Winged Hawks in Moist Karst Forests of Puerto Rico
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J RaptorRes. 39(4):404-416 ¸ 2005 The Raptor ResearchFoundation, Inc. NESTING ECOLOGY AND BEHAVIOR OF BROAD-WINGED HAWKS IN MOIST KARST FORESTS OF PUERTO RICO DEREKW. HENGSTENBERGAND FRANCISCO J. VILELLA • USGSBiological Resources Division, CooperativeResearch Units, MS 9691, Departmentof Wildlifeand Fisheries, MississippiState University, Mississippi State, MS 39762 U.S.A. ABSTRACT.-•The Puerto Rican Broad-wingedHawk (Buteoplatypterus brunnescens) is an endemic and endangered subspeciesinhabiting upland montane forestsof Puerto Rico. The reproductiveecology, behavior,and nestinghabitat of the Broad-wingedHawk were studied in Rio Abajo Forest,Puerto Rico, from 2001-02. We observed 158 courtship displaysby Broad-wingedHawks. Also, we recorded 25 ter- ritorial interactionsbetween resident Broad-wingedHawks and intruding Red-tailed Hawks (Buteoja- maicensisjamaicensis). Broad-winged Hawks displacedintruding Red-tailedHawks from occupiedterri- tories (P = 0.009). Mayfield nest survivalwas 0.67 acrossbreeding seasons (0.81 in 2001, N = 6; 0.51 in 2002, N = 4), and pairs averaged1.1 young per nest (yearscombined). The birds nestedin mixed speciestimber plantationsand mature secondaryforest. Nests were placedin the upper reachesof large trees emergingfrom the canopy.Nest tree DBH, understorystem density,and distanceto karstcliff wall correcdyclassified (77.8%) nest sites. KEYWORDS: Broad-wingHawk; Buteo platypterusbrunnescens; endangered; nest success; prey delivery; hab- itat model;karst forest; Puerto Rico. ECOLOG[AREPRODUCTIVA DE BUTEOPLATYPTERUS BRUNNESCENS EN BOSQUESDE CALIZA H•MEDOS DE PUERTO RICO RESUMEN.--Buteoplatypterus brunnescens es una subespeciede rapaz endSmicaa los bosquesmontanos de Puerto Rico. Investigamosla ecologia reproductivay el comportamiento de B. p. brunnescensen el Bosque de Rio Abajo, Puerto Rico, durante 2001 y 2002. Observamos158 desplieguesde cortejo en Rio Abajo. Observamos25 encuentrosterritoriales entre B. p. brunnescensy B. jamaicensisjamaicensis. B. p. brunnescensdesplaz6 a B. j. jamaicensisde susterritorios el 84% de las veces(P = 0.009). La supervi- venciade los nidos en ambastemporadas fue de 0.67 (0.81 en 2001, N = 6; 0.51 en 2002, N = 4). Los nidos produjeron un promedio de 1.1 volantonespor nido (aftoscombinados). Encontramos nidos en •treasde bosquesecundario maduro y plantacionesforestales. La altura del dosel, di•tmetrodel 5rbol, densidaddel sotobosquey distanciaa faral16nde mogoteclasificaron correctamente (77.8%) losnidos en Rio Abajo. [Traducci6n de los autores] The Broad-winged Hawk ( Buteoplatypterus brun- below the high canopy (Delannoy and Tossas nescens)is an endemic woodland raptor of upland 2002). This insular subspeciesis smaller and dark- montane forestsof Puerto Rico. This subspeciesis er than its North American nominate counterpart listed as endangered (Federal Register 1994) by Buteoplatypterus platypterus, but larger than the the Puerto Rico Department of Natural and Envi- LesserAntillean subspecies(Raffaele 1989, Good- ronmental Resources (DNER) and the U.S. Fish rich et al. 1996). The most recent population es- and Wildlife Service (USFWS). The Broad-winged timate for the Broad-winged Hawk in forest re- Hawk in Puerto Rico is nonmigratory and exhibits serves of Puerto Rico is approximately 125 a limited geographic range with known popula- individuals(Delannoy 1997). tions restricted to montane forests (Delannoy The nesting biology of the Broad-wingedHawk 1997). Breedingin Puerto Rico beginsin late De- in North America has been describedby a number cember,with nestsplaced in the upper reaches,but of authors (e.g., Fitch 1974, Matray 1974, Rosen- field 1984, Titus and Mosher 1987, Crocoll and Corresponding author email address: fvilella@cfr. Parker 1989). However, knowledge on the repro- msstate.edu ductive biology of the insular endemic subspecies 404 DECEMBER 2005 PUERTO RICAN BROAD-WINGED HAWK ECOLOGY 405 BWHA nest sites 2001 2002 Foresttrails /'•',•RoadRivers s Lago Dos Bocas Rio Abajo Forest 900 0 900 Meters Figure 1. Locations of BroadswingedHawk nest sims during the breeding seasonof 2001 and 2002 in Rio Abajo Forest, Puerto Rico. of Puerto RIco, and forest raptors of the West In- maicensisjamaicensis) may have on future plans to dies in general, are limited (Goodrich et al. 1996). establish a second wild population (by releasing Similarly, the available information on nesting be- captive-rearedindividuals) of the critically endan- havior and diet of Broad-wingedHawks in Puerto gered Puerto RIcan Parrot (Amazonavittata) in the RIco is based on a few observationsby Snyder and Rio Abajo Forest (White et al. 2005). Kepler (1987). Additional information on the Puerto RIcan STUDY AREA AND METHODS Broad-winged Hawk's reproductive ecology and StudyArea. We studied BroadswingedHawks in the Rio nest habitat use is required to better understand Abajo Forest and surrounding private lands in Puerto Rico from 2000-02 (Fig. 1). The Rio Abajo Forest current limiting factors, and provide recommen- (18ø20'N, 66ø42'W) is managed by the ForestryDivision dations for future research on habitat conservation of DNER and is in west-central Puerto Rico within the in public and private lands. Herein, we report on moist limestone region of the island (Ewel and Whitmore the nesting ecology and behavior of the Broad- 1973). This forestreserve comprises an area of 2300 ha winged Hawk in a forest reserve in the moist lime- with elevations ranging from 200-420 m. We obtained climate data for the studyperiod from the site closestto stone forest region of Puerto RIco. Specifically,we our study area, the Dos Bocas NOAA weather station provide baseline information on courtship behav- (NOAA 2002). ior and territorial defense, nest successand pro- Annual precipitation during our study averaged 18.3 ductivity,prey delivery rates by nesting pairs, and cm (range = 6.9-34.9 cm) in 2001 and 14.7 cm (range = 5.2-45.1 cm) in 2002. Mean annual temperature was their habitat use. Moreover, we discussthe impli- 25.3øC(range = 19.9-30.6øC)in 2001 and 25.5øC(range cations our results on interactions between the = 20.1-30.9øC) in 2002. Broad-wingedHawk and Red-tailedHawk (Buteoja- The rugged limestone region (i.e., karst) of Puerto 406 HENGSTENBERGAND VILELLA VOL. 39, NO. 4 Rico encompasses27.5% of the island'ssurface (Lugo et Buteos, we observed territorial interactions between al. 2001). Topographyin this region is extreme and char- Broad-wingedHawks and intruding Red-tailed Hawks acterized by karst formations of subterraneandrainages, from limestone hilltops. We recorded the behavior of caves,dome shapedhills or "mogotes,"and deep sink- both birds (aggressiveor passive)and the end result (de- holes. Karst forest containsthe largesttree speciesrich- terred or not deterred). We used a binomial sign test for nessof Puerto Rico (Lugo et al. 2001). Rio Abajo Forest a single sample (Daniel 1990, Sheskin2000) to test the is fragmented on the easternend by a double lane high- hypothesisthat both speciesof raptorsdisplaced the oth- way and in the south-centralpart by a small community er randomly during aerial displays. (Fig. 1). About75% of the forestis within the subtropical Nest Searches and Monitoring. Broad-wingedHawk wet zone, the remaining quarter lies within the subtrop- use areas were delineated through direct observations ical moist zone (Ewel and Whirmore 1973). from limestonehills and hawk locationswere plotted on Previous studies indicated Broad-winged Hawks in USGS topographic quadrangles.We then extensively Puerto Rico have a limited geographicrange, and their searched areas with radio-marked adults and document- abundance is higher in the Karst region compared to ed aerial displaysor other territorialbehavior. Spot maps other life zones on the island (Delannoy 1997). As our and historical nesting information were used to extend primary objectiveswere to expandcurrent knowledgeon our searchesinto other potential nesting territories. All breeding ecology,habitat use, and movementpatterns of potential nestswere monitored for reproductiveactivity Broad-wingedHawks (Hengstenberg and Vilella 2004), beginning in February of each year. we selectedthe Rio Abajo Forest. Moreover,we were in- When a nest site was located, we built observation terested in comparing our results with findings of re- blinds 50-100 m from the nest tree at a location on a cently completedstudies in Rio Abajo Forest (Delannoy nearby cliff wall looking down on the nestwith clear line- and Tossas2002). of-sightvisibility. In one instance,it wasnot possibleto Vegetation of Rio Abajo is comprisedof a mix of sec- build a blind with a direct view because of nest location, ondary growth forestsand timber plantations (Cardona densevegetation, or steeprock walls.In this instance,the et al. 1987). The midstoryof some areasof secondary nest was monitored from the ground at a similar dis- forest was characterizedby abandoned shade-growncof- tance. Nest activities were recorded throughout the fee and citrus plantations.Forest overstories were domi- breeding seasonusing spotting scopes,video cameras, nated by moca (Andirainermis), capa prieto (C0rdiaallio- and binoculars from the observation blind. The distance dora), and guaraguao ( Guareaguidonia). Approximately (m) betweenall occupiednests (nest-site spacing) for the 6.9% of the forest (171.7 ha) are managedtimber plan- 2001 and 2002 breeding seasonswas measured on the tations. Some standsare activelymaintained (open un- ground with surveyingtape and verified on a studyarea derstory), while others