Breeding Biology and Nestling Diet of the Great Black&Hyphen;Hawk

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SHORT COMMUNICATIONS j. RaptorRes. 32(2):175-177 ¸ 1998 The Raptor Research Foundation, Inc.

BREEDING BIOLOGY AND NESTLING DIET OF THE GREAT BLACK-HAWK

NATHANIEL E. SEAVY 17142 LemoloShr. Dr. N.E., Poulsbo, WA 98370 U.S.A.

RtCH?d•D P. GERHARDT

341 N.E. Chestnut St., Madras, OR 97741 U.S.A.

KEY WORDS: GreatBlack-Hawk; Buteogallus urubitinga; Observationsof courtship behavior or of hawkscarry- breedingbiology; diet;, Petgn; Guatemala. ing nest material or prey led to the eventual location of nests.After they were found, nests were checked every 2-3 d to record nesting phenology.During all yearswe The Great Black-Hawk (Buteogallusurubitinga) rang- recorded nest size (diameter and depth) and situation, es from Mexico south to eastern Bolivia, Paraguayand and described nest trees. Observationsof prey deliveries to nests were made with binoculars from observation northern Argentina, inhabiting coastal lowlands and foothills (Brown and Amadon 1968). The few accounts platforms constructed in trees about 35 m from nests. We climbed to nestsweekly to weigh and measure nest- describing its breeding biology have been brief and at lings in 1991; in 1993 and 1994 we avoided climbing to times contradictory(Grossman and Hamlet 1964, Smithe nests,except to verify someclutch sizes,until after fledg- 1966, Brown and Amadon 1968, ffrench 1976, Mader ing. Additional information on clutch size, nesting phe- 1981). Based primarily on isolated observationsof hunt- nology and nestswas obtained from egg-setdata records ing and prey remains collected beneath roosts, a wide from published accounts,the Western Foundation of Ver- variety of prey items has been recorded, including inver- tebrate Zoology (WFVZ), and the Delaware Museum of tebrates,fish, frogs, reptiles,birds, mammalsand carrion Natural History (DMNH). (Dickey and van Rossem 1938, Lowery and Dalquest 1951, Haverschmidt 1962, Wetmore 1965, Mader 1981, RESULTS AND DISCUSSION Olmos 1990, Lewis and Timm 1991). The earliest we observed nesting activity of Great We studied Great Black-Hawks in Tikal National Park, Black-Hawkswas 23 March 1994 when a pair began to Pet6n, Guatemalaas part of The PeregrineFund's "Maya copulate and build a nest. Egg-layingdates were known Project." Two nests were located and studied during for two nests,16-17 April 1991 and 4 May 1994. An •n- 1991, and information collected on food habitswas pub- cubation period of 40 +_2 d was recorded for one nest lished by Gerhardt et al. (1993). We continued to study Basedon this interval, we estimatedan additional laying the speciesduring 1993-94. Here, we present informa- date as 25 March 1994. Known hatch dateswere 27 May tion on its breeding biology, including nesting phenolo- 1991 and 4 May 1994. One nestlingfledged from the nest gy, reproductive successand nest descriptions,and ad- on 28 June 1994, 55 d after hatching. Another nestling ditional dietary data. had not yet fledged at the age of 63 d when observations

STUDY AREA AND METHODS were concludedon 27 July 1991. One nest, found on 6 July 1994, contained a nestling estimatedto be 60 d old Tikal National Park covers 576 km 9 in N.E. Guatemala that had not yet fledged. We know of no other accounts (17ø13'N, 89ø38'W). Elevation averages200-250 m, to- of the duration of incubation or nestling periods for pography is gently rolling, and the climate is tropical, Great Black-Hawks in the wild. For the Common Black- with annual rainfall of about 135 cm. The rainy season begins in June or July, with the highest rainfall in Sep- Hawk (Buteogallusanthracinus), slightly shorter incuba- tember, and a pronounced dry seasonoccurs from Feb- tion (37-39 d) and nestling(43-52 d) periodshave been ruary until June or July. Vegetation, climate and land use reported (Schnell 1994). Substantial geographic varia- patterns of the Tikal area were described by Schulze tion in Great Black-Hawkbreeding phenologymay exist. (1992). At Tikal young fledged at the onset of the rainy season.

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In Surinam (Haverschmidt 1962), Trinidad (ffrench height was 56 cm (SD = 12 cm, N = 3, range = 42-65 1976), and Venezuela(Mader 1981), however,the species CIII). has been observedincubating during, or at the onset of, We found eight neststhat Great Black-Hawkswere con- the rainy season.Egg-laying may occur as early as Decem- structing or embellishing.Four did not contain eggsand ber in E1 Salvador (Brown and Amadon 1968) and as late have been excluded from analysisbecause we were un- asJune in Tamaulipas,Mexico (Martin et al. 1954). Eggs able to determine whether thesepairs nestedat alternate in DMNH and WFVZ were collected in Trinidad and Ven- sites.Of the four nestsin which eggswere laid, two failed ezuela during April and May and in Argentina from Au- during incubation, after 10 and 37 d, respectively.Young gust-January. hatched at two nests;one waskilled by an unknown pred- Our observationsof one juvenile each from two 1994 ator 6.5 wk after hatching and the other fledged. nests indicated an extended dependency period during In each of three nests at which we documented clutch which food was provided by the adults. We observed a size, only one egg was laid. Three additional nests, dis- prey deliveryto one juvenile almost8 mo after fledging, covered after hatching, contained a single nestling each. and another juvenile was consistentlyfound within 500 We documented 27 Great Black-Hawkegg setsfrom Ven- m of the nest tree up to an age of 12 mo, when obser- ezuela (7 sets), Argentina (18 sets) and Trinidad (2 vationswere discontinued.Such an extended dependen- sets)(Swann 1923, Norris 1926, WFVZ, DMNH). All cy period is supportedby Mader (1981) who reportedan clutches contained a single egg. To our knowledge the m•mature Great Black-Hawk, ca. 7 mo postfledging, only evidenceof a larger clutch size is an observationof perched next to an adult, begging for food. In contrast, a "family group" of four individualsin Mexico (Martin the Common Black-Hawk is believed to reach indepen- et al. 1954). dence about 2 mo after fledging (Schnell 1994). Post- Including the 106 prey of Gerhardt et al. (1993), stud- fledgingdependency periods of from 7 mosto more than ies of Great Black-Hawksin Tikal haveidentified 126 prey a year have been documentedfor severalother neotrop- items at least to class.Prey delivered to nestlings have •cal raptors, including Ornate Hawk-Eagles(Spizaetus or- included41 lizards(32.5% of identifiedprey), 34 snakes natus;Madrid et al. 1992), Black Hawk-Eagles(S. Tyran- (27.0%), 24 mammals (19.1%), 16 birds (12.7%), eight nus;D, Whitacre and J. Lopez pers. comm.), Harpy Ea- anurans (6.4%) and three insects (2.4%). Basilisk lizards gles (Harpia harpyja;E. Alvarezpers. comm.), CrestedEa- (Basiliscusvittatus) have made up a large percentage gles (Morphnusguianensis; D. Whitacre pers. comm.), and (70.7%) of all lizards delivered. Other lizards in the diet Gray-backed Hawks (Leucopternisoccidentalis; Vargas include arboreal genera, including Norops(formerly Ano- 1995). lis). Snakes include both arboreal and terrestrial as well All nestswe located were in "bajo" and "transitional" as venomous and nonvenomousspecies. Boa and Oxybelis torest types,which are at the lower end of the topograph- have been the snakegenera most commonlyobserved in ic gradient. Seasonallyflooded, these forest types are the diet. Mammals delivered to the nest have included characterizedby denseunderstory vegetation and a can- eight bats (6.4% of identified prey), nine rodents (7.1%) opy height that is generallylow, but broken by large, iso- and two marsupials(1.6%). Bats are medium-sizedspe- lated, emergent trees (Schulze 1992). Nests were built cies, about 30 g in mass. Although we were unable to within live, emergent trees (Swieteniamacrophylla, Budda identify bats beyond order during nest observations,Ar- buceras,Platymiscium spp., Ceibapentandra and Lonchocar- tibeusspp. remains were collected on several occasions pus castilloi),which had minimal crown contact with the from nests. Rodents have included squirrels (Sciurus surrounding forest canopy.Nest trees (N = 7) had a spp.) and unidentified cricetids.The two small marsupi- mean height of 29 m (SD = 4 m, range = 22-35 m) and als were probably mouse-opossums(Marmosa mexicanus). a mean DBH of 120 cm (SD = 81 cm, range = 66-300 Birds in the diet have ranged in size from an oriole (Ic- cm). Nestswere at a mean height of 25 m (SD = 5 m, terusspp.) to a Pale-billedWoodpecker (Campephilusgua- N = 7, range = 20-31 m) and supported by branches temalensis).Birds most commonlyobserved as prey items and/or vine tangles. are medium-sized Columbaspp. and Clay-coloredRobins Nests were constructed of dry sticks and lined with ( Turdusgrayi). Anurans and insectswere infrequentlyde- green, leafy twigs. Throughout incubation and nestling livered to nests. periods, adults added fi'esh, leafy twigs to nests.Nests The variety of prey delivered to the nestsand the wide were flat throughout the nesting period, with only one range of prey items and foraging habits reported in the having a measurable depression, 10 cm in depth. Smithe literature indicate that Great Black-Hawks are dietary (1966), Brown and Aanadon (1968), and egg records generalistsand opportunists,able to exploit a large di- from Venezuela and Trinidad (DMNH and WFVZ) de- versityof prey types and hunting situations.Midday described nests as "deeply-cupped," but our observations liveriesof nocturnal mammalianprey and reports of pre- of fiat nestsconcur with the descriptionby Grossmanand dation upon nestlings (Lewis and Timm 1991) and eggs Hamlet (1964). Considering only nests that contained (Brown and Amadon 1968, Olmos 1990) suggest that clutches, the mean nest diameter was 83 cm (SD = 21 Great Black-Hawksmay invest time in searching for vul- cm, N = 4, range = 53-100 cm), and mean external nerable and easily captured prey. Insects and anurans JUNE1998 SHORTCOMMUNICATIONS 177

may be captured more frequently than our observations GROSSMAN,M.L. ANDJ. HAMLET.1964. Birds of prey of suggest;adults may eat most such small prey upon cap- the world. Crown Publishers, New York, NY U.S.A. ture rather than transporting them to nests. Although HAVERSCHMIDT,E 1962. Notes on the feeding habits and fish and crustaceansare often mentioned as important food of some hawks in Surinam. Condor 64:154-158 prey items (Grossman and Hamlet 1964, Brown and HOWELL,S.N.G. ANDS. WEBB.1995. A guide to the birds Amadon 1968), we observed no indication that Great of Mexico and northern Central America. Oxford Black-Hawksat Tikal utilized such aquatic prey. Univ. Press, New York, NY U.S.A. Great Black-Hawksexhibit a conservativebreeding bi- LEw•s, S.E. AND R.M. TIMM. 1991. Predation on nestling ology that is characteristicof many large raptors,especial- Bare-throated Tiger-Herons by a Great Black-Hawk ly in the tropics(Newton 1979); clutchsize is smalland OrnitologiaNeotropical 2:37. young appear to have a protracted dependencyperiod. LoWERy,G.H., JR. AN• W.W. DALQUES'•.1951. Birds from Additionally, the fact that the speciesrequires several the State of Veracruz, Mexico. Univ. of Kansas Publ, years to obtain full adult plumage (Howell and Webb Mus. Nat. Hist., Lawrence, KS U.S.A. 1995, R. Gerhardt unpubl. data) suggeststhat there is a MADER,W.J. 1981. Notes on nesting raptors of the llanos substantialpopulation of nonbreeding subadults. of Venezuela. Condor 83:48-51. MADRID M., H.D., R.A. MADRID M., J.R. CRuz E., J.L. RESUMEN.-•Estudiamosla biologia reproductivade Buteo- CORDOVAA., M.C. RivERA, W.E. MARTINEZ A. AND A.R gallusurubitinga en el parque Nacional Tikal, PetEn,Gua- CAM.. 1992. Behaviorand breeding biologyof the Or- temala, durante 1991, 1993 y 1994. Un s61operiodo de nate Hawk-Eagle.Pages 179-191 in D.E Whitacre and incubaci6n fuE de 40 (-+2 dias); el pich6n emplum6 a la R.K. Thorstrom lEDS.], Maya project progressreport edad de 55 dias. Los nidos son construidoscon ramaNy V. The Peregrine Fund, Boise, ID U.S.A. palos grandesen firbolesemergentes. E1 tamafio de la MARTIN, P.S., C.R. ROBBINS AND W.B. HEED. 1954. Birds nidada de 3 nidos en Tikal, 2 en Trinidad, 7 en Vene- and biogeography of the Sierra de Tamaulipas, an iso- zuela y 18 en Argentina fue de un huevo por nido. De lated pine-oakhabitat. WilsonBull. 66:38-57. las 126 presasidentificadas en tres nidos la mayorlafue- NEWTON,I. 1979. Population ecologyof raptors. Buteo ron reptiles(59.5%), mamiferos(19.1%) y aves(12.7%). Books, Vermillion, SD U.S.A. [Traducci6n de CEsarM•rquez] Nov,RIs, J.P., JR. 1926. A catalogue of sets of Accipitres' eggs in the collection of Joseph Parker Norris, Jr, ACKNOWLEDGMENTS Philadelphia, PA U.S.A. Oolo•sts'Rec. 6:25-41. OLMOS,E 1990. Nest predation of Plumbeous Ibis by ca- This is a contribution of the "Maya Project," a conser- vation research effort of The Peregrine Fund. Financial puchin monkeysand Greater Black-Hawk. WilsonBull. support was provided by Robert Berry, Crystal Channel 102:169-170. Foundation, FanwoodFoundation, Gold Family Founda- SCHNELL,J.H. 1994. Common Black-Hawk Buteogallusan- tion, KENNETECH/U.S. Windpower, the John D. and thracinus. In A. Poole and E Gill lEDS.], The birds of Catherine T. MacArthur Foundation, Norcross Founda- North America, No. 122, Acad. Nat. Sci., Philadelphia, tion, Hank and Wendy Paulson, Pew Charitable Trusts, PA and Am. Ornithologists' Union, Washington, DC AndrEs Sada, Joe and Flinda Terteling, and the U.S. U.S.A. Agency for International Development.We are grateful SCHULZF.,M. 1992. A preliminary description of woody to the curatorial staff at the Delaware Museum of Natural plant communities of Tikal National Park. Pages53- History and the Western Foundation of Vertebrate Zo- ologyfor providing data on egg clutches.Lloyd Kiff, Carl 62 in D.E Whitacre and R.K. Thorstrom [Lbs.I, Maya Marti, Dave Whitacre, Rich Glinski, and an anonymous project progressreport V. The Peregrine Fund, Boise, reviewerprovided helpful commentson the manuscript. ID U.S.A. SMITHE, LB. 1966. The birds of Tikal. The Natural His- LITERATURE CITED tory Press,Garden City, NY U.S.A. BROWN,L. ANDD. AMADON.1968. Eagles,hawks, and fal- SWANN, H.K. 1923. Notes on the Gordon collection of cons of the world. McGraw-Hill Book Co., New York, eggs of the Accipiters. Oologists'Rec. 3:25-30. NY U.S.A. VARGAS,H. 1995. Food habits, breeding biology, and sta- DlcI•V, D.R. ANDAJ. vAN ROSSEM.1938. The birds of E1 tus of the Gray-backedHawk (Leucopternisoccidentalis) Salvador. Zool. Ser., Vol. 23. Field Mus. Nat. Hist., in western Ecuador. M.S. thesis, Boise State Univ., Box- Publ. No. 406. Chicago, IL U.S.A. se, ID U.S.A. FFRENCH.1976. A guide to the birds of Trinidad and To- WETMORE,t. 1965. The birds of the Republic of Panama, bago. Harrowood, Valley Forge, PA U.S.A. Part 1. Smiths.Misc. Coil., Vol. 150. Washington,DC GERHARDT,R.P., P.M. HAm•IS AND M.A. VAsQt:F.zM. 1993. U.S.A. Food habitsof nestingGreat Black-Hawksin Tikal Na- tional Park, Guatemala. Biotropica25:349-352. Received17 May 1997; accepted8 February 1998