. Literature Cited Ali, S. and S.D. Ripley. 1978. Handbook of the birds of India and Pakistan. Vol. 1. Oxford Univ. Press, Delhi, India. Baker, E.C.S. 1928. The fauna of British India birds. Vol. V. Taylor and Francis, London, U.K. Bennett, E.L. and Z. Dahaban. 1995. Wildlife response to disturbances in Sarawak and their implications for forest management. Pages 66-86 in R.B. Primack and E. Lovejoy [Eds.], Ecology, conservation and manage- ment of Southeast Asian rainforests. Yale Univ. Press, New Haven, CT U.S.A. Blondel, J. 1990. Long-term studies on bird communi- ties and populations in mainland and island Mediter- ranean forests. Pages 167-182 in A. Keast [Ed.], Bio- geography and ecology of forest bird communities. S.P.B. Academic Publ., The Hague, Netherlands. Burgess, P.F. 1971. The effect of logging on hill dipter- ocarp forests. Malay. Nat. J. 24:231-237. Collar, N.J., M.J. Crosby and A.J. Stattersfield. 1994. Birds to watch. The world list of threatened birds. Birdlife International, Cambridge, U.K. Collins, N.M., J.A. Sayer and T.C. Whitmore [Eds]. 1991. The conservation atlas of tropical forests, Asia and the Pacific. IUCN and MacMillan Press, London U.K. Daniels, R.J.R. 1996. Landscape ecology and conserva- tion in the western Ghats, south India. Ibis 138:64—69. Deignan, H.G. 1963. Checklist of the birds of Thailand Bull. U.S. Natl. Mus. 226, Smithsonian Inst., Washing- ton DC U.S.A. del Hoyo, J., A. Elliott and J. Sargatal. 1994. Hand- book of the birds of the world. Vol. 2. Lynx Edicions, Barcelona, Spain. Delacour, J. and P. Jabouille. 1931. Les oiseaux de l’lndochine Fran^aise. Exposition coloniale interna- tionale, Paris, France. FAO. 1988. An interim report on the state of forest re- sources in the developing countries. FAO, Rome, It- aly. Henry, G.M. 1971. A guide to the birds of Ceylon. Ox- ford Univ. Press, London U.K. Hill, J.K., K.C. Hamer, L.A. Lace and W.M. Banham. 1995. Effects of selective logging on tropical forest butterflies on Burn, Indonesia./. Applied Ecol. 32:754— 760. IUCN. 1990. IUCN directory of South Asian protected areas. IUCN, Gland and Cambridge, U.K. Johns, A.D. 1985. Selective logging and wildlife conser- vation in tropical rainforest: problems and recom- mendations. Biol. Conserv. 31:355-375. . 1986. Effects of selective logging on the ecolog- ical organization of a peninsular Malaysian rainforest avifauna. Forktail 1:65-79. . 1987. The use of primary and selectively logged rainforest by Malaysian Hornbills ( Bucerotidae ) and im- plications for their conservation. Biol Conserv. 40:179- 190. . 1989. Recovery of a peninsular Malaysian rain- forest avifauna following selective timber logging: the first twelve years. Forktail 4:89—105. . 1992. Species conservation in managed tropical forests. Pages 15-53 in T.C. Whitmore and J.A. Sayer [Eds.], Tropical deforestation and species extinction. Chapman and Hall and IUCN, London, U.K. King, B., M. Woodcock and E.C. Dickinson. 1975. A field guide to the birds of Southeast Asia. Collins, London, U.K. Lekagul, B. and P.D. Round. 1991. A guide to the birds of Thailand. Saha Karn Bhaet Co., Bangkok, Thai- land. Mace, G. and S. Stuart. 1994. Draft IUCN Red List Cat- egories. Species 21—22:13—24. MacKinnon, J. and K. Philipps. 1993. A field guide to the birds of Borneo, Sumatra, Java, and Bali. Oxford Univ. Press, Oxford, U.K. Medway, L. and D.R. Wells. 1976. The birds of the Ma- lay peninsula. Witherby, London U.K. 52 Thiollay Vol. 32, No. 1 Morris, G.E. 1986. Birds of prey in Vietnam. Birds of Prey Bull 3:163-169. Neu, L.W., C.R. Byers and J.M. Peek. 1974. A technique for analysis of utilization-availability data. J, Wildl. Manage. 38:541-545. Panayotou, T. and P.S. Ashton. 1992. Not by timber alone. Economics and ecology for sustaining tropical forests. Island Press, Washington, DC U.S.A. Sarker, S.U. 1986. Population dynamics of raptors in the Sundarban forests of Bangladesh. Birds of Prey Bull. 3: 157-162. Smythies, B.E. 1953. The birds of Burma, 2nd Ed. Oliver and Boyd, Edinburgh U.K. . 1981. The birds of Borneo. The Sabah Society, Kota Kinabalu. Sokal, R.R. AND F.J. Rohlf. 1981. Biometry. Freeman and Co., San Francisco, CA U.S.A. Thiollay, J.M. 1978. Distribution des Falconiformes ni- cheurs autour du massif de l’Annapurna (Himalaya central). Oiseau Rev. Fr. Omithol. 48:291-310. . 1983. Evolution actuelle du peuplement de ra- paces diurnes dans le nord de Borneo. Alauda 51: 109-123. . 1989. Censusing of diurnal raptors in a primary rainforest: comparative methods and species detect- ability. J. Raptor Res. 23:72—84. . 1993. Response of a raptor community to shrinking area and degradation of tropical rainforest in the southwestern Ghats (India). Ecography 16:97- 110 . . 1995. Rainforest raptor communities in Suma- tra: the conservation value of traditional agroforests. In D.M. Bird, D.E. Varland andJ.J. Negro [Eds.], Rap- tor adaptations to human influenced environments. Academic Press, London U.K. . 1996. The raptor community of Nias Island, Su- matra: survey and conservation. Rukila 8:113-116. . 1997. Distribution and abundance patterns of bird community and raptor populations in the An- daman Archipelago. Ecography 20:67-82. and B.-U. Meyburg. 1988. Forest fragmentation and the conservation of raptors: a survey of the island of Java. Biol. Conserv. 44:229-250. Usher, M.B. 1986. Wildlife conservation evaluation: at- tributes, criteria and values. Pages 3-44 in M.B. Usher [Ed.], Wildlife conservation evaluation. Chapman and Hall, London, U.K. van Balen, B. 1995. Red data bird: Javan Hawk-eagle. World Birdwatch 17:20-21. , I.S. Suwelo, D.S. Hadi, D. Soepomo, R. Marlon and Mutiarina. 1993. The decline of the Brahminy Kite, Haliastur indus, on Java. Forktail 8:83-88. van Marle, J.G. and K.H. Voous. 1988. The birds of Sumatra. B.O.U. Checklist No. 10. British Ornitholo- gists’ Union, Tring, U.K. Whistler, H. 1941. Popular handbook of Indian birds. Gurney and Jackson, London U.K. Whitmore, T.C. 1985. Tropical rainforests of the Far East, 2nd Ed. Clarendon Press, Oxford U.K. Wildash, P. 1968. Birds of south Vietnam. Charles Tutde Publ., Tokyo, Japan. Wilson, W.L. and A.D. Johns. 1982. Diversity and abun- dance of selected animal species in undisturbed for- est, selectively logged forest, and plantations in East Kalimantan, Indonesia. Biol. Conserv. 24:205-218. Wright, S.J. 1980. Density conservation in island avifau- nas. Oecologia 45:385—389. Received 16 October 1996; accepted 9 October 1997 March 1998 Status of Raptors in Tropical Asia 53 Appendix 1. Status and distribution of raptors in South Asia. B = breeding but may include additional, wintering nonresident populations; W = only wintering migrants, usually from Eastern Palearctic zone. Number of subspecies in parentheses after species names. N Pen. Sri NE Anda- Bur- Thai- Viet- Malay Sara- S Bor- Suma- India India Lanka India mans MA land NAM Pen. WAK neo IRA Java Osprey Pandion haliaetus (2) Jerdon’s Baza W W W W w W W W w W w B Aviceda jerdoni (3) Black Baza B B B B B B W B B B Aviceda leuphotes (3) Oriental Honey-buzzard B W B B B W W W W Pernis ptilorhynchus (4) Bat Hawk B B B B W B B B B B B B B Macheiramphus alcinus (1) Black-winged Kite B B B B Elanus caeruleus (2) Black Kite B B B B B B B B B B B B Milvus migrans (2) Brahminy Kite B B B B B B B W Haliastur Indus (2) B B B B B B B B B B B White-bellied Sea-eagle Haliaeetus leucogaster (1) Pallas’s Sea-eagle B B B B B B B B B B B B B Haliaeetus leucoryphus (1) B B B Lesser Fish-eagle Ichthyophaga humilis (1) Grey-headed Fish-eagle B B B B B B B B Ichthyophaga ichthyaetus (2) Egyptian Vulture B B B B B B B B B B B B Neophron percnopterus ( 1 ) Indian Black-backed Vulture B B Gyps bengalensis (1) Long-billed Vulture B B B B B B B Gyps indicus (2) Eurasian Griffon B B B B B B Gyps fulvus (1) Eurasian Black Vulture B B Aegypius monachus (1) Red-headed Vulture W Sarcogyps calvus (1) Short-toed Snake-eagle B B B B B B B Circaetus gallicus (1) Crested Serpent-eagle B B W W W W Spilomis cheela (16) B B B B B B B B B B B B B Nicobar Serpent-eagle Spilomis klossi (2) Kinabalu Serpent-eagle Spilomis kinabaluensis (1) Andaman Serpent-eagle B B Spilomis elgini (1) B Pallid Harrier Circus macrourus (1) Pied Harrier W W W W W W Circus melanoleucos (1) W W W W 54 Thiollay Vol. 32, No. 1 Appendix 1. Continued. N Pen. Sri NE Anda- Bur- Thai- Viet- Malay Sara- S Bor- SUMA- India India Lanka India MANS MA land NAM Pen. WAK NEO TRA Java Montagu’s Harrier Circus pygargus (1) Western Marsh Harrier W W W W Circus aeruginosus (1) Eastern Marsh Harrier W W w W Circus spilonotus (1) Crested Goshawk w W w W w w Accipiter trivirgatus (7) Shikra B B B B B B B B B B B B Accipiter badius (4) Nicobar Sparrowhawk Accipiter butleri (2) Chinese Goshawk B B B B B B B B Accipiter soloensis (1) Japanese Sparrowhawk W W W W W W W W W Accipiter gularis (2) Besra W w w w W W W W W Accipiter virgatus (7) Eurasian Sparrowhawk B B B B B B B B B B B B B Accipiter nisus (1) White-eyed Buzzard W W W W W W W Butastur teesa ( 1 ) B B B B Rufous-winged Buzzard Butastur liventer (1) Grey-faced Buzzard B B B Butastur indicus ( 1 ) Common Buzzard W W W W W W W Buteo buteo (2) Long-legged Buzzard W W W W W W W W Buteo rufinus (1) w Indian Black Eagle Ictinaetus malayensis (2) Lesser Spotted Eagle B B B B B B B B B B B B Aquila pomarina (1) Greater Spotted Eagle B B Aquila clanga (1) W W W W W Tawny Eagle Aquila rapax (1) Steppe Eagle B B B B W Aquila nipalensis (1) Eurasian Imperial Eagle W W W W Aquila heliaca (1) Bonelli’s Eagle W w W Hieraaetus fasciatus (I) Booted Eagle B B B B B B Hieraaetus pennatus ( 1 ) Rufous-bellied Eagle W W W W W Hieraaetus kienerii (2) B B B B B B B B B B B Changeable Hawk-eagle Spizaetus cirrhatus (5) Mountain Hawk-eagle B B B B B B B B B B B B B Spizaetus nipalensis (2) B B B B B W W March 1998 Status of Raptors in Tropical Asia 55 Appendix 1. Continued. N Pen. Sri NE Anda- Bur- Thai- Viet- Malay Sara- S Bor- SUMA- India India Lanka India MANS ma land NAM Pen. WAK neo IRA JAVA Blyth’s Hawk-eagle Spizaetus alboniger (1) Javan Hawk-eagle Spizaetus bartelsi (1) Wallace’s Hawk-eagle B B b B B Spizaetus nanus White-rumped Pygmy Falcon B B B B Polihierax insignis (3) B B B Collared Falconet Microhierax caerulescens (2) Black-thighed Falconet Microhierax fringillarius (1) White-fronted Falconet B B B B B B B B B B Microhierax latifrons (1) Pied Falconet B Microhierax melanoleucus (1) B B B Lesser Kestrel Falco naumanni (1) Common Kestrel W W W Falco tinnunculus (2) Spotted Kestrel Falco moluccensis (1) Red-necked Falcon W B B W W W W B Falco chicquera (1) B B Merlin Falco columbarius (2) Eurasian Hobby W w Falco subbuteo (2) W W W W W w Oriental Hobby Falco severus (1) Laggar Falcon B W W B B B B B Falco jugger ( 1 ) Saker Falcon B B B B Falco cherrug (1) Peregrine Falcon W W Falco peregrinus (5) B B B B w B W W B B B B B Total breeding 31 28 17 31 8 31 23 23 19 20 18 18 18 Total wintering 16 11 9 14 5 13 15 15 10 6 5 6 4 Pen = Peninsular, Andamans = + Nicobars, Thailand = +Cambodia, Vietnam = +Laos, Sarawak — + Sabah. f. Raptor Res. 32(l):56-63 © 1998 The Raptor Research Foundation, Inc. TROPICAL FOREST RAPTORS IN INDONESIA: RECENT INFORMATION ON DISTRIBUTION, STATUS, AND CONSERVATION S. (Bas) van Balen BirdLife International— Indonesia Programme, P. O. Box 31 0, BOGOR 1 6003, Indonesia; and Department of Terrestrial Ecology and Nature Conservation, Wageningen Agricultural University, Bornsesteeg 69, 6708 PD Wageningen, The Netherlands Abstract. — In Indonesia (subregions Sumatra, Kalimantan, Java/Bali, Sulawesi, Lesser Sundas, Moluc- cas, and Irian Jaya) , 68 species of diurnal birds of prey, nine of which are endemic to the region, are known to occur. Ever-growing pressure on natural habitats, in particular the timber-rich, tropical rain- forests, as well as hunting and the use of pesticides, is believed to have major impacts on the numbers and distribution of these species. The status of 20 species is a matter of concern since three species are considered globally Threatened and eleven others are Near Threatened. The status of yet another five species is of local concern as they are widespread and relatively common elsewhere. Key Words: status diurnal birds of prey; conservation ; distribution', Indonesia. Aves rapaces de bosques tropicales en Indonesia: informacion reciente sobre su distribucion, estado, y conservacion Resumen. — En Indonesia (subregiones de: Sumatra, Kalimantan, Java/Bali, Sulawesi, Lesser Sundas, Moluccas, y Irian Jaya), existen 68 especies de aves rapaces diurnas, nueve de las cuales son endemicas a la region. La presion constante sobre los habitats, en particular sobre los bosques de lluvia ricos en madera, como tambien la caza y el uso de pesticidas han tenido un impacto importante en la poblacion y distribucion de estas especies. El estado de 20 especies es un aspecto alarmante debido a que tres especies son consideradas como globalmente amenazadas y otras once como cercanamente amenazadas. El estado de las otras cinco especies es preocupante a nivel local unicamente debido a que tienen una distribucion amplia y son relativamente comunes en otras areas. [Traduccion de Cesar Marquez] Straddling the equator over a distance of 4500 km, the republic of Indonesia has about 1539 bird species. Of these, 381 are endemic (Andrew 1992, 1993) and the country ranks among the five bird- richest in the world (Groombridge 1994). There are 68 species of birds of prey, and many are forest dwellers, occupying many niches, with ^30 species on the largest islands, and at least two or three species on even the smallest oceanic islands (Ap- pendix 1). Many of Indonesia’s species are among the world’s least known (Meyburg 1986) because they are restricted to remote, little known places. Most research on these species has been taxonom- ic and was carried out in the decades before or around the turn of this century. Our knowledge on the biology and ecology on Indonesian raptors has been and is still fragmentary. In this paper, I try to summarize work done so far on raptors in Indonesia and identify gaps in our knowledge, as- sess the status of a number of selected species, eval- uate the current protection system, and give rec- ommendations for future research and protection measures. Although only part of the Indonesian raptor fauna is presumed to be dependent on for- ests, none can survive without some kind of wood- land. Therefore, I have considered both forest and nonforest raptors. Studies, Past and Present The first written scientific report of raptors in Indonesia was made by a merchant, Baron Von Wurmb (1779-82) who described a hawk and the Brahminy Kite ( Haliastur indus). In subsequent centuries, most studies of birds in Indonesia fo- cused on collecting specimens; nevertheless, they produced a relatively accurate distribution map for raptors across the archipelago, with standard an- notated checklists developed for Sulawesi, Lesser 56 March 1998 Raptors in Indonesia 57 Table 1. Recent raptor studies conducted in Indonesia. References Research Subject Taxa 1. Ash 1984, 1993 Autumn migration in NW Bali Migratory raptors 2. Thiollay and Meyburg 1988 Forest fragmentation and raptor conservation on Java Falconiformes 3. van Balen et al. 1993 Conservation status on Java Haliastur indus 4. Meyburg and van Balen 1994 5. van Balen and Meyburg 1994; Conservation on Sulawesi Falconiformes Sozer and Nijman 1995 Conservation, ecology and distribution Spizaetus bartelsi 6. Thiollay 1996a, b Conservation, ecology and distribution on Sumatra Falconiformes Sundas and Moluccas (White and Bruce 1986), and Sumatra (van Marie and Voous 1988). Similar checklists are currently being written for Java, Bali, and Borneo, but locations such as Irian Jaya (e.g., western New Guinea) still need to be studied. Dur- ing the past decade, the study of raptors in Indo- nesia has experienced considerable growth and studies are currently being undertaken on the Gur- ney’s Eagle ( Aquila gurneyi) and Javan Hawk-eagle ( Spizaetus bartelsi ) (N. R0v and J.O. Gjershaug pers. comm.; T. Yamazaki pers. comm.). National Overview Indonesia’s 17 000 islands show a great variety in habitats, influenced by climate, edaphic factors, to- pography, and human interference. On Java and most of the Lesser Sundas, high human population pressure, dating back from ancient times, has had far-reaching effects on natural ecosystems. In the past few decades, high technology and extensive exploitation of natural resources have increased this pressure from people at an alarming rate (Ta- ble 2). Java and Bali. Largely deforested, Java and Bali are essentially a mosaic of paddy fields, cities, and villages in the low lands, with most forest fragments covering the numerous volcanoes on the two is- lands. The most developed forest reserve system in Indonesia is located on these islands. However, due to extreme population pressure, many reserves es- tablished in the first half of this century are small and have probably lost most of their species diver- sity. Because of extreme habitat fragmentation, the endemic Javan Hawk-eagle is considered to be En- dangered (Collar et al. 1994). Development has been most concentrated in open woodland areas, where five raptor species, the Brahminy Kite, Ru- fous-winged Buzzard ( Butastur liventer), Black- winged Kite ( Elanus caeruleus), Changeable Hawk- eagle ( Spizaetus cirrhatus), and Spotted Kestrel (Fal- co moluccensis) , are disappearing (Thiollay and Mey- burg 1988, van Balen et al. 1993, Holmes 1995). Lesser Sundas. Burning, wood collection, and inefficient subsistence agriculture have long been destructive forces to the forests of these islands. The coundess islands of the Lesser Sundas are cov- ered in a parklike landscape of extensive grass- lands, thickets, and wooded river valleys. The most extensive forests are on Sumbawa and north Flores where there are few people. The conservation area network in the Lesser Sundas is insufficient (Sujat- nika et al. 1995), but the typical park landscapes seem to favor a relatively diverse raptor fauna in- Table 2. Land use and the human population in Indonesia (after Sujatnika et al. 1995). Total Area (X10 6 HA) Forest (%) Development Area (%) Population (X10 6 ) Density (per km 2 ) Growth Rate (%) Kalimantan 53.95 74 23 9.10 17 3.5 Irian Jaya 42.20 85 28 1.65 4 3.5 Sulawesi 18.92 59 40 12.52 66 2.0 Sumatra 47.34 49 45 36.51 77 3.0 Java/Bali 13.77 10 72 110.36 801 1.8 Lesser Sundas 8.29 18 49 7.39 89 2.3 Moluccas 7.45 74 38 1.86 24 2.8 58 VAN BALEN Vol. 32, No. 1 eluding Short-toed Eagles ( Circaetus gallicus), Bo- nelli’s Eagles ( Hieraaetus fascialus). Changeable Hawk-eagles, Variable Goshawks ( Accipiter novaehol- landiae), and Brown Goshawks {A. fasciatus) . Sumatra. The larger part of the Sumatran low- land rainforest has been logged or is under a pro- duction regime. Mountain forests are relatively se- cure, though degradation and encroachment on lower slopes is widely reported (Sujatnika et al. 1995). Small islands with distinct races of raptors are found off west Sumatra, notably Nias Island (e.g,, Thiollay 1996b). Separated by mainland Ma- laysia by only a narrow sea strait, Sumatra is an important passage and wintering area for southern bound migratory raptors from northern temperate regions. Kalimantan (Former Dutch Borneo). Until re- cently, extensive rainforest covered most of the is- land, but conversion into plantations, wasteland (notably alang-alang [ Impel ata cylindrica\ grass- fields) , and secondary growth is beginning to dom- inate the landscape. The island’s endemic raptors, the Mountain Serpent-eagle ( Spilornis kinabaluen- sis ) and White-fronted Falconet ( Microhierax lato- frons) are restricted to the northern, non-Indone- sian part of the island. Sulawesi (Former Celebes). Extensive lowland rainforest still covers Sulawesi, although mainly at moderate altitudes. In southwest Sulawesi, the sparse forest is only on high mountain tops, where- as extensive lowlands and hill forests still cover cen- tral Sulawesi. Agriculture, transmigration, and log- ging continue to absorb large areas of lowland for- est. Montane forest is relatively untouched, al- though some encroachment occurs on the lower slopes (Sujatnika et al. 1995). Despite harsh cli- matic conditions and deforestation, Coomans de Ruiter (1947) reported more raptors in southwest Sulawesi than anywhere else. Nevertheless, the for- merly common Black Kite ( Milvus migrans) has de- creased in numbers (Meyburg and van Balen 1994). The status of the three endemic Accipiters is largely unknown, partly because of identification problems. Moluccas, Originally, the numerous islands of Moluccas were covered in tropical rainforest. High quality timber, and local climatic and edaphic con- ditions attracted loggers and led to the develop- ment of plantations of nutmeg and cloves. As a result, lowland forests have rapidly disappeared, and little virgin forest remains. The largest stretch- es of remaining original forest are found on Ser- am, Buru, and Halmahera (Poulsen pers. comm.). These areas support Rufous-necked Sparrowhawks (Accipiter erythrauchen) , Moluccan Goshawks (A. henicogrammus) , and a number of endemic races. Irian Jay a (Former Dutch New Guinea). Irian Jaya comprises the second largest and most pristine region in Indonesia. However, development from mining, logging, resettlement schemes, and infra- structure improvement is proceeding rapidly. Two large forest eagles in this area, the New Guinea Harpy Eagle ( Harpyopsis novaeguineae ) and Gur- ney’s Eagle { Aquila gurneyi ), are presumably still safe, but monitoring is needed to ensure that they continue to do well. Conservation of Indonesian Raptors There are 20 species of raptors that are globally Threatened and Near Threatened with extinction (Collar et al. 1994), or may be classified as such, pending further field research. The number of open woodland and forest edge species that are listed is striking. One of the greatest threats to these raptors is the loss of habitat. Large trees for perches, roosts, and nests are important components of raptor habitat (Janes 1985). Huge islands like Kalimantan and Irian Jaya seem well-forested and secure (Table 2), but present human population growth and habitat conversion may soon alter this. Therefore, man- agement of the existing forest reserve systems needs to be improved without delay. Hunting also poses a serious threat to raptors on Java and most other regions in Indonesia. With the immensely increased number of air rifles accom- panying the nation’s overall improved standard of life, raptors, like many other birds, have been dec- imated, especially near settlements. With more people that can afford the high price and care of pet eagles and hawks, the demand for wild birds is high, despite legal protection of all diurnal raptors. It is not clear if pesticides have played a major role in the decline of Brahminy Kites on Java. Nev- ertheless, a recent decline in the species reported from Laos indicates that the species may be un- dergoing a widespread decline in Southeast Asia (Thewlis et al. 1995). A coincident increase in use of pesticides on Java and Bali in the 1970s (van Balen et al. 1993) and the overall decline of Pond Herons ( Ardeola speciosa ) and egrets ( Egretta spp., van Helvoort 1981) is striking. The ban on 57 va- rieties of organophosphate chemicals since 1986 March 1998 Raptors in Indonesia 59 Table 3. Indonesian raptors of conservation concern, threats to their survival, and research needs. Species Threats 1 Trend RDB Status Research Needs End- emic Taxa Refs. Aviceda jerdoni L2 ?< ?NT monitoring 2 2 Pernis ptilorhynchus 1,2,3 ?= - taxonomy 1 3 Macheiramphus aldnus 1,4,7 ?<(B) - monitoring - 4 Elanus caerulaeus 4,5 «(J,S) - monitoring 1 4, 5 Haliastur Indus ?1, ?4, 5, 6 «(J) - monitoring - 1 Ichthyophaga humilis 1, 2 < ?NT monitoring - 2 Ichthyophaga ichthyaetus 1, ?4, ?5 ?< ?NT monitoring - 2 Accipiter henicogrammus 1, ?3 < p ecology 1 6 Accipiter erythrauchen 1, 2, 3 ?< p ecology 2 6 Accipiter nanus ?3 ?< NT ecology 1 2 Accipiter buergersi 7 ?= DD ecology - 2 Accipiter doriae 1, 2, 3 ?= NT ecology - 2 Butastur liventer 1, ?3, ?4 <(J) NT monitoring - 2 Harpyopsis novaeguineae ?3, 4, 7 ? = V ecology - 2 Aquila gurneyi 1 ?= NT ecology - 2 Spizaetus cirrhatus 1, 4, ?5 <(J) - monitoring 2 4 Spizaetus bartehi 1, 3, 4, 7 ?« E ecology 1 2 Spizaetus nanus 1,3 ?< V ecology 1 2 Spizaetus lanceolatus 1,2 ? = NT ecology 1 2 Falco moluccensis 4, ?5 <(J,S) - monitoring 2 5 1 Threats: 1 — habitat loss; 2 — habitat degradation; 3 — forest fragmentation; 4 — hunting; 5 — pesticides; 6 — improved sanitation; 7 — low density/ small population. Trend: < — decrease; << — dramatic decrease; = — (presumably) stable; J — Java; S — Sulawesi; B — Borneo. RDB Status or Global Conservation Status (after Collar et al. 1994): E — Endangered; V — Vulnerable; DD — data-deficient; NT — Near Threatened; ? — possible candidate. Refs.: 1 — Van Balen et al. 1993; 2 — Collar et al. 1994; 3 — del Hoyo et al. 1994; 4 — van Balen 1994; 5 — Holmes 1995; 6 — M. Poulsen and D. Permiasa pers. comm. (Whitten et al. 1996) has not yet resulted in an increase in either of these species. Decreased food availability associated with the growing human population and increased sanita- tion have perhaps affected only a few, scavenging raptors (e.g., Brahminy Kite in large cities on Java [van Balen et al. 1993]). Research There are several specific areas where further re- search is necessary on Indonesian raptors (Table 3) . It is not yet known what the long-term impact of logging will be on raptor populations in Indo- nesia. Selective logging leaves a relative high forest cover which supports widespread populations of raptors. Several species of raptors in Indonesia are so poorly studied that their taxonomic status may change as information on them increases (Sujat- nika et al. 1995) . A number of island races, some vulnerable to extinction, may prove to be full spe- cies, such as Accipiter griseogularis comprising A. no- vaehollandiae obiensis of Obi Island, A. n. griseogularis of Bacan and Halmahera Islands, and A. n. mortyi of Morotai Island (White and Bruce 1986); Spizae- tus cirrhatus floris of Sumbawa and Flores Islands, Spizaetus nanus stresemanni of Nias Island, and Per- nis p. ptilorhynchus of Java and P. p. torquatus of Su- matra, Borneo, and peninsular Malaysia (del Hoyo et al. 1994) . Only fragmentary information is avail- able on migration routes of northern migratory raptors. Bali Strait (Ash 1984, 1993), the eastern part of the Java Sea (J.H. Becking pers. comm.), and the Puncak pass in west Java are known as pas- sage points, but huge stretches in between and be- yond need to be investigated. Recommendations and Conclusions Future raptor studies in Indonesia should focus on the causes of declines of formerly common open woodland species, habitat use and measure- ment of home ranges of raptors, effects of habitat degradation and forest fragmentation, and migra- tion routes of raptors over Indonesia. Large areas of Sulawesi, the Moluccas, Kaliman- 60 VAN BALEN Vol. 32, No. 1 tan, and other areas virtually are unexplored. The possible addition of additional natural areas that might be added to the current network is urgent. Raptors are either persecuted because they are suspected of killing domestic fowl or because they are sought as pets, especially the rarer ones. Public education needs to be undertaken at all levels through the use of posters, television spots, and newspaper articles to increase awareness of the im- portance of raptors. Local bird-watching clubs should be supported in monitoring the passage of migratory raptors as well resident raptors. Captive breeding has become a popular issue, and often too quickly considered the most impor- tant solution to preserve a particular rare species. The Javan Hawk-eagle has recently been indicated as a candidate for such a program. Before steps are taken towards establishing a captive population, further consideration should be given to the con- tribution of such a program to conservation of the species, its effect on the wild population, and the availability of secure habitat for reintroduced birds. A steering committee of local and international experts in the field of raptor ecology and conser- vation should supervise any future conservation ac- tion. Major steps have already been made by the adoption of an Action Plan by the “Focus Group for Javan Hawk-eagle Conservation” and the estab- lishment by the State Minister of Environment in January 1997 of the “Working Group for Manage- ment of Indonesian Raptor Areas.” Both initiatives are cooperative activities of the State Ministry of Environment, the Directorate General of Forest Protection and Nature Conservation (PHPA) and the Indonesian Institute of Sciences (LIPI) involv- ing various national and international organiza- tions, including BirdLife International-Indonesian Programme. The commitment of Indonesia’s Government to preserve its local wildlife is attested to by the adop- tion of two of its birds of prey as the mascot of Jakarta (Brahminy Kite) and as a national symbol (Javan Hawk-eagle) . The former is Indonesia’s most ubiquitous raptor and the second is one of its rarest birds of prey. Acknowledgments I would like to thank the Peregrine Fund and Dr. Rick Watson in particular for financial support to attend the symposium. Paul Jepson, Michael Poulsen, Prof. C.M. White and an anonymous reviewer gave useful comments on drafts of the paper, for which I am very grateful. Literature Cited Andrew, P. 1992. The birds of Indonesia. A checklist (Peters’ Sequence). Indonesian Ornithological Soci- ety, Jakarta, Indonesia. . 1993. The birds of Indonesia, Kukila checklist No. 1. Additions, corrections and notes — 1. Kukila 6: 47-52. Ash, J.S. 1984. Bird observations on Bali. Bull. Br. Orni - thol. Club 104:24-35. . 1993. Raptor migration on Bali. Forktaii 9:3-11. Collar, N.J., M.J. Crosby and A.J. Stattersfield. 1994. Birds to watch: the world list of threatened birds. BirdLife International, Cambridge, U.K. Coomans de Ruiter, L. 1947. Roofvogelwaarnemingen in Zuid-Celebes. Limosa 20:213-229. del Hoyo, J., A. F. i.i. iott and J. Sargatal. 1994. Hand- book of the birds of the world. Vol. 2. Lynx Edicions, Barcelona, Spain. Groombridge, B. [Ed.]. 1994. Biodiversity data source- book. World Conservation Monitoring Centre, World Conservation Press, Cambridge, U.K. Holmes, D.A. 1995. Editorial. Kukila 7:85-90. Janes, S.W. 1985. Habitat selection in raptorial birds. Pages 159-188 in M.L. Cody [Ed.], Habitat selection in birds. Academic Press, Orlando, FL U.S.A. MacKinnon, J. and J. Wind. 1980. Birds of Indonesia. FAO, Bogor, Indonesia. Meyburg, B.-U. 1986. Threatened and near-threatened diurnal birds of prey of the world. Birds of Prey Bull 3:1-12. and S. van Ba i.f. n. 1994. Raptors on Sulawesi (In- donesia) : the influence of rainforest destruction and human density on their populations. Pages 269-276 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. Sozer, R. and V. NlJMAN. 1995. Behavioral ecology, dis- tribution and conservation of the Javan Hawk-eagle Spizaetus bartelsi Stresemann, 1924. Verslagen en Technische Gegevens No. 20. Institute of Systematics and Population Biology, Univ. Amsterdam, The Neth- erlands. Sujatnika, P., P.R. Jepson, T. Soehartono, M.J. Crosby and A. Mardiastuti. 1995. Conserving Indonesian biodiversity: the endemic bird area approach. PHPA and BirdLife International-Indonesia Programme, Ja- karta, Indonesia. Thewlis, R.M., J.W. Duckworth and G.Q.A. Anderson. 1995. Ornithological records from Laos. Forktaii 11: 47-100. Thiollay, J.-M. 1996a. Rainforest raptor communities in Sumatra: the conservation value of traditional agro- forests. Pages 245-260 in D.M. Bird, D.E. Varland and J.J. Negro [Eds.], Raptor adaptations to human influ- enced environments. Academic Press, London, U.K. March 1998 Raptors in Indonesia 61 . 1996b. The raptor community of Nias Island, Sumatra: survey and conservation. Kukila 8:113-116. and B.-U. Meyburg. 1988. Forest fragmentation and the conservation of raptors: a survey on the is- land of Java. Biol. Conse rv. 44:229—250. van Balen, S. 1994. The status and conservation of birds of prey in the Sondaic and Wallacean regions of In- donesia. Pages 245-254 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey and Owls, London, U.K. and B.-U. Meyburg. 1994. The Java Hawk-eagle Spizaetus bartelsv. results of recent research on distri- bution status and ecology. Pages 89-92 in B.-U. Mey- burg and R.D. Chancellor [Eds.], Raptor conserva- tion today. World Working Group for Birds of Prey and Owls, London, U.K. , I.S. Suwelo and D.S. Hadi. 1993. The decline of the Brahminy Kite Haliastur indus on Java. Forktail 8:83-88. van Helvoort, B. 1981. Bird populations in the rural ecosystems of West Java. Student Report No. 560. Ag- ricultural Univ., Wageningen, The Netherlands. VAN Marle, J.G. and K.H. Voous. 1988. The birds of Sumatra. B.O.U. Check-list 10. British Ornithologists’ Union, London, U.K. VON WuRMB, F. 1779-1782. Verhandelingen van het Ba- taviaasche Genootschap van Kunsten en Wetenschap- pen. Vol. 1-4. White, C.M.N. and M.D. Bruce. 1986. The birds of Wal- lacea (Sulawesi, the Moluccas and Lesser Sunda Is- lands, Indonesia): an annotated check-list. B.O.U Check-list 7. British Ornithologists’ Union, London, U.K. Whitten, T., R.E. Soeriaatmadja and S.A. Afiff. 1996 The ecology of Java and Bali. Periplus, Singapore. Received 16 October 1996; accepted 11 November 1997 62 VAN BALEN Vol. 32, No. 1 Appendix 1. Distribution of raptors in Indonesia. Sequence, nomenclature, and subregional boundaries follow Andrew (1992). R 1 S Distribution Habitat Osprey ( Pandion haliaetus) <> Jerdorfs Baza (Aviceda jerdoni) < Pacific Baza ( Aviceda subcristata) > Black Baza ( Aviceda leuphotes) < Long-tailed Buzzard ( Henicopernis longicauda) P Oriental Honey-buzzard ( Pernis ptilorhynchus ) < Barred Honey-buzzard ( Pernis celebensis) < Bat Hawk ( Macheiramphus alcinus)

Whistling Kite ( Haliastur sphenurus ) > Brahminy Kite (Haliastur indus) <> White-bellied Sea-eagle ( Haliaeetus leucogaster) <> Lesser Fish-eagle ( Ichthyophaga humilis) < Grey-headed Fish-eagle ( Ichthyophaga ichthyaetus ) < Short-toed Eagle ( Circaetus gallicus ) < Crested Serpent-eagle ( Spilomis cheela) < Sulawesi Serpent-eagle ( Spilomis rufipectus ) E Spotted Harrier ( Circus assimilis ) > Pied Harrier ( Circus melanoleucos) < Western Marsh-harrier ( Circus aeruginosus) < Eastern Marsh-harrier (Circus spilonotus)

Crested Goshawk ( Accipiter tiivirgatus) < Sulawesi Goshawk ( Accipiter griseiceps ) E Shikra (Accipiter badius) < Chinese Goshawk (Accipiter soloensis) < Spot-tailed Goshawk (Accipiter trinotatus) E Brown Goshawk (Accipiter fasciatus) > Grey Goshawk (Accipiter novaehollandiae) > Black-mantled Goshawk (Accipiter melanochlamys) P Moluccan Goshawk (Accipiter henicogrammus) E Grey-headed Goshawk (Accipiter poliocephalus) P Japanese Sparrowhawk (Accipiter gularis) < Besra (Accipiter virgatus ) < Small Sparrowhawk (Accipiter nanus ) E Collared Sparrowhawk (Accipiter cirrocephalus ) > Rufous-necked Sparrowhawk (Accipiter erythrauchen ) E Vinous-breasted Sparrowhawk (Accipiter rhodogaster) E Meyer’s Goshawk (Accipiter meyerianus ) > Chestnut-shouldered Goshawk (Accipiter buergersi) P Doria’s Hawk (Accipiter doriae) P Rufous-winged Buzzard (Butastur liventer) < Grey-faced Buzzard (Butastur indicus) < Common Buzzard (Buteo buteo) < New Guinea Eagle (Harpyopsis novaeguineae ) P Black Eagle (Ictinaetus malayensis) < Spotted Eagle (Aquila clanga ) < Gurney’s Eagle (Aquila gurneyi) P Wedge-tailed Eagle (Aquila audax) > Bonelli’s Eagle (Hieraaetus fasciatus) < Booted Eagle (Hieraaetus pennatus) < R/M JB T S K C M I C R - - S K C M - LR/SR/SS R - T - - c M I LM/R/SR/SS M J - s - - - - SS/A R I LR/SR R/M JB T s K c M - LR/FE/SS R - - - - c - - LR/FE R - - s K c - - LM/LR/FE R JB T s K c - I LM/FE/G/SS R/M - T s - c M I LM/FE/G/SS R I F/G/SS R JB T s K c M I F/C/G/SS R JB T s K c M I F/C R - ■ - s K c - - F/C R J - s K c - - F R JB T LM/FE/G/SS R JB - s K - - - LR/SR/SS R - - - - c - - LR/SR/G/SS R/M - T - - c - - G/SS V - - - K - - - F/G/SS V - - s - - - - F/G M - - s - - - I F/G M I F/S R J - s K - - - LR R - - - — c - - LR M — - s — — — — LR M JB T s K c M I LM/FE/LR R - - - - c - - LR/SR R - T - - c M I LR/SR R - T - - - M I F/LR/G/FE/SS R I MR R - - — - - M - LR R I LR/FE/SS M JB T s K c - - LM/LR/FE/SS R JB T s K - - - LR/SR/MR R - - - ■ c - - LR/SR/MR R - - - - - - I LR R - - - - - M - LR R - - - - c - - LR R - - — - — M I LR R - - - - — - I SR R I LR/SR R J - - - c - - LM/LR/SR/G/SS M J p s - c M I FE/LM/LR/G/SS M JB FE/G/SS R I LR/SR/MR/FE R JB - s K c M - LR/SR V - - s - - - - C R - - - - - M I C/F/LR/G R I F/G R - T LM/FE V J LR March 1998 Raptors in Indonesia 63 Appendix 1. Continued. R 1 S Distribution Habitat Litde Eagle ( Hieraaetus morphnoides ) > R - - - - - M I LR/SR Rufous-bellied Eagle ( Hieraaetus kienerii) < R JB T S K C M - LR Changeable Hawk-eagle ( Spizaetus drrhatus) < R JB T S K - - - LM/LR/FE/G/SS Javan Hawk-eagle ( Spizaetus bartelsi) E R J LR/SR Sulawesi Hawk-eagle ( Spizaetus lanceolatus) E R - - - - c - - LR/SR Blyth’s Hawk-eagle ( Spizaetus alboniger) < R - - s K - - - SR/MR Wallace’s Hawk-eagle ( Spizaetus nanus ) < R - - s K - - - LR Black-thighed Falconet ( Microhierax fringillarius) < R JB - s K - - - LR/FE/SS Brown Falcon ( Falco berigora) > R I LR/SR/MR/G/SS Eurasian Kestrel {Falco tinnunculus) < M - - s - - - - G/SS Spotted Kestrel {Falco moluccensis ) E R JB T - K c M I G/SS Australian Kestrel {Falco cenchroides ) > V J T - - - M I G Eurasian Hobby {Falco subbuteo) < V J T FE/G/SS Oriental Hobby {Falco severus) <> R JB T s K c M I LR Australian Hobby {Falco longipennis ) > R/M - T - - c M I FE Peregrine Falcon {Falco peregrinus) <> R/M JB T s K c M I FE/G 1 R (Range): < — western extension; > — eastern extension; E — endemic Indonesia; P — found in PNG. (S) Status: R — resident; M — migrant; V — vagrant. Distribution: JB — -Java/Bali; T — Lesser Sundas; S — Sumatra; K — Kalimantan; C — Sulawesi; M — Moluccas; I — Irian Jaya (after Andrew 1992, 1993). Habitat: LM — lowland monsoon forest; LR — lowland rainforest; SR — submontane rainforest; MR — montane rainforest; C— coastal and mangrove forest; F — freshwater bodies; S — swamp forest; FE — forest edge; SS — secondary forest/ scrub; G — grass and open woodland; A— agricultural land (after MacKinnon and Wind 1980, own observations). J. Raptor Res. 32(l):64-73 © 1998 The Raptor Research Foundation, Inc. STATUS AND CONSERVATION OF RAPTORS IN AUSTRALIA’S TROPICS Nick Mooney Parks and Wildlife Service, GPO Box 44A, Hobart 7001, Tasmania, Australia Abstract. — All of Australia’s 34 raptors are found in the tropics. No full species and only one subspecies, an island endemic owl, are extinct. All of Australia’s three threatened, diurnal species are endemic to the continent. One, the Vulnerable Red Goshawk ( Erythrotriochis radiatus), is endemic to Australia’s tropical forests and is under threat from loss of habitat, persecution, and egg collecting. Conservation efforts include legal protection, education, and keeping nest sites secret. A second species, the rare Square-tailed Kite ( Lophoictinia isura) is widely distributed and, except for clearing of woodland, threats are not obvious. Many raptors from arid areas, including the endemic Grey Falcon ( Falco hypoleucos), “winter” in tropical woodlands. For adequate conservation, critical habitats of the Grey Falcon must be identified. Grey Falcons should be helped in the long term by the anticipated reduction of rabbits in arid Australia by rabbit calicivirus disease, but widespread clearing of tropical woodlands for agriculture continues as does local, heavy use of pesticides. Although no species of owls are threatened, five sub- species are; two are subspecies of the endemic Rufous Owl ( Ninox rufa, one rare and one insufficiently known) and two are subspecies of the Masked Owl (Tyto novaehollandiae, both insufficiently known). Threats include loss of critical habitat to fire and agriculture. On Christmas Island, the small populations of endemic subspecies of the Brown Goshawk ( Acdpiter fasciatus ) and Moluccan Hawk-owl (N. squamipila) are Vulnerable and threatened by loss of habitat to urbanization and formerly mining. Besides legal protection, conservation efforts have included education and habitat preservation. The tropical Eastern Grass-owl ( T. longimembris ) is secure although some populations are under pressure from agriculture (including rodenticides) and urbanization. Coastal, tropical raptors appear secure even though per- manent reductions in some populations have been caused by agriculture, urbanization, and coastal developments such as aquaculture and marinas. Key Words: tropical raptors; Australia; conservation; status. Estado y conservacion de las aves rapaces en los tropicos de Australia ReSUMEN. — Todas las 34 especies de aves rapaces de Australia se encuentran en los tropicos. Una de las tres especies amenazadas de Australia, Erythrotriorchis radiatus es endemico a los bosques tropicales de Australia y esta amenazada por la perdida de habitat, persecucion, y saqueo de huevos. Los esfuerzos de conservacion han incluido su proteccion legal, educacion, y proteccion de nidos. Una segunda especie Lophoictinia isura esta ampliamente distribuida, excepto por la deforestacion, las demas amenazas no son obvias. Varias aves rapaces de regiones aridas incluyendo al endemico Falco hypoleucos emigran hacia los bosqnes tropicales. Desde el punto de vista de la conservacion, los habitats criticos de F. hypoleucos deben de ser identificados debido a que la deforestacion para la agricultura continua al igual que el uso intensivo de pesticidas. Aunque ningun buho ha sido identificado como amenazado, en Australia, dos raras y poco estudiadas subespecies del endemico Ninox rufa y dos subespecies de Tyto novaehollandiae estan amenazadas en las areas tropicales. Las amenazas incluyen la perdida de habitats criticos debido al fuego y a la agricultura. En la Isla Christmas, las reducidas poblaciones de Acdpiter fasciatus y de N. squamipila estan amenazadas debido a la perdida de habitat por actividades mineras y el desarrollo urbanistico. Aparte de la proteccion legal, los esfuerzos de conservacion han incluido educacion y proteccion del habitat. Aunque aparentemente estables algunas poblaciones de T. longi- membris estan bajo presion debido a la agricultura (incluyendo el uso de rodenticidas) y el desarrollo urbano. Aunque las poblaciones de aves rapaces costeras estan estables, estas han tenido continuadas reducciones causadas por la agricultura, la urbanizacion, y por desarrollos costeros tales como proyectos de acuacultura y marinas. [Traduccion de Cesar Marquez] 64 March 1998 Status of Tropical Australian Raptors 65 Background Australia is a continent with an area totaling about 7.7 million km 2 . It includes a few small is- lands, one of which is Christmas Island in the In- dian Ocean, approximately 1400 km northwest of Australia near the Indonesian island of Java. Aus- tralia is the driest continent and less than 15% is covered by tropical forests located on the north and northeastern coasts. Although people have slowly modified these forests using fire over at least 60 000 yr, it has only been in the last 200 yr since Europeans arrived that this change has ac- celerated. While the size of the continent has shel- tered most raptors from threats to tropical forests, small islands have not escaped these effects. The wet, tropical forests of Australia show distinct sea- sonality but have long-term ecological stability. The occurrence of irregular, severe inland droughts make the continent’s biogeographv unique (Nix 1972, Flannery 1994). This instability has resulted in many arid and semiarid birds, rap- tors included, becoming migrants during drought periods with many showing both inland-coastal and south-north movements (Baker-Gabb and Fitzherbert 1989). These local movements may be small in relation to the size of the continent but they are still large in absolute terms and their ap- parent dispersive rather than routed nature makes their study difficult. Another unusual movement is the irruptive ten- dency of a few raptor species. This type of move- ment usually follows heavy rains marked by increas- es in rodent numbers, and rapid breeding in pop- ulations of Letter-winged Kites (Elanus scriptus) and Black Kites ( Milvus migrans) which then disperse when dry conditions return and rodent numbers crash (Hollands 1977, Lavery and Johnson 1993). This paper examines raptors in wet tropics (a strongly seasonal environment) and dry tropics (an environment stable in the short to medium term but unstable in the longer term). It is important to make this distinction because tropical raptors may be largely dependent on a system with long- term stability and they may be slow to adapt to en- vironmental changes. The Australian wet tropics contain both open and closed forests, including rainforest on the con- tinent’s northeast coast. Mainly coastal, these for- ests extend from nearly half-way down the east coast and all the way to the northwest corner of Australia. Tropical areas also spread inland along permanent rivers and wetlands and include some areas of highlands such as the Atherton tablelands. Adjacent islands off these areas including Christ- mas Island are also wet and support rainforests. The dry tropics are comprised of mainly savanna woodlands and a variety of permanent and tem- porary waterways and wetlands. They are located south of the wet tropics but north of a line extend- ing across the continent from the mid-east to the mid-upper west coast from about Brisbane to about Broome (Fig. 1). Raptors are very mobile birds and most species are able to utilize a variety of habitats. Therefore, I was cautious when assigning species to habitats and used broad definitions. Use of Australia’s Tropics by Raptors Presence and Core Range. All 34 species of Aus- tralian raptors are found in the tropics. Of Austra- lia’s 10 endemic raptor species, only two, the monotypic Red Goshawk ( Erythrotriorchis radiatus) and Lesser Sooty-owl ( Tyto multipunctata) , are en- demic to the tropics as are four and eight subspe- cies of Falconiformes and Strigiformes, respective- ly. One other monotypic species, the Square-tailed Kite ( Lophoictinia isura ) , six subspecies of Falconi- formes, and six of Strigiformes have large portions of their ranges in the tropics (Blakers et al. 1984, Debus and Czechura 1988b, Marchant and Higgins 1993). Australia is relatively close to Melanesia and some raptors interchange between the two includ- ing Swamp Harriers ( Circus approximans) , Austra- lian Hobbies ( Falco longipennis) , and Nankeen Kes- trels (F. cenchroides) . Numbers of raptors that inter- change are not known but neither location is known to be a major wintering area for interchang- ing species (Baker-Gabb and Fitzherbert 1989). Wet vs. Dry Tropics. More species of Strigifor- mes rely on the wet tropics than do Falconiformes. Seven species of Strigiformes are endemic while only three species of Falconiformes are exclusively found in the wet tropics. All told, 10 of Australia’s 19 species and subspecies of Strigiformes but only 10 of 28 species and subspecies of Falconiformes are either endemic to or a large segment of their core ranges lies within the wet tropics. Conversely, the dry tropics are more important to the majority of Australia’s tropical Falconiformes. Six species are endemic to the dry tropics while this area has no endemic owls. 66 Mooney Vol. 32, No. 1 I CO 2 O J-H O w & ^ 2 £ 2 w o 2 O o m S (j Q § « £ 3 £ g o owoqShH 0^g2Sh» fs5 O ffi O H < ffi CQ j h B <1 U h CO o CL o QC LD CO O 0_ O DC h- > DC Q O DC (D O cr LD >- LD CO LD > DC LU CO LD DC DC O —> < CO DC LU I- I- < Q_ D . >* w pq l: o pq < u h Figure 1. Key raptor wintering grounds (after Marchant and Higgins 1993), reserves, and movement patterns (after Nix 1972, Baker-Gabb and Fitzherbert 1989) in the Australian tropics. March 1998 Status of Tropical Australian Raptors 67 Movements. In the Australian tropics, there are 13 resident Falconiformes, 10 regular “winter” mi- grants, and one species, the Black-breasted Buz- zard ( Hamirostra melanosternon) , that moves to the tropics and other coastal areas in response to ir- regular, inland droughts. Although two regular mi- grants, the Black-shouldered Kite ( Elanus axillaris) and the Black Kite are also irruptive, only the Let- ter-winged Kite appears to be adapted for such movements through its colonial and early breeding (Marchant and Higgins 1993). Interpreting move- ments for many species is difficult because their extensive ranges include populations which are resident while others are migratory on either a reg- ular or irregular basis. This difficulty does not exist for Australia’s Strigiformes. Although there are sev- eral irruptive species, the Barn Owl ( Tyto alba), possibly the Grass Owl (T. longimmibris) , and the Southern Boobook ( Ninox novaeseelandiae ) on Tas- mania seem to be at least partially migratory (Schodde and Mason 1980, Blakers et al. 1984, Hollands 1991). After becoming concentrated in the best breed- ing habitats in spring, populations of some species barely leave inland breeding areas in the heat of summer (e.g., Wedge-tailed Eagles [Aquila audax \ and Black Falcons [jF subniger] ; Marchant and Hig- gins 1993). Other species such as Black Kites and Brown Falcons (F berigora) make partial migrations forming concentrations in spring to undertake northward and coastward through summer and au- tumn. Other species like the Nankeen Kestrel and Spotted Harrier ( Circus assimilis) disperse in early winter and rapidly return to breeding areas through late winter. The Pacific Baza ( Aviceda sub- cristata) migrates altitudinally, making it inland and coastal at the same time. This is best seen in the north-south migration of the Swamp Harrier w r here virtually the entire breeding population is in the far southeast (mostly in Tasmania) while non- breeding immatures are concentrated in grassy woodlands to the north. Habitats and Areas of Raptor Concentration. A higher proportion of Strigiformes (9 of 17 species) than Falconiformes (7 of 28 species) rely on trop- ical closed forests. Conversely, woodlands support a higher proportion of Falconiformes (13 of 28 species) than Strigiformes (6 of 7 species) . During the dry season and during droughts, tr opical woodlands become particularly important for both residents and wintering migrants. Inland grassy, w r oodland areas of the Kimberleys and the lower (coastward) catchments of the larger rivers around the Gulf Region are particularly important since they support the greatest raptor diversity and share the greatest abundance (Fig. 1). Grassy woodlands support mainly Brown Falcons, Swamp Harriers, Spotted Harriers, Black Kites, and Black- shouldered Kites while open forests along perma- nent waterways support Red Goshawks, Australian Hobbies (F longipennis) , Peregrine Falcons (F per- egrinus) , and Grey Falcons (F. hypoleucos ) . Open for- ests throughout the area support many Brown Gos- hawks ( Accipiter fasciatus) and Collared Sparrow- hawks ( Accipiter cirrocephalus ) . During this autumn/ winter dry period, White-bellied Sea Eagles (Ha- liaeetus leucogaster) and Ospreys ( P and, ion haliaetus ) sometimes travel far inland up permanent rivers (Blakers et al. 1984, Marchant and Higgins 1993) and Brahminy Kites ( Haliastur indus ) disperse into the Torres Strait (Draffon et al. 1983). Other areas of the tropics are important as win- tering grounds. For example, the developed grass- lands of the Darling Downs area in southeastern Queensland support high densities of wintering Nankeen Kestrels (Baker-Gabb 1996) and any ar- eas of open forest where small passerines are com- mon are used by Square-tailed Kites (Debus and Czechura 1988b). Except for a few large areas of rainforest along the north coast, most of Australia’s tropical forests and woodlands are fragmented, with indistinct bor- ders caused by combinations of soil fertility and fire regimes (Flannery 1994). Consequently, in most of the Australian tropics, raptors normally as- sociated with closed forests such as Rufous Owls (N. rufa) and Grey Goshawks ( Accipiter novaehollan- diai) , may range far along rivers and between wet- lands using riparian forests. Status Australia’s small tropical islands have dispropor- tionate numbers of threatened (Rare, Vulnerable, or Endangered, after Garnett 1992) or insufficient- ly known species. The only extinct Australian rap- tor, the Lord Howe Island Boobook (N. n. albana), was endemic to an island as are five of Australia’s seven Endangered or Vulnerable raptors. The highest proportion of Australia’s threatened or in- sufficiently known raptors are those that rely on forests, particularly tropical forests (four of five Fal- coniformes and six of seven Strigiformes) . Austra- lia’s tropical woodlands contain most threatened Falconiformes (three of four species) and closed, 68 Mooney Vol. 32, No. 1 wet tropical forests contain most threatened Stri- giformes (four of seven species) . Nearly all of Australia’s threatened and insuffi- ciently known raptors are endemic to or have their core ranges in the tropics (three of five Falconi- formes and five of seven Strigiformes; Raker-Gabb 1994). Three species of Strigiformes are classified as insufficiently known most likely because they are difficult to survey (Clark and Mikkola 1989, Gar- nett 1992). In reality, many species of raptors re- garded as secure need further surveys and actually should be classified as insufficiently known. Of Australia’s four threatened, endemic and monotypic species, only one, the tropical Red Gos- hawk, is Vulnerable (IUCN 1994). The other two, the largely tropical Square-tailed Kite and the semi- arid Grey Falcon, are both Rare. All three of the species have naturally low population densities (Garnett 1992, Marchant and Higgins 1993). For example, the current Red Goshawk population consists of about 350 pairs as compared to its his- toric population of about 440 pairs spread over nearly 1 million ha. Its population decline has been more the result of a range contraction rather than density reduction (Aumann and Baker-Gabb 1991b, Debus 1993). Conversely, the Vulnerable Brown Goshawk (Ac- dpiterf natalis ) and Moluccan Hawk-owl { N. squam- pila natalis ), both endemic to Christmas Island, have high population densities but very limited dis- tributions. There are currently an estimated 50- 100 pairs of goshawks and about 600 pairs of owls on the 13 650 ha island (Hill 1996). The goshawk’s naturally small and very tame population is Vul- nerable in every sense but the owl has the advan- tage of nocturnal seclusion. There is some ques- tion as to the identity of the owl and whether the goshawk is simply a subspecies of the Grey Gos- hawk (Debus 1994a). These taxonomic questions must be settled in which case more attention may need to be given to the security of these isolated raptors. Threats and Conservation Precaution and Prevention. Threats to Austra- lia’s tropical raptors are similar to those which threaten raptors in other countries (e.g., Thiollay 1994). However, Australia has a great advantage in that much of the management necessary to pre- serve these species is preventative. Loss of Habitat — Agriculture. Both productivity and survival can be affected by loss of critical hab- itat (Newton 1979). In the Australian tropics, this loss has been mosdy due to agriculture. Draining of wetlands is an important issue. The Convention on Wedands of International Importance Especial- ly as Waterfowl Habitat (the Ramsar Convention, Davis 1994) lists several large Australian tropical wetlands for conservation but most are unprotect- ed. However, wetlands have been successfully re- stored by deliberate management (e.g., Anony- mous 1994) and changing public attitudes may give more such opportunities. Like many species, tropical raptors are heavily reliant on remnant wet- lands during the dry season or droughts. Acceler- ated destruction is potentially harmful both to common species, such as the White-bellied Sea-ea- gle, Brahminy Kite, Whisding Kite {Haliastur sphen- urus ) and Barking Owl ( N . connivens ), and to al- ready Vulnerable species, such as the Red Gos- hawk. Perhaps more than any other habitat, Australia’s woodlands have suffered enormously since Euro- pean colonization. In temperate areas, 80% has been cleared (Robinson and Traill 1996) and rapid inroads are being made into tropical woodlands for agriculture, especially in southern Queensland (Anonymous 1996). Square-tailed Kites, Red Gos- hawks and Pacific Bazas are the species most at risk from this deforestation. Dry tropical regions are also disappearing due to the development of irrigation in desert areas. This practice has the potential to increase saltation and change tiooding regimes with widespread ef- fects on many thousands of km 2 . Extensive mono- culture crops are susceptible to pests requiring the use of pesticides (insecticides and rodenticides) which may have consequences for raptors. How- ever, Australia’s National Landcare Program (McLennan 1996) is slowing this and other large- scale land degradation problems such as saltation and erosion. Loss of Habitat — Forestry. Forestry has had lim- ited effects on Australia’s tropics but the increasing demand for woodchips from native hardwood for- ests may change this. Plantations of exotic pines make areas unsuitable for most raptors ( acdpiters occasionally use them; Marchant and Higgins 1993) and, even if the original forest is retained, often the structure and age is so altered that rap- tors can no longer rely on them. This is a problem particularly for large owls needing large tree hol- lows such as the Rufous Owl ( N , rufa), Powerful March 1998 Status of Tropical Australian Raptors 69 Owl (N. strenua) and Sooty Owl ( T tenebricosa) (De- bus 1994b). Codes of Practice have been established for most Australian forest operations (Anonymous 1993) and contain protocols and recommendations based on specific research (e.g., Mooney and Hold- sworth 1991, Brereton and Mooney 1994). In Tas- mania, for example, state codes regulating every operation selling timber has been effective in con- serving nesting Wedge-tailed Eagles (Mooney and Taylor 1996). Survey techniques must be improved to make pre-logging planning effective (Kavan- augh and Peake 1993, Mooney 1996). Loss of Habitat — Urbanization. The penchant of Australians to live on coastal and subcoastal resi- dential blocks is a further local threat to raptors. This type of land use is becoming increasingly de- structive to tropical habitats in Queensland and New South Wales and brings with it all the hazards of people beyond the initial alteration of habitats (e.g., pets, persecution, egg collecting, overhead wires, vehicles, and fuel reduction burns). These rural-residential developments are particularly problematic for large owls and aggressive raptors, the latter which occasionally attack stock. Such ur- banization should be more conservatively planned and education programs aimed at reducing peo- ple’s expectations of what they can do in these en- vironments should be undertaken (Wood 1996). Living With Wildlife Programs (Anonymous 1988, Temby 1992) should be more aggressively promot- ed by wildlife authorities. Loss of Habitat — Other Development. The clearing of mangroves and reclamation of coastal wetlands for aquaculture, marinas, and urban areas are growing problems, most likely to affect Brah- miny Kites and White-bellied Sea-eagles. Ospreys in Australia, as elsewhere, seem remarkably tolerant of humans and have adapted to many artificial nesting sites (Marchant and Higgins 1993, Ewins 1996). Mining is potentially a problem for raptors through its resulting habitat loss and pollution. For example, 25% of Christmas Island was badly de- graded from mining (Hill 1996) and a large ura- nium mine (Ranger Mine) exists in Kakadu Na- tional Park, Australia’s premier tropical reserve. Loss of Habitat — Fire. Fire is more a qualitative problem than other more recent threats to habitat; closed forest is turned to open forest, open forest is turned to woodlands and woodland to grassland. Fire is a fundamental ecological shaper in Austra- lia’s tropics (Flannery 1994). Although some spe- cies directly benefit from it, such as Black Kites, Whistling Kites, and Brown Falcons that forage in front of fires (Marchant and Higgins 1993), fire occasionally kills nestling raptors (Aumann and Ba- ker-Gabb 1991a). However, more serious is fire’s attritive affects on remnants of closed forest that are so important to raptors such as the Rufous Owl. Not only can traditional nest trees be burned and nest stands reduced in size but they can also be degraded by weeds and introduced predators (Ba- ker-Gabb 1994). Fire is a traditional and necessary land management tool in much of Australia (Flan- nery 1994) but it is not always the right took Disturbance. Disturbance can have severe effects on raptor productivity. Tourism is now one of the most important industries in the Australian tropics and amongst raptors its disturbing effects are most felt by breeding White-bellied Sea-eagles on large wetlands and coastal rivers which are favorite tour stops. The ground-nesting Grass Ow4 may be at spe- cial risk from tourism (Gamauf 1994). National park authorities need to consider temporary zon- ing restrictions and tour operators should develop other tours that do not come into close proximity with nesting owls. There is a great deal of oppor- tunity for innovation in such ecotourism. Inexpe- rienced photographers can also cause nesting fail- ures, possibly effecting marginal populations (Gar- nett 1992). Loss of Prey. The rapid spread of rabbit calici- virus disease is expected to decimate populations of rabbits in Australia (Coman 1996). Rabbits do occur in the dry tropics (Williams et al. 1995) but in far lower densities than in more southern regions. Hence, they are far less important in the diets of tropical raptors. Species most likely to be affected are Wedge-tailed Eagles and Little Eagles (Hieraaetus morphnoides ) , but the effects will prob- ably be small and restricted to locally diminished productivity. The effect of this disease on raptor populations is being monitored both locally and nationwide using a road transect survey called Bird of Prey Watch (Baker-Gabb 1996). In the long term, the removal of rabbits should allow vegeta- tion to recover in large areas increasing its value for wildlife (Mutze 1996). However, there are pri- vate plans to increase livestock grazing (the density of sheep and cattle) in these areas immediately af- ter the rabbits are eradicated, perhaps negating some of the benefits for wildlife. Agricultural au- thorities should consider stocking restrictions to balance these pressures. Black-breasted Buzzards, 70 Mooney Vol. 32, No. 1 Black Falcons, Grey Falcons, and Letter-winged Kites are the most susceptible to grazing pressures. Collecting. Egg collecting is a small but persis- tent problem targeting uncommon species. Once nests of species such as Red Goshawks or Grey Fal- cons become well known they may be robbed (Cupper and Cupper 1981, Hollands 1984, Garnett 1992, Raker-Gabb 1993, Marchant and Higgins 1993) and it is the responsibility of researchers and photographers to restrict this knowledge. At worst, it is a local problem for a few species. Recreational falconry is not allowed in Australia and there is little consequent nest robbing and no pet trade. Persecution. Australia is a member of CITES and all raptors in Australia have full legal protection under state and territory laws (Aumann et al. 1989). However, wildlife authorities retain the right to issue special permits to control raptors. Perse- cution of raptors in the Australian tropics is only a local problem. Beyond random vandalism the big- gest problems are associated with the defense of stock. Accipiters are commonly shot near poultry and aviary birds; the Brown Goshawk was heavily persecuted on Christmas Island (Van Tets 1967) but this now has diminished. Wedge-tailed Eagles are still reported to cause losses of lambs and even calves (Keough 1994), and are frequently shot, trapped, or poisoned. Such shooting is still com- mon in western Queensland and several stockmen shoot eagles from helicopters (Czechura, pers. comm.). Persecution maybe an important contrib- uting factor to the range contraction of the Red Goshawk from northern New South Wales and most of southeastern Queensland (Debus and Cze- chura 1988a, Baker-Gabb 1993). Besides policing, education (especially use of peer pressure) is one of the few ways to limit such persecution. Present efforts (Holdsworth and Marmion 1993) should be consolidated and expanded. Australia’s new, strict, national gun laws can only help reduce persecu- tion of raptors. Pesticides. Pesticides can affect both productivity and survival of raptors. The effects of some persis- tent organochlorine pesticides such as DDT and dieldrin are reasonably well known (Riseborough 1987) but today they are probably of little conse- quence for populations of Australian raptors (Ful- ler and Maples 1993). The increasing problem of malaria in the nearby Solomon Islands brings greatly increased use of DDT and if climates change, the possibility of malaria establishing in Australia exists. The effect of most pesticides, especially when used as cocktails, is little known and perhaps pre- sents new dangers for Australian raptors. This es- pecially applies to cotton crops on which large amounts of pesticides are traditionally used. Some pesticides (rodenticides and insecticides) are highly toxic and raptors have been killed by them in Australia. Experiences in Israel have shown how catastrophic the use of topical, organo- phosphate rodenticides or insecticides can be for raptors (Mendelssohn and Puz 1977). A recent ac- cidental kill of up to 20 000 Swainson’s Hawks (Bu- teo swainsoni ) by the organophosphate insecticide monocrotophos on their Argentine wintering grounds (Krapovickas and Lyons 1997) shows the potential of organophosphate insecticides to seri- ously affect populations of Nankeen Kestrels, Brown Falcons, Little Eagles, Masked Owls ( T. no- vaehalla ndiae) , Barn Owls, Grass Owls, Whistling Kites, Black Kites, Black-shouldered Kites, and Let- ter-winged Kites in Australia. The use of pesticides should be under severe restrictions in areas where these species are likely to winter or concentrate to breed. Even in established agricultural areas, large concentrations of wintering raptors can be found (e.g., Nankeen Kestrels on the Darling Downs), benefiting from limited agriculture, but the use of highly-toxic chemicals must be controlled in these places. Agricultural Codes of Practice along the lines of Forestry Codes of Practice could have ob- vious use. Dingo/feral dog (Canis familiaris ) and feral pig ( Sus scrofa ) poisoning programs use sodium mon- oflouroacetate (1080), a chemical to which Austra- lian raptors are somewhat resistant (Mcllroy 1984, 1986). Nevertheless, illegal baiting using organo- phosphates (lucijet, a fenthion-ethyl derivative, and phosdrin) can and does kill raptors (DuGuesclin et al. 1983). Exotic Species. Besides the risks of nontarget poisoning, introduced species present many prob- lems to raptors. The cane toad (Bufo marinaris ) is highly toxic and it may have killed raptors after its release. Today, it is a common prey item of the Black Kite (Marchant and Higgins 1993). Foxes ( Vulpes vulpes) and feral cats (Felis domesticus ) can compete with and prey on raptors and both are found in the dry and wet tropics. Their potential effect is greatest for ground nesters such as the Grass Owl and small, colonial nesters such as the Letter-winged Kite (Cupper and Cupper 1981). Such depredation may increase because of prey March 1998 Status of Tropical Australian Raptors 71 switching when rabbit calicivirus disease decimates rabbits. Beyond plantation forestry or agriculture, intro- duced plants can change large areas. Afro-Asian prickly acacia ( Acacia nilotica) has changed nearly 50 000 km 2 of grassland in Queensland to tall scrub (Anonymous 1990). Much of this was potentially valuable wintering habitat for raptors. Under the federal Endangered Species Protec- tion Act 1992, research and management phases of Recovery Programs for species threatened nation- ally are funded by the Australian Nature Conser- vation Agency. Funding to carry out follow-up re- commnedations is not guaranteed. Fortunately, there is no remedial action urgently needed for any threatened species except perhaps for the Red Goshawk in New South Wales and southeast Queensland (Auman and Baker-Gabb 1991b). Reserved Status. A common conservation mea- sure for wildlife is to protect land through the es- tablishment of reserves. For this to be effective, the most important areas of habitat must first be iden- tified. Some rare species, such as the Square-tailed Kite, have an enormous distribution, are migratory through dispersive rather than routed mecha- nisms, and seem always at low population densities making accurate assessment of their status and identification of crucial wintering areas very diffi- cult. The large home ranges of many breeding rap- tors necessitates enormous reserves to support vi- able populations (Thiollay 1989). About 5.9% of Australia is under reservation for conservation, compared to 10.5% for the U.S., 7% for Canada and a world average of 4.9% (McLennan 1996). Although Australia’s proportion seems small, pro- tected areas in the north are well connected by relatively intact habitat. Most of the islands, Christ- mas Island included, are now well reserved. Al- though all Australian raptors are found in at least one reserve, Australia has no large reserves dedi- cated to raptors similar to those found in Israel, Spain, and the U.S. This is due to the fact that Australia’s reserves are generally for the conserva- tion of mammals, a group in far more need of spe- cial attention than raptors (Flannery 1994), Conclusion The vast majority of tropical Australia is still sparsely populated by people and environmental degredation is at a low level. As a result, problems for raptors in Australia’s tropics are generally lo- calized but development has resulted in significant contraction of range for at least one species, the Red Goshawk. Fortunately, no raptor in Australia is in need of intensive management. Beyond mak- ing deliberate efforts to preserve a few subspecific island populations, further researching insuffi- ciently known and rare species, and confirming the security of most species through regular monitor- ing of populations and threats, aggressive educa- tion and development of Codes of Practice are probably all Australia’s raptors need for security well into the 21st century. Ackn o wledgments I would like to thank Henry Nix, Richard Hill, Penny Olsen, and Stephen Debus for their thoughts on raptors in Australia’s tropics. Will Steele, Dave Baker-Gabb, and Richard Hill kindly let me use their unpublished data Dave Baker-Gabb and Jean-Marc Thiollay kindly reviewed the text and the Raptor Research Foundation helped me attend the conference in Boise where this paper was pre- sented. Literature Cited Anonymous. 1988. Living with hawks. Wildlife Note- sheet, TASPAWS, Hobart, Australia. . 1990. Atlas of Australian resources: vegetation. AUSLIG, Canberra, Australia. . 1993. Forest practices code. Forestry Tasmania, Hobart, Australia. . 1994. Wetland Restoration. In D. Navid [Ed,], Ramsar Newsletter 17. . 1996. Australia: State of the environment 1996, biodiversity. ANCA, Canberra, Australia. Aumann, T. and D.J. Baker-Gabb. 1991a. The ecology and status of the Red Goshawk in Northern Australia. RAOU Report 75. RAOU, Melbourne, Australia. and . 1991b. A management plan for the Red Goshawk. RAOU, Melbourne, Australia. , N.J. Mooney and P.D. Olsen. 1989. The legal status of birds of prey in Australia. Pages 591-596 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptors in the modern world. World Working Group for Birds of Prey, London, U.K. Baker-Gabb, D.J. and K. Fitzherbert. 1989. An overview of raptor movement and wintering places in Australia and New Zealand. Pages 159-166 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptors in the modern world. World Working Group for Birds of Prey, Lon- don, U.K. . 1994. Threatened raptors of Australia’s tropical forests. Pages 241-254 in B.-U. Meyburg and RD Chancellor [Eds.], Raptor conservation today. World Working Group for Birds of Prey, London, U.K. . 1996. When rabbit calicivirus strikes, whither the raptors? Wingspan 6. RAOU, Melbourne, Australia. Blakers, N., S.J.J.F. Davies and P.N. Reilley. 1984. Atlas 72 Mooney Vol. 32, No. 1 of Australian birds. Melbourne Univ. Press, Mel- bourne, Australia. Brereton, R.N. and N. Mooney. 1994. Conservation of the nesting habitat of the Grey Goshawk Accipiter no- vaehollandiae in Tasmanian state forests. Tasforests 6: 79-91. Coman, B.J. 1996. Environmental impacts associated with the proposed use of rabbit calicivirus disease in Australia, CSIRO, Canberra, Australia. Cupper, J. and L. Cupper. 1981. Hawks in focus. Jaclin Enterprises, Mildura, Australia. Clark, R.J. AND H. Mikkola. 1989. A peliminary revision of threatened and near threatened nocturnal birds of prey in the world. Pages 371-388 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptors in the modern world. World Working Group for Birds of Prey, Lon- don, U.K. Davis, T.J. 1994. The Ramsar convention manual. Ram- sar Convention Bureau, Gland, Switzerland. Debus, S.J.S. 1993. The status of the Red Goshawk Ery- throtriorchis radiatus in New South Wales. Pages 182- 191 in P. Olsen [Ed.], Australian raptor studies. RAOU, Melbourne, Australia. . 1994a. What is the Christmas Island goshawk? Aust. Bird Watcher 15:377-379. . 1994b. The Sooty Owl Tyto tenebricosa in New South Wales. In P. Roberts [Ed.], Large forest owls of New South Wales. Australian Birds 28 (Supplement). and G.V. Czechura, 1988a. The Red Goshawk Erythrotriorchis radiatus: a review. Aust. Bird Watcher 12: 175-199. and . 1988b. The Square-tailed Kite Lo- phoictinia isura: a review. Aust. Bird Watcher 13:81-98. Draffon, R.D.W., S.T. Garnett and G. Malone. 1983. Birds of Torres Strait. Emu 83:207-234. DuGuesclin, P.B., W.B. Emison and I.B. Temby 1983. Deliberate misuse of the organophosphate pesticide, fenthion-ethyl, to poison birds in Victoria. Corella 7: 37-39. EwiNS, P.J. 1996. The use of artificial nests by an increas- ing population of Ospreys in the Canadian Great Lake Basin. Pages 109-125 in D. Bird, D. Varland and J. Negro [Eds.], Raptors in human landscapes. Aca- demic Press, London U.K. Flannery, T. 1994. The future eaters. Reed Books, Lon- don, U.K. Fuller, P. And T.G. Maples. 1993. Pesticide-related egg- shell thinning in Australian raptors. Emu 93:1-11. Gamauf, A, 1994. The influence of tourism on Marsh Harriers Circus aeruginosis in the Neusiedlersee-See- winkel National Park, Austria. Pages 103-108 in B.-U. Meyburg and R.D. Chancellor [Eds.], Raptor conser- vation today. World Working Group for Birds of Prey, London, U.K. Garnett, S. [Ed.]. 1992. Threatened and extinct birds of Australia. RAOU, Melbourne, Australia. Hill, F.A.R. 1996. The Christmas Island hawk-owl: its distribution, population size and conservation status. Unpubl. Rep., Australian Nature Conservation Agen- cy, Darwin, Australia. Holdsworth, M. and P. Marmion. 1993. Raptors and education in Tasmania. Pages 220-226 in P. Olsen [Ed.], Australian raptor studies. RAOU, Melbourne, Australia. Hollands, D. 1977. The Letter-winged Kite irruption of 1976-77. Aust. Bird Watcher 8:1-18. . 1984. Eagles, hawks and falcons of Australia. Nelson, Melbourne, Australia. . 1991. Birds of the night. Reed Books, London, U.K. IUCN. 1994. IUCN relist categories. IUCN Species Sur- vival Commission, Gland, Switzerland. Kavanaugh, R.P. and P. Peake. 1993. Distribution and habits of nocturnal forest birds in south-eastern New South Wales. Pages 101-125 in P. Olsen [Ed.], Austra- lian raptor studies. RAOU, Melbourne, Australia. Keough, H. 1994. Easy-care herefords suit prairie con- ditions. Aust. Hereford Quarterly Feb. Krapovickas, S. And J.A. Lyons. 1997. Swainson’s Hawk in Argentina: international crisis and cooperation. World Birdwatch 19:12-15. Lavery, H.J. and P.H. Johnson. 1993. The Black Kite Milvus migrans in the Townsville District of Queens- land: a comparison or irruption and nonirruption years. Pages 209-219 in P. Olsen [Ed.], Australian rap- tor studies. RAOU, Melbourne, Australia. Marchant, S. and P.J. Higgins [Eds.]. 1993. Handbook of Australian, New Zealand and Antarctic birds. Vol 2: Raptors to lapwings. RAOU, Melbourne, Australia. McLennan, W. 1996. The Ramsar convention on wet- lands: its history and development. Ramsar Conven- tion Bureau, Gland, Switzerland. Mendelssohn, H. and U. Puz. 1977. Mass mortality of birds of prey caused by azodrin, an organophospho- rus insecticide. Biol. Conserv. 11:163-170. McIlroy, J.C. 1984. The sensitivity of Australian animals to 1080 poison. VII. Native and introduced birds. Aust. Wildl. Res. 11:373-385. . 1986. The sensitivity of Australian animals to 1080 poison. IX. Comparisons between the major groups of animals, and the potential danger nontar- get species face from 1080-poisoning campaigns. Aust. Wildl. Res. 13:39-48. Mooney, N. 1996. Establishing priorities for searching forestry coupes for nests of Wedge-tailed Eagles. ARA News 17 ( 1 ) :9— 1 1 . and M. Holdsworth. 1991. The effects of dis- turbance on nesting Wedge-tailed Eagles Aquila audax fleayi in Tasmania. Tasforests 3:15-33. and R. Taylor. 1 996. Value of nest-site protec- tion in ameliorating the effects of forestry operations on Wedge-tailed Eagles in Tasmania. Pages 275-282 in D. Bird, D. Varland and J. Negro [Eds.], Raptors in human landscapes. Academic Press, London U.K. March 1998 Status of Tropical Australian Raptors 73 Mutze, G. 1996. Flinders ranges: rabbit-free and thriv- ing. Rabbit Calicivirus Update, May. Newton, I. 1979. Population ecology of raptors. Buteo Books, Vermillion, SD U.S.A, Nix, H.A. 1972. Environmental control of breeding, post breeding dispersal and migration of birds in the Aus- tralian region. Proc. Int. Ornithol. Congr. 16:272-305. Riseborough, R.W. 1987. Toxic chemicals and birds of prey: discussion at Eilat in 1987. In B.-U. Meyburg and R.D. Chancellor [Eds.], Raptors in the modern world. World Working Group for Birds of Prey, Berlin, Ger- many. Robinson, D. and B.J. Traill. 1996. Conserving wood- land birds in the wheat and sheep belts of southern Australia. RAOU Conservation Statement 10. SCHODDE, R. AND IJ- Mason. 1980. Nocturnal birds of Australia. Landsowne, Melbourne, Australia. Temby, I. 1992. A guide to living with wildlife. Dept. Con- serv. and Environ., Victoria Government Publ. THIOLLAY, J.M. 1989. Area requirments for the conser- vation of rainforest raptors and game birds in French Guinea. Conserv. Biol. 3:128-137. . 1994. A world review of tropical forest raptors Current trends, research objectives and conservation strategy. Pages 231-232 in B.-U. Meyburg and R.D. Chancellor [EDS.], Raptor conservation today. World Working Group for Birds of Prey, London, U.K. Van Tets, G.F. 1967. A report on the resident birds of the territory of Christmas Island. Emu 66:309-319 Williams, K., I. Parer, B. Coman, J. Burley and M Braysher. 1995. Managing vertebrate pests: rabbits. Bureau of Resource Sciences/CSIRO, Aust. Govt. Printers, Canberra, Australia. Wood, D. 1996. Tassie’s five-acre nightmare. The Sun- day Tasmanian, 4/7/96: 6-7, Hobart, Australia. Received 16 October 1996; accepted 15 October 1997 BOOK REVIEW Edited by Jeffrey S. Marks Buteo Books is pleased to sponsor the Book Review section of the Journal of Raptor Research. Buteo Books stocks a comprehensive selection of ornithology books, both new and used. J. Raptor Res. 32(1): 74-75 © 1998 The Raptor Research Foundation, Inc. The Prairie Falcon. By Stanley H. Anderson and John R. Squires. 1997. University of Texas Press, Austin, TX. viii + 162 pp., 19 color photos, 10 fig- ures, 5 tables. ISBN 0-292-70473-9. Cloth, $29.95.— When I discovered that the cover of this book bears a photograph of my hunting companion of 10 years, I knew we were off to a good start. Like many people, I had been waiting for someone to amass under one cover much of the existing but disconnected information about the only falcon in- digenous solely to North America. Finally, Ander- son and Squires have shouldered the burden, cre- ating a useful addition to the literature that focuses on the subject and broaches many of the topics one associates with the field of raptor biology. Be- cause the Prairie Falcon ( Falco mexicanus) is neither a glamour species nor ecologically endangered, this compilation has been a long time coming. Stanley H. Anderson, Professor of Zoology and leader of the Wyoming Cooperative Wildlife Re- search Unit, and John R. Squires, Research Scien- tist for the U.S. Forest Service, bring strong aca- demic credentials and a fresh approach to under- standing this under-explored raptor. Those of us who have searched for in-depth information on the Prairie Falcon and struggled through the works of Bent, Gabrielson and Jewett, Brown and Ama- don, and other dated sources, owe the authors a debt of gratitude. For those who seek information on the Prairie Falcon, this is the new starting point. In format, the book is a hybrid between popular and scientific literature, one that may expose the reader to new topics without burying him or her in technical terminology. It is arranged in accessi- ble, well-defined sections. As one might expect, The Prairie Falcon offers much natural history informa- tion and provides scientific documentation. I par- ticularly liked the brief inserts that highlight vari- ous topics. Some general, some specific, the inserts focus on many of the curious aspects of raptor bi- ology. For compulsive readers, sources of infor- mation are conveniently listed by topic at the end of each chapter. Starting with general life-history information, the book explores the Prairie Falcon's behavior, hunting and food habits, ecological and interspe- cific interactions, and its predatory nature. In ad- dition to familiar topics like breeding distribution, territoriality, and reversed sexual size dimorphism, the unusual topic of tree nesting by Prairie Falcons is discussed. Under Coloration (p. 5), the authors refer to several color morphs of the Prairie Falcon, which may be misleading. Prairie Falcon plumages vary between light and dark, but there are no distinct color morphs as found in Gyrfalcons (F. rusticolus ) and some other raptors. In the next section (p. 15), the authors discuss nest-site fidelity, a strong characteristic in this species. Although this aspect of behavior is well documented, there is no evi- dence that Prairie Falcons mate with the same part- ner from year to year, a point that should have been included. I particularly enjoyed the section on Hunting Techniques (p. 28) , which includes information on foraging behavior, kleptoparasitism, and food cach- ing. However, I should note a slight discrepancy in the interpretation of data from my study of forag- ing behavior. Although most of the attacks I ob- served were of the low coursing type at moderate speeds, some attacks were initiated from a soar and ended in an extremely fast, low-trajectory ap- proach close to the ground. The phenomenon is described by the aeronautical term “ground ef- fect.” Clearly, these two methods were not the same. The soaring technique, when applied to prey spotted on the ground, gave the falcon great in- coming speed and made it difficult for prey to de- tect the approaching falcon. I suspect that all large falcons use this soaring technique to some degree. 74 March 1998 Book Review 75 Table 2 describes the food habits of Prairie Falcons collected throughout the western United States. It clearly shows that these falcons are adaptable pred- ators, able to glean whatever food resources the local environment has to offer. Chapter 2 covers seasonal patterns of courtship, breeding, and nesting. Information on fledging ac- tivities and fall and winter dispersal patterns also are included. I wish Figure 6 (p. 82) had included a map of current band-recovery information for all age groups, as well as data from breeding popula- tions in the southern part of the falcon’s range. Data for this figure were collected in 1964. How- ever, most of the information on the species has been collected since the 1970s. Jim Enderson was the first researcher to recognize the importance of the Great Plains to the wintering ecology of this species, including falcons from as far west as Cali- fornia and Oregon. Considering that most Prairie Falcons nest west of the Continental Divide, and that human-caused alterations in the Plains States have vastly changed the ecology of that region, there is still much to be learned by investigating the eastern movements during fall and winter. The authors’ comments on the fragility of the Prairie Falcon’s environment lead one to conclude that winter habitat may, in fact, be one of the most crit- ical elements for the long-term survival of the spe- cies. Early on, the authors mention that nestlings fledge at two months of age. But on p. 73, they correctly state that fledging age is between 33 and 35 days of age. It seemed curious that no mention was made of the adult male’s role in incubation (p. 63) . Although undoubtedly there is wide variation among pairs, it should be noted that males share this duty during daylight hours. The third chapter concentrates on the juxtapo- sition of the Prairie Falcon in relation to other rap- tors found in North America and around the world. It serves primarily as a review of nomencla- ture, physiology, and evolution. The Taxonomy sec- tion (p. 100) seems to support the conclusion that Tom Cade reached in The Falcons of the World : that Prairie Falcons are part of the “desert falcon” group that includes Gyrfalcons, Saker Falcons (F cherrug), and Lanner Falcons (F. biarmicus ). How- ever, at the very end of the book (p. 138), the au- thors acknowledge that recent DNA studies show that Prairie Falcons are more closely related to Per- egrine Falcons (F. peregrinus ) . Chapter 4 begins with an overview of the Prairie Falcon’s role in falconry and captive breeding. The authors state that stocks of Prairie Falcons can be managed to provide both viable populations of wild birds and a falconry harvest. Under Captive Breeding (p. 119), a method of semen collection is described that would be appropriate for large poultry, but not for raptors. (It should be noted that Prairie Falcons breed readily in captivity. Yet, since the initial flurry of captive-breeding technol- ogy two decades ago, little effort has been expend- ed on breeding them in any number.) The authors then present current information on the effects of pesticides, human influences, and mitigation mea- sures. The book concludes by suggesting that the spe- cies be viewed holistically — as one important link in the food chain of its habitat — linking predator with prey, prey with habitat, and habitat with hu- manity. We must make this connection if Prairie Falcons, and indeed all raptors, are to have a fu- ture on this continent. Finally, the Raptor Etiquette section in the Ap- pendix offers sound suggestions on how to watch nesting falcons without disturbing them. Overall, I appreciated the high quality of the book, including good paper and a sturdy cover. Simple line draw- ings, augmented by striking photographs by LuRay Parker and Rick Kline, enhance its visual appeal. Although the angled layout of the photographs may be artistically appealing to some, I found them mildly distracting. The book comes complete with a well-stocked section of references. There were some minor typos, but these are hard to avoid. The Prairie Falcon belongs in the libraries of serious scholars, budding researchers, fans of western wild- life, and avid birders of all descriptions. At $29.95, it is a bargain. — Bruce Haak, Idaho Department of Fish and Game, 3101 South Powerline Road, Nam- pa, ID 83686 U.S.A. J Raptor Res. 32(1) :76 © 1998 The Raptor Research Foundation, Inc. Manuscript Referees The following people reviewed manuscripts for the Journal of Raptor Research in 1997. Peer review plays a vital role in the publishing process and in improving the quality of the Journal. The names of individuals who reviewed two or more manuscripts are followed with an asterisk. Beatriz Arroyo*, Eric C. Atkinson, James C. Bednarz, Maria Isabel Bellocq*, James R. Belthoff, Keith L. Bildstein*, David M. Bird, Clint W. Boal*, Gary R. Bortolotti*, Thomas Bosakowski, Joseph B. Buchanan, Javier Bustamente*, Tom J. Cade*, William S. Clark, Randy Craft, Dick Dekker, Stephen DeStefano, Alfred M. Dufty, Jr., Gary E. Duke*, James H, Enderson*, Allen M. Fish*, Eric Forsman*, Alan B. Franklin, Frederick R. Gehlbach*, Richard P. Gerhardt, James A. Gessaman, Richard Glinski, Daniel N. Gossett, Teryll Grubb, Ralph J. Gutierrez*, Alan R. Harmata*, Eliza- beth Haug, Gregory D. Hayward, William Heinrich, Charles J. Henny*, Geoffrey L. Holroyd, Denver W. Holt*, C. Stuart Houston*, C. Hubert, Jerome A. Jackson, Fabian M, Jaksic*, Paul James*, Robert S. Kennedy, Steven Knick, Jeffrey S. Marks*, Carl D. Marti*, John M. Marzluff, Carol L. McIntyre, Michael McGrady*, Heimo Mikkola*, Brian A. Milsap, Helmut C. Mueller*, Ian Newton*, Alan Poole*, Charles R. Preston*, Patrick T. Redig*, Gary Ritchison, James A. Rodgers, Jr,, Robert N. Rosenfield, Sievert A. Rohwer, Philip F. Schempf, Josef K. Schmutz*, Dwight G. Smith*, Karen Steenhof*, Stephen Sweeney*, Theodor Swem, Paul W. Sykes, Jr., Ethan J. Temeles, Jean-Marc Thiollay, Kimberly Titus*, Helen M. Ulmschneider, S. Van Balen, Daniel E. Varland, Ian G. Warkentin*, Richard T. Watson, David Whitacre, Clayton M. White, Per Widen. [ BUTEO BOOKS The following Birds of North America Species Accounts are available through Buteo Books, 3130 Laurel Road, Shipman, VA 22971. TOLL-FREE ORDERING: 1-800-722-2460; FAX: (804) 263-4842. Barn Owl (1). Carl D. Marti. 1992. 16 pp. Boreal Owl (63). G.D. Hayward and P.H. Hayward. 1993. 20 pp. Broad-winged Hawk. (218). L.J. Goodrich, S.C. Crocoll and S.E. Senner. 1996. 28 pp. Burrowing Owl (61). E.A. Haug, B.A. Millsap and M.S. Martell. 1993. 20 pp. Common Black-hawk (122). Jay H. Schnell. 1994. 20 pp. Cooper’s Hawk (75). R.N. Rosenfield andj. Bielefeldt. 1993. 24 pp. Crested Caracara (249). Joan L. Morrison, 1996. 28 pp. Eastern Screech-owl (165). Frederick R. Gehlbach. 1995. 24 pp. Ferruginous Hawk (172). MarcJ. Bechard and Josef K. Schmutz. 1995. 20 pp. Flammulated Owl (93). D. Archibald McCallum. 1994. 24 pp. Great Gray Owl (41). Evelyn L. Bull and James R. Duncan. 1993. 16 pp. Gyrfalcon (114). Nancy J. Clum and Tom J. Cade. 1994. 28 pp. Harris’ Hawk (146). James C. Bednarz. 1995. 24 pp. Long-eared Owl (133). J.S. Marks, D.L. Evans and D.W. Holt. 1994. 24 pp. Merlin (44). N.S. Sodhi, L. Oliphant, P. James and I. Warkentin. 1993. 20 pp. Northern Saw-whet Owl (42). Richard J. Cannings. 1993. 20 pp. Northern Goshawk (298). John R. Squires and Richard T. Reynolds. 1997. 32 pp. Northern Harrier (210). R. Bruce MacWhirter and Keith L. Bildstein. 1996. 32 pp. Red-shouldered Hawk (107). Scott T. Crocoll. 1994. 20 pp. Red-tailed Hawk (52). C.R. Preston and R.D. Beane. 1993. 24 pp. Short-eared Owl (62). D.W. Holt and S.M. Leasure. 1993. 24 pp. Snail Kite (171). P.W. Sykes, Jr., J.A. Rodgers, Jr. and R.E. Bennetts. 1995. 32 pp. Snowy Owl (10). David F. Parmelee. 1992. 20 pp. Spotted Owl (179). R.J. Gutierrez, A.B. Franklin and W.S. Lahaye. 1995. 28 pp. Swainson’s Hawk (265). A. Sidney England, MarcJ. Bechard and C. Stuart Houston. 1997. 28 pp. Swallow-tailed Kite (138). Kenneth D. Meyer. 1995. 24 pp. White-tailed Hawk (30). C. Craig Farquhar. 1992. 20 pp. White-tailed Kite (178). Jeffrey R. Dunk. 1995. 16 pp. 76 THE RAPTOR RESEARCH FOUNDATION, INC. (Founded 1966 ) OFFICERS PRESIDENT: Michael N. Kochert SECRETARY: Patricia A. H ai.i. VICE-PRESIDENT: David E. Andersen TREASURER: Jim Fitzpatrick BOARD OF DIRECTORS NORTH AMERICAN DIRECTOR #1: Brian A. Millsap NORTH AMERICAN DIRECTOR #2: Petra Bohall Wood NORTH AMERICAN DIRECTOR #3: Karen Steenhof INTERNATIONAL DIRECTOR #1: Massimo Pandolfi INTERNATIONAL DIRECTOR #2: Reuven Yosef EDITORIAL STAFF EDITOR: MarcJ. Bechard, Department of Biology, Boise State University, Boise, ID 83725 U.SA. ASSOCIATE EDITORS Allen M. Fish Fabian Jaksic Gary R. Bortolotti Daniel E. Varland Charles J. Henny BOOK REVIEW EDITOR: Jeffrey S. Marks, Montana Cooperative Research Unit, University of Montana, Missoula, MT 59812 U.SA. SPECIAL PUBLICATIONS EDITOR: Daniel E. Varland, Rayonier, 3033 Ingram Street, Hoquiam, WA 98550 SPANISH EDITOR: Cesar MArquez Reyes, Instituto Humbolist Colombia, AA. 094766, Bogota 8, Colombia The Journal of Raptor Research is distributed quarterly to all current members. Original manuscripts dealing with the biology and conservation of diurnal and nocturnal birds of prey are welcomed from throughout the world, but must be written in English. Submissions can be in the form of research articles, letters to the editor, thesis abstracts and book reviews. Contributors should submit a typewritten original and three copies to the Editor. All submissions must be typewritten and double-spaced on one side of 216 X 278 mm (8A£ X 11 in.) or standard international, white, bond paper, with 25 mm (1 in.) margins. The cover page should contain a title, the author’s full name(s) and address(es). Name and address should be centered on the cover page. If the current address is different, indicate this via a footnote. A short version of the title, not exceeding 35 characters, should be provided for a running head. An abstract of about 250 words should accompany all research articles on a separate page. Tables, one to a page, should be double-spaced throughout and be assigned consecutive Arabic numer- als. Collect all figure legends on a separate page. Each illustration should be centered on a single page and be no smaller than final size and no larger than twice final size. The name of the author (s) and figure number, assigned consecutively using Arabic numerals, should be pencilled on the back of each figure. Names for birds should follow the A.O.U. Checklist of North American Birds (6th ed., 1983) or another authoritative source for other regions. Subspecific identification should be cited only when pertinent to the material presented. Metric units should be used for all measurements. Use the 24-hour clock (e.g., 0830 H and 2030 H) and “continental” dating (e.g., 1 January 1990). Refer to a recent issue of the journal for details in format. Explicit instructions and publication policy are oudined in “Information for contributors,”/. Raptor Res., Vol. 27(4), and are available from the editor. INTERNATIONAL DIRECTOR #3: Michael McGrady DIRECTOR AT LARGE #1: Patricia L. Kennedy DIRECTOR AT LARGE #2: John A. Smallwood DIRECTOR AT LARGE #3: James C. Bednarz DIRECTOR AT LARGE #4: Cesar MArquez Reyes DIRECTOR AT LARGE #5: Lloyd Kiff DIRECTOR AT LARGE #6: Robert Kenward 1998 ANNUAL MEETING The Raptor Research Foundation, Inc. will hold its 1998 annual meeting in Ogden, Utah from 30 September through 4 October at the Ogden Egyptian Conference Center. Details about the meeting and a call for papers will be mailed to members in the spring of 1998. A site on the World Wide Web contains information about the meeting and setting (http://www.weber.edu/ rrf). Contact Carl D. Marti for more information (801/626-6172; cmarti@weber.edu). Raptor Research Foundation, Inc., Awards Recognition for Significant Contributions 1 The Dean Amadon Award recognizes an individual who has made significant contributions in the field of systematics or distribution of raptors. Contact: Dr. Clayton White, 161 WIDB, Department of Zoology, Brigham Young University, Provo, UT 84602 U.S.A. Deadline August 15. The Tom Cade Award recognizes an individual who has made significant advances in the area of captive propagation and reintroduction of raptors. Contact: Dr. Brian Walton, Predatory Bird Research Group, Lower Quarry, University of California, Santa Cruz, CA 95064 U.SA. Deadline: August 15. The Fran and Frederick Hamerstrom Award recognizes an individual who has contributed significantly to the understanding of raptor ecology and natural history. Contact: Dr. David E. Andersen, Department of Fisheries and Wildlife, 200 Hodson Hall, 1980 Folwell Avenue, University of Minnesota, St. Paul, MN 55108 U.SA. Deadline: August 15. Recognition and Travel Assistance The James R. Koplin Travel Award is given to a student who is the senior author of the paper to be presented at the meeting for which travel funds are requested. Contact: Dr. Petra Wood, West Virginia Cooperative Fish and Wildlife Research Unit, P.O. Box 6125, Percival Hall, Room 333, Morgantown, WV 26506-6125 U.SA Deadline: established for conference paper abstracts. The William C. Andersen Memorial Award is given to the student who presents the best paper at the annual Raptor Research Foundation Meeting. Contact: Ms. Laurie Goodrich, Hawk Mountain Sanctuary, Rural Route 2, Box 191, Kempton, PA 19529-9449 U.SA. Deadline: Deadline established for meeting paper abstracts. Grants 2 The Stephen R. Tully Memorial Grant for $500 is given to support research, management and conservation of raptors, especially to students and amateurs with limited access to alternative funding. Contact: Dr. Kimberly Titus, Alaska Division of Wildlife Conservation, P.O. Box 20, Douglas, AK 99824 U.SA. Dead- line: September 10. The Leslie Brown Memorial Grant for $500-$l,000 is given to support research and/or the dissemination of information on raptors, especially to individuals carrying out work in Africa. Contact: Dr. Jeffrey L. Lincer, 15644 Kingman Rd., Poway, CA 92064 U.SA. Deadline: September 15. 1 Nominations should include: (1) the name, title and address of both nominee and nominator, (2) the names of three persons qualified to evaluate the nominee’s scientific contribution, (3) a brief (one page) summary of the scientific contribution of the nominee. 2 Send 5 copies of a proposal (^5 pages) describing the applicant’s background, study goals and methods, anticipated budget, and other funding.