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Ecological Relationship of Two Todies in Hispaniola: Effects of Habitat and Flocking

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Ecological Relationship of Two Todies in Hispaniola: Effects of Habitat and Flocking The Condor98:169-719 0 The CooperOrnithological Society 1996 ECOLOGICAL RELATIONSHIPS OF TWO TODIES IN HISPANIOLA: EFFECTS OF HABITAT AND FLOCKING’ STEVEN C. LATTA~ AND JOSEPH M. WUNDEIUE, JR. International Institute of Tropical Forestry, U.S.D.A. Forest Service, P.O. Box 490, Palmer, Puerto Rico 00721 Abstract. We studied microhabitat use, foraging and social behavior of Broad-billed (Todus subulatus)and Narrow-billed (T. angustirostris)Todies in two areasof sympatry in the Cordillera Central of the Dominican Republic. Solitary Broad-billed and Narrow-billed Todies occupied distinct microhabitats in both shade coffee plantations and native pine forest while generally sharing similar foraging strategies.In both habitats, Broad-billed Todies foraged higher in the vegetation and occurred in more outer horizontal positions with lower foliage density than did their congener.Movement ratesand feedingrates differed significantlybetween the two species,with the Narrow-bill being the more active species. Changesin foragingbehavior by both speciesof todies were observedwhen they associated with mixed-speciesflocks in pine forest. We noted a decreasein some measuresof spatial overlap of todies in interspecificflocks, but other feeding behaviors tended to converge. Key words: foraging behavior;mixed-species flocks;Hispaniola; Todus; insectivores;hab- itat use. Sinopsis. Estudiamos el uso de1microhabitat, comportamiento alimenticio y social de Todussubulatus y T. angustirostrisen dos areas simpatricasde la Cordillera Central de la Republica Dominicana. Aves solitarias de ambas especiesocupan distintos microhabitates en plantacionesde cafe de sombra y en 10sbosques nativos, mientras comparten estrategias de forrajeo generalmentesimilares. En ambos habitates T. subulatusse alimenta m&s alto en la vegetation mientras se encuentraen posicioneshorizontales m&s hacia afuera, con una densidad de follaje menor que su congenere.Las razones de movimiento y alimentaci6n difieren grandementeentre las dos especies,siendo el T. angustirostrisla especiem&s activa. Se observaron10s cambios en el comportamiento de alimentaci6n cuando estasespecies se asociabanen bandadas de especiesmixtas en 10sbosques de pinos. Notamos una merma en el solapamiento espacial en las bandadas interespecificas,pero otros comportamientos de alimentacmn mostraron una tendencia a converger. INTRODUCTION solitary individuals with individuals in mixed- Patterns of resourceuse among avian sympatric species flocks (Morse 1970, Austin and Smith congenersmay changein the presenceof mixed- 1972, Alatalo 1981), and color-banded birds species foraging flocks. Plocking behavior, hy- which forage solitarily and as group members pothesized to increase fitness through enhanced (Valburg 1992). Plocking may result in the con- foraging or decreasedlikelihood of predation (re- vergence of foraging behavior of flock partici- viewed by Powell 1985, Hutto 1994) may fa- pants (Pearson 197 1, Buskirk 1972 cited in Pow- cilitate the exploitation of foraging locations or ell 1985, Valburg 1992) and a reduction in re- tacticsthat a bird would not otherwiseuse (Krebs source partitioning (Morse 1970, Powell 1985), 1973,Buskirk 1976,Valburg 1992),ormayforce indicating possible social learning and copying a bird to modify foraging behavior because of (Krebs 1973, Morse 1978). Decreased foraging potential competition with so many birds in close overlap in flocks is generally interpreted as in- proximity (Hutto 1988). Changesin foraging be- dicating interference competition (Alatalo 198 1). havior have been suggestedin studiescomparing Studies of how resource partitioning is affected by the interaction of species,social organization, and habitat have not been published for the trop- ics (but see With and Morrison 1990). I Received11 March 1996. Accepted17 July 1996. * Present Address: Division of Biological Sciences, Hispaniola supportstwo closely related species 110 Tucker Hall, University of Missouri, Columbia, of todies and provides an ideal system for study- MO 65211, e-mail: [email protected] ing resourcepartitioning and the effectsof habitat [7691 170 STEVEN C. LATTA AND JOSEPH M. WUNDERLE, JR. and social organization on foraging behavior of ious citrus species(Citrus spp.), and banana or congeners. Broad-billed (Todussubulatus) and plantains (Muss spp.) were also scattered Narrow-billed (T. angustirostris)Todies, thought throughout some plantations where they provid- to have evolved in isolation when Hispaniola ed an intermediate layer above the coffee. In a was separatedinto two islands during Pleistocene few plantations, an occasionalpine (Pinussp.) or glacier melts (Kepler 1977), continue to be geo- palm (Roystoneasp.) extended into the oversto- graphically isolated over much of the island (Ke- ry. The predominant variety of coffee (Coffea pler 1977, Dod 1981). The Broad-billed Tody arabica)in the plantations was the traditional occursfrom sealevel to 1700 m and prefersxeric “tipica” variety, although “caturra” predomi- scrub habitats and broadleaf and pine forests. nated in some of the larger plantations. Most of The Narrow-billed Tody is found at higher ele- the shadecoffee plantations were relatively small vations (800-3,200 m) and most often occursin (X = 1.9 ha, range = 0.1-9.7 ha). moist forests and ravines. The two speciesare Coffee is cultivated in areas that were origi- sympatric, however, in mangroves and wet forest nally pine (P. occidentalis)forest. Pine forests, on the Samana Peninsula, over a broad range many of which have been selectively logged, re- (465-1,730 m) in the Sierra de Baoruco, and in main on the steeper slopes and ridge-tops, with portions of the Cordillera Central (Kepler 1977; variable amounts of broadleaf understory that pers. observ.). Both speciesmost frequently oc- may be affected by fire, cutting, or grazing. cur as single birds or in pairs throughout their Broadleaf specieswere not sampled but included ranges, but join mixed-species flocks in the pine Ribessp., I. Vera,mango (A4. indica),citrus (Cit- forestsof the Cordillera Central (Latta and Wun- russp.), Cecropiasp., Syzygiumjumbos, Chia derle 1996). sp., Miconia sp., as well as numerous unidenti- In this study we compared the foraging be- fied shrubs. Pine forest was sampled at six sites havior of Broad-billed and Narrow-billed Todies on slopesnear Manabao (90%1,050 m) and six in two areasof sympatry in the Cordillera Central sites near Jarabacoa(643-779 m). Although the of the Dominican Republic to determine if re- area of the pine forest was not measured, the sourceuse and foraging behavior varied by spe- remnant patches were all estimated to be > 100 cies, habitat, or social organization of the birds. ha. We compared foragingbehavior between the two Foliage height profileswere determined for each species in shade coffee plantations and native habitat where feeding observations were collect- pine forest, and within specieswhen foraging sol- ed (Wunderle and Latta, in press).Foliage height itarily and in mixed-speciesflocks in pine forests. profiles for the two habitat types showedgeneral We hypothesized that flocking allows similar structural similarity with a fairly open canopy, a speciesto copy foraging locations and behaviors, densercoffee or mixed broadleaf understory, and and that todies gain foraging benefits from such very little in the way of an intermediate layer. In behavior. We predicted that (1) when foraging the shadecoffee plantations, Zngaprovided most solitarily in similar habitat each specieswould of the shadeoverstory, with greatestcover in the occupy a distinct microhabitat and interspecific lo-12 m height category. Pine foliage predom- behavioral differences would be consistent be- inated in the canopy of the pine forest, although tween habitats, (2) foraging behavior and micro- scatteredbroadleaf trees also extended into the habitat use by the two specieswould converge canopy, and pine sites had a slightly higher can- in mixed-species flocks, and (3) foraging rates opy with greatest cover in the 15-20 m height would increase in the presenceof mixed-species category. The broadleaf understory of the pine flocks. forest provided a sparserfoliage cover within the different height categoriesthan that provided by STUDY AREA AND METHODS coffee in the shade coffee plantations, except Todies were studied in fourteen shade coffee within the first 1.0 m where foliage cover was plantations in the vicinity of Manabao and Jar- greater in the pine sites. abacoa, La Vega Province, Dominican Republic at elevations of 540-850 m. Shade coffee plan- FEEDING BEHAVIOR tations were characterized by a predominant Foraging observations were made in the non- overstory of ZngaVera, although mango (Man- breedingseason from October-March 1993-l 995 giferaindica), avocado (Perseaamericana), var- while walking slowly through the sites. Obser- ECOLOGICAL RELATIONSHIPS OF TODIES 171 vations were made from 07:00-12:00 in coffee Becausecanopy heights varied somewhat be- plantations, but extended to 18:OOin pine forests tween sites,bird height relative to canopy height where feeding activity occurredall day. Foraging was calculated and used in further analyses. In observations were made by walking slowly all statistical analysesfoliage density scoresof O- through the habitat until a foraging bird was lo- 2 and 4-5 were combined to reduce the number cated. The first foraging event five secondsafter of sparsecells. Similarly, for analyses of feeding an individual was initially detectedwas recorded behavior
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