Avian Research in the Caribbean: Past Contributions and Current Priorities Steven C

Journal of Field Ornithology

J. Field Ornithol. 83(2):107–121, 2012 DOI: 10.1111/j.1557-9263.2012.00361.x

Avian research in the Caribbean: past contributions and current priorities Steven C. Latta1

National Aviary, Allegheny Commons West, Pittsburgh, Pennsylvania 15212, USA Received 6 December 2011; accepted 12 February 2012

ABSTRACT. The islands of the Caribbean contain habitat of critical importance to a large number of endemic and resident birds, as well as many overwintering Neotropical migrants, and they rank as a globally outstanding conservation priority ecoregion and biodiversity hotspot. Considerable research from the region has focused on the ecology of permanent resident species, and these studies have had particular signiﬁcance for threatened species management, especially parrot biology and conservation, but also for tropical community ecology in general. Work by ornithologists in the Caribbean has been instrumental in improving our understanding of the ecology of overwintering Neotropical migrants and in developing long-term monitoring programs. Although Caribbean-based studies of birds have resulted in signiﬁcant contributions in many important areas of ecological research, there is a great need for additional research. Especially needed are studies with application to the management of resident species, and studies of how bird populations may be affected by pathogens, parasites, plants, and other types of biotic interactions. Studies focusing on how bird species and populations are affected by global climate change, and cumulative, landscape-level changes in land use are also needed. Along with additional research, scientists have an important role to play in building capacity to prepare the next generation of biologists in the region who will need to address mounting challenges related to biodiversity protection. As with many conservation efforts, funding is a critical need for almost all organizations and agencies involved in research, conservation action, and capacity building in the West Indies.

RESUMEN. La investigacion´ ornitologica´ en el Caribe: contribuciones pasadas y prioridades actuales Las islas del Caribe contienen habitat´ de importancia cr´ıtica para un gran numero´ de aves endemicas´ y residentes, as´ı como muchos migrantes Neotropicales durante el invierno, y figuran como una ecorregion´ prioritaria para la conservacion´ y de importancia para la biodiversidad a nivel mundial. Muchas investigaciones en la region´ se han concentrado en la ecolog´ıa de las especies residentes permanentes, y estos estudios han tenido un significado particular para la gestion´ de especies amenazadas, especialmente en cuanto a la biolog´ıa y conservacion´ de loros, pero tambien´ para la ecolog´ıa de comunidades tropicales en general. El trabajo de ornitologos´ en el Caribe ha sido instrumental en mejorar nuestro conocimiento de la ecolog´ıa de las aves migratorias Neotropicales durante el invierno y en el desarrollo de programas de monitoreo a largo plazo. Aunque los estudios de aves basadas en el Caribe han hecho contribuciones significativas en muchas areas´ importantes de investigacion´ ecologica,´ hay una gran necesidad de investigacion´ adicional. Se necesitan en especial estudios con aplicacion´ a la gestion´ de las especies residentes, e investigaciones de como´ las poblaciones de aves pueden ser afectadas por las interacciones con patogenos,´ parasitos,´ plantas, y otros tipos de interacciones bioticas.´ Estudios enfocados en como´ las especies y poblaciones de aves son afectadas por el cambio climatico´ global y por cambios cumulativos a nivel del paisaje en el uso de la tierra tambien´ son necesarios. Ademas´ de realizar mas investigaciones, los cient´ıficos tienen un papel importante que desempenar˜ en capacitar la proxima´ generacion´ de biologos´ de la region,´ en prepararlos para enfrentar los crecientes desaf´ıos relacionados con la proteccion´ de la biodiversidad. Al igual que muchos esfuerzos de conservacion,´ el financiamiento es una necesidad cr´ıtica para casi todas las organizaciones y agencias involucradas en la investigacion,´ accion´ conservacionista y capacitacion´ en las Indias Occidentales. Key words: biodiversity, capacity building, endemic birds, Greater Antilles, Neotropical migrants, West Indies

The islands of the Caribbean contain habitat 90% of the land area, and contain the greatest of critical importance to a large number of biological diversity in the region (FAO 1991, resident bird species as well as many winter- Keith et al. 2003). The Bahamas, Turks and ing Neotropical migrants. In the Caribbean, Caicos, Bermuda, and the Lesser Antilles are the Greater Antillean islands of Cuba, Jamaica, also home to many bird species, including some Hispaniola, and Puerto Rico encompass nearly endemics. Of the ∼770 bird species found on Caribbean islands, 148 are endemic to the region, and 105 are conﬁned to a single island 1Email: [email protected]

C 2012 The Author. Journal of Field Ornithology C 2012 Association of Field Ornithologists 107 108 S. C. Latta J. Field Ornithol. (BirdLife International 2008). In the region, 54 understanding of the ecology of overwinter- species are threatened with extinction, and 12 ing Neotropical migrants, developing long-term are considered Critically Endangered (BirdLife monitoring programs, and understanding how International 2008). In addition, many migra- evolutionary and ecological processes, including tory species visit the Caribbean, with some biotic interactions, generate biological diversity. populations wintering either primarily or ex- Below I address each of these areas of research. clusively in the region (Faaborg and Terborgh My review is not intended to be encyclopedic, 1980). Global biodiversity assessments recog- and my focus is on key papers that have made nize the importance of the region (Myers 1988, the most significant contributions to ornithol- Stattersfield et al. 1998, Mittermeier et al. ogy. In addition, I suggest areas of research 1999, WCPA Caribbean 2003) and rank the where ornithologists in the Caribbean can make Caribbean as a globally outstanding conserva- important contributions, and conclude with tion priority ecoregion and biodiversity hotspot, ideas on how the research community can help distinguished by high terrestrial endemism and advance conservation efforts. little pristine vegetation. The early history of research and bird moni- RESEARCH CONTRIBUTIONS toring in the Caribbean basin followed a similar pattern on different islands (Latta et al. 2003a). Threatened species management. Early Most work prior to the 1970s was limited studies of the ecology of Caribbean birds often to species lists and natural history observations, focused on endemic species, including Kepler’s and these were usually published as notes, in- (1977) monograph on todies and Short’s (1974) cluded in checklists, or summarized in bird study of West Indian woodpeckers. Other work guides and special publications. A number of focused on the ecology of threatened species and annotated checklists were published that sum- provided the basis for management and conser- marized the basic biology of birds, includ- vation activities. Many studies were conducted ing occurrence, distribution, breeding periods, in Puerto Rico where an unfortunate wealth of and habitat use (Keith 1997, Bradley 2000, threatened species coincided with funding avail- Keith et al. 2003, Buckley et al. 2009). These ability and legal mandates for action. Species were followed by summaries of the distri- of particular concern included Puerto Rican bution and status of birds including com- Nightjars (Caprimulgus noctitherus) threatened pendiums of seabird breeding sites (Bradley by deforestation and habitat change (Kepler and Norton 2009) and important bird areas and Kepler 1973, Vilella and Zwank 1993, (BirdLife International 2008) that were de- Vilella 2008), and Yellow-shouldered Blackbirds veloped for conservation planning purposes. (Agelaius xanthomus) that were endangered pri- Although previous papers have included sum- marily because of brood parasitism by Shiny maries of advances within narrowly defined Cowbirds (Molothrus bonariensis;PostandWiley research themes such as Neotropical migratory 1977, Post 1981, Wiley et al. 1991), as were bird ecology (Holmes 2007, 2011, Faaborg Puerto Rican Vireos (Vireo latimeri; Woodworth et al. 2010b) or biogeographic patterns of bird 1997, Woodworth et al. 1998). distribution (Ricklefs 2011), a comprehensive Elsewhere, Staus (1998) studied West Indian summary of ornithological research in the region Whistling-Ducks (Dendrocygna arborea)inthe has never been published. Bahamas, and emphasized the importance of My objective was to fill that void and explain protected wetlands to their conservation. This how studies of the ecology and evolution of birds work was instrumental in the establishment of a in the Caribbean islands have contributed to West Indian Whistling-Duck conservation pro- the broader field of ornithology. Although much gram focused on community education and pro- work from the region has focused on the ecology tection of coastal wetlands (Sutton et al. 2004). of resident species, particularly important stud- Another declining coastal species, the White- ies have focused on management of threatened crowned Pigeon (Columba leucocephala), was species, parrot biology and conservation, and studied in Puerto Rico, leading to an impressive tropical community ecology. Theoretical studies monograph (Wiley and Wiley 1979b). A study by ornithologists working in the Caribbean of Caribbean Flamingos (Phoenicopterus ruber) have also been instrumental in advancing our by Allen (1956) led to conservation efforts on Vol. 83, No. 2 Avian Research in the Caribbean 109 Great Inagua in the Bahamas, as well as similar serve Hispaniolan Parrots (Amazona ventralis). efforts in Cuba, Bonaire, and elsewhere. Efforts In addition, Butler (1992) described innovative to conserve flamingos continue through broad methods for conserving parrots in the Lesser An- census and monitoring programs (Wiley and tilles. Now exported worldwide, Butler’s (1992) Wiley 1979a, Paulino et al. 2010). pride program identifies endangered birds as Other studies have also had important man- unique symbols of nationhood, changing hu- agement and conservation implications, includ- man attitudes and ultimately leading to increases ing studies of Montserrat Orioles (Icterus oberi), in parrot populations. a species that faces unique threats from vol- Tropical community ecology. As sug- canic activity (Hilton et al. 2003, Dalsgaard gested above, studies conducted in the et al. 2007), as well as two Hispaniolan species, Caribbean have generated a rich literature on Ridgway’s Hawks (Buteo ridgwayi;Wileyand avian brood parasites. This is in part a re- Wiley 1981), and White-necked Crows (Corvus sponse to the arrival of Shiny Cowbirds in the leucognaphalus; Wiley 2006), that are threatened Caribbean and their exploitation of the island by hunting and habitat loss. Also on Hispan- avifaunas (Cruz et al. 1985, Wiley 1985, 1988, iola, Latta et al. (2000) examined the nesting Post et al. 1993). Comparing host responses to and foraging habitats of Hispaniolan Crossbills brood parasitism in areas with a long history (Loxia megaplaga), emphasizing the need to of parasite–host interaction to those in areas control wildfires affecting mature pine forests. where cowbirds recently arrived, Cruz and Wiley Similarly, Hayes et al. (2004) noted the impor- (1989) found changes in nest placement, nest tance of mature stands of Caribbean pines (Pinus defense, and productivity, as well as differences caribaea) for conserving the remaining popula- in egg acceptance and rejection by hosts. In tion of Bahama Nuthatches (Sitta pusilla). addition, Cruz et al. (2008) examined how these Parrot biology and conservation. Be- behaviors changed over time with increased cause of their charismatic nature and their exposure to cowbirds. often threatened population status, psittacids Classic studies of community assemblages have been the focus of numerous studies in were conducted in Jamaica (Diamond 1974, the Caribbean. Eleven of the surviving parrot Cruz 1974), Grand Bahama Island (Emlen species in the West Indies are endemics, and 1977), and Cuba (Acosta Cruz and Mugica all are of conservation concern. Gnam (1991) Valdes´ 1988). The low species richness of birds studied the breeding biology of Bahama Parrots on Caribbean islands allowed detailed stud- (Amazona leucocephala bahamensis), and other ies of hummingbird-plant webs (Lack 1976, investigators have examined the distribution, Dalsgaard et al. 2008, 2009), and of a tropical abundance, and threats facing several other par- forest food web in the Luquillo Experimental rot species in the Lesser Antilles (Gochfeld 1974) Forest (Puerto Rico), with the latter resulting and Greater Antilles (Wiley 1991). Puerto Rican in the most complete summary of the role of Parrots (Amazona vittata) have been the focus of birds in a tropical forest food web (Waide 1984). considerable research and management efforts. More recently, Arendt (2006) studied Pearly- A classic monograph on the species (Snyder eyed Thrashers (Margarops fuscatus) and argued et al. 1987) was followed by a study by Beissinger that they are preeminent avian “supertramps.” et al. (2008) focusing on the relative impor- With generalized nesting and foraging strategies, tance of genetic, demographic, environmental, these thrashers are able to adapt to the ever- and catastrophic processes in maintaining a changing conditions in the region’s natural and population bottleneck. In addition, efforts to anthropogenic environments, and they now oc- reintroduce Puerto Rican Parrots have led to cur on often disturbed, species-poor islands and the development of methods for captive-rearing, habitats throughout the Caribbean. In another nest-site management, and relocation (Snyder important study, Mugica Valdes´ et al. (2006) et al. 1987, Brock and White 1992, Collazo examined the ecology and conservation status et al. 2000, 2003, White and Vilella 2004, of wetland birds in Cuba, and described strate- White et al. 2005, Koenig et al. 2007). At gies for conserving the birds and their wetland the interface between research and conservation, habitats. White et al. (2011) described how local residents Studies conducted in the Caribbean have also could become involved in research to help con- contributed to our understanding of the effects 110 S. C. Latta J. Field Ornithol. of hurricanes on birds (Wunderle 1995b, 1999). were more important than patch area and patch The greatest impacts occur after a hurricane, isolation in determining local colonization and with nectarivores and frugivores affected by extinction probabilities. Together, these stud- interrupted food supplies, and highland birds ies reinforce the importance of differentiat- impacted by relatively slow plant regrowth at ing among land cover and land uses in frag- higher, cooler elevations (Waide 1991, Askins mentation research and management. Lessons and Ewert 1991, Wunderle et al. 1992, Wiley from landscape-level studies have been invoked and Wunderle 1993, Wunderle 2005). in proposed conservation measures for threat- A number of important studies in the ened Bicknell’s Thrushes (Catharus bicknelli); Caribbean have addressed the impacts of an- Kerchner et al. (2009) modeled benefits of a thropogenic habitats on tropical birds, especially voluntary incentive program for landowners in their responses to monocultures and forests the Dominican Republic where changes in land dominated by introduced tree species (Cruz use would yield a cost-effective network of pro- 1987, 1988). Other investigators have compared tected habitat that would sustain overwintering avian communities in native forests to those thrush populations. found in Caribbean pine plantations (Collazo Ecology of Neotropical migratory birds. and Bonilla Martinez 1988), sun and shade Beginning in the mid-1970s, the ecology of coffee plantations (Wunderle and Latta 1996), Neotropical migrants overwintering in the and several agricultural habitats in Jamaica, Caribbean attracted increased attention (Keast including coffee and citrus (Johnson 2000, and Morton 1980). The results of early stud- Johnson et al. 2006). The results of these studies on overwintering ecology were summa- ies suggest that arboreal agricultural habitats rized by Holmes (2007, 2011) and Faaborg (e.g., coffee, citrus, and cacao) can play a role et al. (2010b), with these studies generally focus- in the conservation of generalist, insectivorous ing on habitat specific demographics and the site birds, and that plantations may aid efforts to fidelity of warblers (Holmes et al. 1989, 1996, reforest abandoned agricultural fields. Studies Wunderle 1995a, Marra et al. 1998, Wunderle on Hispaniola, and later Jamaica, were among and Latta 2000, Marra and Holmes 2001, Latta the first to reveal the value of shade grown coffee and Faaborg 2001, 2002). These studies often plantations as bird habitat (Wunderle and Latta documented strong winter territoriality, gener- 1996, 1998, 2000, Johnson 2000), and follow- ally with nonoverlapping territories (Holmes up studies in Jamaica were the first to show that et al. 1989, Wunderle 1995a, Wunderle and birds can control coffee borers (Hypothenemus Latta 2000, Latta and Faaborg 2001, 2002), hampei), an important coffee pest (Kellermann differences in sex and age ratios among habitats et al. 2008), and to place a monetary value on the resulting from dominance interactions (Holmes ecosystem services provided by birds in coffee et al. 1989, Parrish and Sherry 1994, Wunderle plantations (Johnson et al. 2009, 2010). 1995a, Marra 2000, Wunderle and Latta 2000, Investigators conducting studies in the Marra and Holmes 2001, Latta and Faaborg Caribbean have also examined how species and 2002), and site faithfulness between winters communities respond to variation in landscape (Holmes et al. 1989, Marra and Holmes 2001, characteristics. Tossas (2002) analyzed source Latta and Faaborg 2001). Researchers developed and sink dynamics of Puerto Rican Vireos, models to determine when during the annual and Acevedo and Restrepo (2008) showed how cycle populations of migratory passerine birds changes in land cover and land use across might be limited (Sherry and Holmes 1995, Puerto Rico influenced the large-scale organi- 1996, Latta and Baltz 1997, Sherry et al. 2005), zation of bird assemblages. In Jamaica, Kennedy and hypothesized that winter food availability et al. (2010) found that species richness, limits population sizes via habitat quality. Sup- community composition, and abundance were port for this hypothesis was provided by studies matrix-dependent, with agricultural landscapes demonstrating the relationship between food supporting greater avian diversity and more availability and the physical condition of birds intact community assemblages than either peri- (Strong and Sherry 2000, Latta and Faaborg urban or bauxite-mined landscapes. Kennedy 2001, Studds and Marra 2005, Brown and et al. (2011) found that within-patch vegetation Sherry 2006a, Diggs et al. 2011), decreased rates structure and the matrix type between patches of site persistence in habitats with reduced food Vol. 83, No. 2 Avian Research in the Caribbean 111 resources (Johnson and Sherry 2001, Latta and itoring programs in the Caribbean and Latin Faaborg 2002), and linkages of both population America (Rivera-Milan´ 1992, 1995a, b, 1997, responses and individual condition of nonbreed- Latta et al. 2003a, 2005, Faaborg et al. 2007). ing birds to prevailing ecological conditions Monitoring programs have also contributed to across divergent habitats (Latta and Faaborg our understanding of the impacts of stochastic 2002). The results of other studies suggested events such as hurricanes (Waide 1991, Askins a relationship between rainfall or local moisture and Ewert 1991, Wunderle et al. 1992, Wiley levels and habitat quality (Latta and Faaborg and Wunderle 1993, Wunderle 2005) and vol- 2001, Studds and Marra 2007). canic eruptions (Dalsgaard et al. 2007) on avian Other investigators focused more on carry- populations, and more intensive, often species- over effects, especially the impacts of winter specific, monitoring efforts have improved our habitat quality on later reproductive success. understanding of the ecology and conservation Marra and Holberton (1998) and Reudink et al. (Townsend et al. 2010), and population trends (2009) showed that nonbreeding season events (Rimmer et al. 2011) of threatened species such could have fitness consequences during both the as Bicknell’s Thrushes. nonbreeding and subsequent breeding season, Generating biological diversity. Carib- and that these seasonal interactions can play an bean islands have also been important in at- important role in regulating populations (Runge tempts to better understand how ecological and Marra 2005). For example, compared and evolutionary processes generate biological to those in optimal habitats, American Red- diversity. Studies conducted in the West Indies starts (Setophaga ruticilla) in suboptimal habitats were central to debates in community ecology had elevated corticosterone levels (Marra and about factors influencing bird diversity on is- Holberton 1998), lost mass over winter (Marra lands (MacArthur et al. 1966, Case et al. 1983, and Holmes 2001), departed later on spring Ricklefs and Bermingham 2001, Ricklefs 2011). migration (Marra et al. 1998, Studds and Marra Lack (1976) used Jamaican birds to illustrate his 2005), and had lower annual survival (Johnson theory of island biology, now discredited, that et al. 2006). These effects carry over to the breed- rested on the idea that habitats on an island ing grounds with redstarts from suboptimal determined the number of species present, with habitats arriving in poorer condition, settling on distance from the source of colonists playing territories later, and fledging fewer young (Marra little role. Ecological patterns of community et al. 1998, Norris et al. 2004). Stable-hydrogen structure included a search for “assembly rules” isotope ratios have also been used to show that such that the composition of birds in foraging habitat use during the first overwintering period guilds was not random but repeatable, and that plays a role in determining the direction and guilds on larger islands could become saturated distance traveled during the first spring migra- (Terborgh and Faaborg 1973, Terborgh et al. tion (Studds et al. 2008), further extending 1978, Faaborg 1980). In addition, coexisting the role of carry-over effects to natal dispersal guild members were shown to consistently differ patterns. in body size by body weight ratios of 2.0 Avian monitoring. Complementing field (Faaborg 1982, 1985, Case et al. 1983). These studies of overwintering migrants, investigators observations were used to argue that the overall in the Caribbean region have led in the develop- arrangement of species on islands is nonrandom, ment of monitoring strategies and implementa- and that these patterns are consistent with the tion of long-term avian monitoring programs. hypothesis that competition is a driving force in One of the longest bird monitoring programs structuring communities (Case et al. 1983). in the hemisphere was initiated in 1973 at Because of their small size, geographical iso- Guanica,´ Puerto Rico, and the results of this lation with defined boundaries, and simplified work have contributed to our understanding of communities, islands have also been central to the ecology of migrants during the nonbreed- improving our understanding of the role of evo- ing season (Faaborg and Arendt 1989, Faaborg lutionary processes in generating biological diet al. 2000, Faaborg 2002, Dugger et al. 2004). versity (Ricklefs and Bermingham 2001, 2007, In addition, lessons learned from this pioneering 2008, Ricklefs 2011). Although Caribbean birds monitoring effort were critical in developing rec- are not as well known and have not been used as ommendations for other constant-effort mon- textbook examples of adaptive radiation, Burns 112 S. C. Latta J. Field Ornithol. et al. (2002), in a study designed to identify RESEARCH PRIORITIES the sister taxon of Darwin’s finches, showed that their closest living relatives were largely Although studies of Caribbean birds have Caribbean endemics. Caribbean birds also show made numerous important contributions, some a remarkable diversity of bill types and feed- areas of investigation have received less attention ing behaviors similar to that observed among (Latta et al. 2003a, Faaborg et al. 2010a). Below Darwin’s finches. Studies conducted in the West I summarize some of the more critical needs. Indies have also illustrated the importance of Ecology of permanent resident species. allopatry for speciation, with Townsend et al. Much of the work on West Indian birds has (2007) and Sly et al. (2010, 2011) using phylo- focused on several key threatened species or a few genetic techniques and a knowledge of geologic species of overwintering Neotropical migrants. events to eliminate the possibility that sympatric As a result, little is known about the basic natural speciation explained the presence of sister species historyofmanyspecies,includingsomeofthe on Hispaniola. The results of these studies also most threatened endemics and most of the birds lend support to Diamond’s (1977) assertion that currently considered game species. However, avian speciation is unlikely to occur on single this lack of information is not equally shared islands smaller than New Guinea. Nevertheless, among all Caribbean islands; more is known, Coyne and Price (2000) noted that the existence in general, about the natural history of birds of endemic sister species of hummingbirds (Red- in Puerto Rico, Cuba, and Jamaica than those billed [Trochilus polytmus] and Black-billed [T. on some of the smaller or more impoverished scitulus] streamertails) on Jamaica that may be islands. evidence of sympatric speciation. To effectively manage bird populations, we Finally, many of these concepts come together need species-specific studies to determine cur- in the idea of the taxon cycle (Wilson 1961), an rent distributions, habitat associations, timing of idea closely associated with West Indian birds breeding, reproductive rates, and sex- and age- (Ricklefs and Cox 1972). Studies of Caribbean specific survival rates. In addition, few studies birds suggest that species progress through cycles have used a multiscale perspective to determine of geographic expansion via dispersal followed how landscape-level habitat features may inter- by adaptation to local environments (espe- act with nest-site, territory, habitat, and local cially on the margins of their range), diver- patch effects to impact reproductive success and gence and extinction of some populations, and survival. Studies that include factors such as greater ecological specialization of remaining habitat patch size, proximity to edge, and charac- populations, especially in forest-interior habitats teristics of the matrix surrounding a patch have (Ricklefs and Cox 1972). Some species may rarely been conducted in the Caribbean, despite evolve secondary distributions and initiate new their potential importance for understanding cycles of expansion to continue the cycle. the ecology of many species. Ricklefs and Bermingham (2001) used molec- Biotic interactions. Biotic interactions, ular phylogenetic analyses of small land birds such as host–pathogen, competitive bird–bird, in the Lesser Antilles to lend support for the and mutualistic bird–plant interactions, have taxon cycle concept, while rejecting MacArthur not been well studied in Caribbean ecosys- and Wilson’s (1967) equilibrium theory of bio- tems. Although some work has been conducted geography based on nonequilibrium patterns on brood parasitism (Cruz and Wiley 1989), of colonization and extinction at evolutionary, patterns of distribution of unique strains of rather than ecological, time scales. In recent avian malaria among hosts (Ricklefs et al. 2004, years, molecular phylogenetic evidence has also Ricklefs and Bermingham 2008), and relation- been used to confirm the temporal sequence ships between overwintering migrants and per- of distribution patterns representing phases of manent residents (Latta and Wunderle 1998, species expansion and contraction (Ricklefs and Johnson et al. 2005), few investigators have Bermingham 2004), and to further suggest that studied how biotic interactions affect population the cycles may be generated by coevolutionary dynamics and the distribution of species. Failure relationships between species and their predators to incorporate biotic interactions into ecological and pathogens (Ricklefs and Bermingham 2002, studies is currently viewed as a major reason Losos and Ricklefs 2009, Ricklefs 2011). why many models fail to fully describe bird Vol. 83, No. 2 Avian Research in the Caribbean 113 populations and their distribution (Kissling et al. overwintering migrants in the Caribbean have 2012). Describing biotic interactions through, been conducted at just a few sites in a few for example, network analyses, is not only use- habitats. In addition, because most work has ful for understanding distributional patterns, focused on just a few species, little is known but for clarifying interdependencies between about other overwintering migratory species, or plants and their bird and insect pollinators, whether models developed for the better-studied functional specialization, resilience, and stability species apply to others. There is a need for a more (Dalsgaard et al. 2008, Kissling et al. 2012). widespread assessment of which species occupy Network analysis is also useful for evaluating which habitats, and a broader understanding of and predicting ecosystem responses in the face how variation in occupancy and survival across of habitat change, species loss, and global climate the entire wintering range impacts each species. change. Although most widespread natural habitats have The Caribbean also offers opportunities received some attention from researchers, some for studies to improve our understanding of habitat types are almost completely unstudied, competitive relationships among and between including oil palms, cacao and other agricultural resident and migratory species, and how habi- habitats other than coffee (but see Johnson et al. tats support overwintering migrants in addition 2006), pastures and mesquite-dominated shrub- to permanent residents. Recognizing that in- lands, grasslands, wetlands, lowland forests, and sects may control the size of bird populations, urban parks. Although wildlife managers may Greenberg (1995) proposed the “breeding cur- be most interested in knowing what levels of rency hypothesis” to explain the apparent para- disturbance migratory birds will tolerate, with dox of arthropod food sources for birds being the exception of shade coffee (Wunderle and at their lowest during the nonbreeding season Latta 2000), few studies of survival have focused when bird numbers peak. The hypothesis con- on anthropogenic habitats (but see Johnson et al. trasts the abundance of large, high quality, soft- 2006) or the value of early successional habi- bodied arthropods that may limit numbers of tats for migrants (but see Wunderle et al. permanent resident birds during reproduction, 2010). Finally, as with studies conducted dur- with the abundance of relatively low-quality, ing the breeding season (see “Ecology of per- small insects that may maintain greater num- manent resident species”), application of a bers of adult residents as well as overwintering multiscale perspective is required to fully un- migrants during the nonbreeding period. In derstand the nonbreeding season ecology of the only test of the hypothesis, Johnson et al. migrants. (2005) found general support, but suggested Temporal expansion of studies. Few that other factors are also operating because studies of resident birds in the Caribbean the ratio of migrants to residents was signifi- have focused on their ecology outside of the cantly higher than predicted in some habitats. breeding season, and few published studies of Additional data are needed concerning how overwintering migrants encompass the entire changes in bird distributions, and seasonal shifts nonbreeding season. In addition, use of stopover in habitat or diet by permanent residents, may sites by passage migrants in the Caribbean is create ecological “vacancies” for overwintering surprisingly understudied. Although shorebird migrants (Ricklefs 1992, Johnson et al. 2005). counts have been conducted on a number of We also need a better understanding of how islands, few results of these counts appear to have intratropical migrants such as some populations been published (Wunderle et al. 1989, Collazo of Gray Kingbirds (Tyrannus dominicensis)and et al. 1995). We know from observational Black-whiskered Vireos (Vireo altiloquus)arein- records that many landbird migrants use a tegrated into breeding bird communities in the wide variety of habitat types and even habitat Caribbean (Johnson et al. 2005). In addition, fragments during migration, but little is known we need to better understand how relationships about the distribution or quality of these sites for between permanent residents and overwintering migrants in the Caribbean, or whether there are migrants may be affected by habitat alteration concentrations of migrants at particular points and changes in habitat quality. during the migratory periods (but see Rodr´ıguez Habitat-specific survival of overwinter- et al. 1994, Rodr´ıguez and Sanchez´ 1995, Latta ing migrants. Most studies of the ecology of and Brown 1999, McNair et al. 2002). 114 S. C. Latta J. Field Ornithol. Citizen science projects such as eBird Carib- uncovered elsewhere will help determine if the bean (http://ebird.org/content/caribbean), decline in numbers of migrants at Guanica,´ Caribbean BirdWatch (http://conserveonline. Puerto Rico, are general or specific to that org/workspaces/caribbeanbirdwatch), and the site. Caribbean Waterbird Census (http://www. We also need monitoring to document the scscb.org/) use simple monitoring or obser- spread of exotic diseases such as West Nile vational data to “visualize” migration of species virus or avian influenza, and to understand on maps by plotting concentrations of birds at the impact of disease and parasites on avian different sites and habitats during migrations. populations. Other than studies of avian malaria Such projects should contribute to our under- (Ricklefs and Fallon 2002, Ricklefs et al. 2004, standing of the location and use of stopover sites. Fallon et al. 2004, 2005, Latta and Ricklefs However, we also need a better understanding 2010), botfly infections (Arendt 1985a, b), and of migrant ecology during the period of arrival scaley-leg mites (Latta and O’Connor 2001, and territory establishment. We do not know Latta 2003), few sampling efforts have fo- how territories are selected, how territorial indi- cused on emerging diseases or parasites in the viduals prepare behaviorally and physiologically Caribbean (but see Dupuis et al. 2003, Komar for spring migration, or if they abandon terri- et al. 2003), or on monitoring their impacts. tories to use different habitats during the spring Climate change. Climate change, re- fattening period prior to migration. flected in warming trends and more frequent Finally, not all individuals or species are extreme weather events, is affecting avian dis- territorial during the nonbreeding period. We tributions and the timing of breeding and need a better understanding of what drives the migration by birds around the world (Møller movements of floaters during the nonbreeding et al. 2010, Cox 2010). From a Caribbean period (but see Brown and Sherry 2008), the perspective, only recently have a limited number occurrence of altitudinal migration, and how of papers addressed climate change and birds, wandering or seasonal movements influence and these have focused primarily on how rainfall survival. may affect habitat quality for overwintering Documenting population trends. Des- migrants, thereby impacting spring departure pite some prominent efforts described above, schedules and breeding success through carry- there has been insufficient monitoring to fully over effects (Sillett et al. 2000, Smith et al. assess the status of vulnerable bird populations 2010, Wilson et al. 2011, Studds and Marra and their habitats in the Caribbean or to guide 2011). These studies have emphasized the need conservation strategies. Some highly threatened to understand within-season and annual vari- species such as the Puerto Rican Parrot are ation in precipitation to model the impacts monitored regularly (Beissinger et al. 2008), of climate change (Smith et al. 2010, Wilson and citizen-based efforts such as Caribbean et al. 2011). As Studds and Marra (2011) BirdWatch, Caribbean Waterbird Census, pointed out, migrants in the Caribbean may and eBird Caribbean have been inaugurated, be informative in modeling impacts resulting but long-term, locally intensive efforts (Latta from global climate change because they face et al. 2005) using mist nets are also needed. both declining rainfall in the overwintering areas The long-term monitoring study described by and increased temperatures on the breeding Faaborg et al. (2007) has been invaluable for grounds. This approach, using events through- detecting population changes, but similar efforts out the annual cycle to study population impacts are needed elsewhere. Constant-effort mist- and adaptive responses to climate change, is netting in a variety of habitat types should be unique. encouraged, but monitoring of birds in relatively Other than these studies that focus on mi- unperturbed native habitats is preferred; popu- grants and studies mentioned previously that lation declines in habitats assumed to be optimal have focused on the impacts of hurricanes, few would best signal the need for management or authors have investigated how climate change conservation activities (Faaborg 2000). Such may be affecting birds in the West Indies, or monitoring is needed to better understand how weather patterns (especially rainfall) may population trends of both permanent resident affect reproduction and survival of resident and overwintering migratory birds, and patterns species in the region (but see Faaborg and Vol. 83, No. 2 Avian Research in the Caribbean 115 Arendt 1989, Dugger et al. 2004). We need conservation organizations. Such collaboration data to better understand interactions among will result in studies more likely to reflect local climate change and global climate patterns such needs and interests, and can also help build as the El Nino˜ Southern Oscillation (ENSO) capacity. In addition, working with local citizens andLaNina˜ events. Several global warming can provide them with a sense of ownership models are consistent in predicting increased and personal investment in programs, and affirm summer droughts in the Caribbean basin over the perception that some benefits will accrue to the next 50 yr (Neelin et al. 2006), and recent them for their efforts (White et al. 2011). rainfall declines in the Bahamas (Martin and Funding is critical for almost all organizations Weech 2001), Puerto Rico (Heartsill-Scalley and agencies involved in research, conservation et al. 2007), and Jamaica (Studds and Marra action, and capacity building in the West Indies, 2007) are consistent with the predictions of and the funding devoted to these efforts can these models. The expected effects of these sum- have important impacts. For example, Latta and mer droughts include phenological disruptions, Faaborg (2009) analyzed patterns of funding declines in food availability, and an increase for research and monitoring using examples in fire frequency (Weaver and Gonzalez 2005). from Puerto Rico and the Dominican Republic, Models to help evaluate how Caribbean birds demonstrating that research has had an under- might respond to these combined threats are appreciated effect on the development of conser- needed, as is empirical data about bird con- vation capacity and conservation efforts in host dition and population trends. As previously countries (e.g., Ewert et al. 2009). Investments mentioned, studies that use species interaction in research contributed to an increase in the distribution models to better understand biotic training of students and wildlife professionals, interactions within guilds or across trophic levels promoted conservation awareness, played an at large spatial scales may be particularly infor- important role in the growth and profession- mative with respect to the effects of a changing alization of key environmental organizations, climate (Dalsgaard et al. 2008, Kissling et al. spawned a growing ecotourism industry for bird- 2012). watching, and drove planning and conservation efforts for birds in national parks. BUILDING CAPACITY FOR RESEARCH AND Researchers in the tropics should also be CONSERVATION encouraged to use protocols that provide the most information about all species of birds in To meet research needs, we must prioritize their study areas (e.g., Latta et al. 2003b, Brown building capacity within indigenous conserva- and Sherry 2006b). By broadening the scope of tion groups and among the next generation studies beyond a single focal species, additional of biologists who will address the growing cri- opportunities are created for collaborations, stu- sis in biodiversity protection. Although some dent involvement, and generating results rele- islands such as Puerto Rico and Cuba have vant to resource managers. With an awareness a number of excellent field biologists, many of unique histories and cultural and political other countries have few, if any, trained research traditions among islands, by partnering with biologists to develop research or monitoring local organizations and individuals, and with programs. Although excellent field assistants increased funding, ornithologists and conserva- are available in most countries, managers are tionists working in the Caribbean region will often dependent on outside researchers to design have a greater chance of conserving Caribbean and direct research programs, and to interpret birds and their habitats in a challenging relevant research. Thus, building capacity, espe- environment. cially in terms of Ph.D.-level ornithologists with permanently supported positions, is a critical need in many countries across the region. This ACKNOWLEDGMENTS may be dependent on funding, but ornithologists based outside the region can have an I thank B. Dalsgaard, J. Faaborg, B. Mulvihill, C. Rimmer, T. W. Sherry, L. Sorenson, J. Wunderle, important impact by collaborating with local Jr., and two anonymous reviewers for their insightful organizations including universities, museums, comments on an early version of this manuscript. Their governmental agencies, and nongovernmental kind criticisms were greatly appreciated. 116 S. C. Latta J. Field Ornithol.

LITERATURE CITED sity in Darwin’s finches and their relatives. Evolution 56: 1240–1252. CEVEDO,M.A.,AND C. RESTREPO. 2008. Land-cover and BUTLER, P. J. 1992. Parrots, pressures, people and pride. land-use change and its contribution to the large- In: New World parrots in crisis: solutions from scale organization of Puerto Rico’s bird assemblages. conservation biology (S. R. BEISSINGER AND N. F. R. Diversity and Distributions 14: 114–122. SNYDER, eds.), pp. 25–46. Smithsonian Institution ACOSTA CRUZ,M.,AND L. MUGICA VALDES´ . 1988. Press, Washington, D.C. Estructura de las comunidades de aves que habitan CASE, T. J., J. FAABORG, AND R. SIDELL. 1983. The role los bosques Cubanos. Ciencia Biologica´ ACC 19–20: of body size in the assembly of West Indian bird 9–19. communities. Evolution 37: 1062–1074. ALLEN, R. P. 1956. The flamingos: their life history COLLAZO,J.A.,AND G. I. BONILLA MARTINEZ. 1988. and survival, with special reference to the American Comparacion´ de la riqueza de aves entre plantaciones or West Indian Flamingo (Phoenicopterus ruber). de pino hondureno˜ (Pinus caribea)yareas´ de bosque Research Report No. 5, National Audubon Society, nativo en el Bosque Estatal de Carite, Cayey, Puerto New York, NY. Rico. Caribbean Journal of Science 24: 1–10. ARENDT, W. J. 1985a. Philornis ectoparasitism of Pearly- ———, B. A. HARRINGTON,J.S.GREAR, AND J. A. eyed Thrashers. I. Impact on growth and develop- COLON´ . 1995. Abundance and distribution of shore- ment of nestlings. Auk 102: 270–280. birds at the Cabo Rojo salt flats, Puerto Rico. Journal ———. 1985b. Philornis ectoparasitism of Pearly-eyed of Field Ornithology 66: 424–438. Thrashers. II. Impact on adults and reproduction. ———, F. J. VILELLA,T.H.WHITE,Jr.,AND S. Auk 102: 281–292. GUERRERO. 2000. Survival, use of habitat, and ———. 2006. Adaptations of an avian supertramp: dis- movements of captive reared Hispaniolan Parrots tribution, ecology, and life history of the Pearly-eyed released in historical, occupied habitat: implications Thrasher (Margarops fuscatus). General Technical for the recovery of the Puerto Rican Parrot. North Report 27, U.S. Department of Agriculture, Forest Carolina Cooperative Fish and Wildlife Research Service, International Institute of Tropical Forestry, Unit, Raleigh, NC. San Juan, PR. ———, T. H. WHITE,Jr.,F.J.VILELLA, AND S. A. ASKINS,R.A.,AND D. N. EWERT. 1991. Impact of Hur- GUERRERO. 2003. Survival of captive-reared Hispan- ricane Hugo on bird populations in Virgin Islands iolan Parrots released in Parque Nacional del Este, National Park. Biotropica 23: 481–487. Dominican Republic. Condor 105: 198–207. BEISSINGER, S. R., J. M. WUNDERLE,Jr.,J.M.MEYERS, COX, G. W. 2010. Bird migration and global change. B. E. SAETHER, AND S. ENGEN. 2008. Anatomy of Island Press, Washington, D.C. a bottleneck: diagnosing factors limiting population COYNE,J.A.,AND T. D. P RICE. 2000. Little evidence for growth in the Puerto Rican Parrot. Ecological Mono- sympatric speciation in island birds. Evolution 54: graph 78:185–203. 2166–2171. BIRDLIFE INTERNATIONAL. 2008. Important bird areas in CRUZ, A. 1974. Feeding assemblages of Jamaican birds. the Caribbean: key sites for conservation. BirdLife Condor 76: 103–107. Conservation Series No. 15, BirdLife International, ———. 1987. Avian community organization in a Cambridge, UK. mahogany plantation on a Neotropical island. BRADLEY, P. E. 2000. The birds of the Cayman Islands. Caribbean Journal of Science 23: 286–296. British Ornithologists’ Union, Tring, UK. ———. 1988. Avian resource use in a Caribbean pine ———, AND R. L. NORTON. 2009. An inventory of plantation. Journal of Wildlife Management 52: breeding seabirds of the Caribbean. University Press 274–279. of Florida, Gainesville, FL. ———, AND J. W. WILEY. 1989. The decline of an BROCK,M.K.,AND B. N. WHITE. 1992. Application adaptation in the absence of a presumed selection of DNA fingerprinting to the recovery program of pressure. Evolution 43: 55–62. the endangered Puerto Rican Parrot. Proceedings of ———, T. MANOLIS, AND J. W. WILEY. 1985. The Shiny the National Academy of Sciences USA 89: 11121– Cowbird, a brood parasite expanding its range in 11125. the Caribbean region. In: Neotropical ornithology BROWN,D.R.,AND T. W. S HERRY. 2006a. Food sup- (P.A.BUCKLEY,M.S.FOSTER,E.S.MORTON,R. ply controls the body condition of a migrant bird S. RIDGELY, AND F. G . B UCKLEY, eds.), pp. 607–620. wintering in the tropics. Oecologia 149: 22–32. Ornithological Monograph 36, American Ornithol- ———. 2006b. Population response of resident Jamaican ogists’ Union, Washington, D.C. birds to food supplementation: evidence of dry- ———, J. W. PRATHER,J.W.WILEY AND P. F. W EAVER. season limitation. Biotropica 38: 91–99 2008. Egg rejection behavior in a population exposed ———. 2008. Alternative strategies of space use and to parasitism: Village Weavers on Hispaniola. Behav- response to resource change in a wintering migrant ioral Ecology 19: 398–403. songbird. Behavioral Ecology 19: 1314–1325. DALSGAARD,B.,G.M.HILTON,G.A.L.GRAY,L. BUCKLEY, P. A., E. B. MASSIAH,M.B.HUTT,F. AYMER,J.BOATSWAIN,J.DALEY,C.FENTON,J. G. BUCKLEY, AND H. F. HUTT. 2009. The birds MARTIN,L.MARTIN,P.MURRAIN,W.J.ARENDT, of Barbados. British Ornithologists’ Union, Tring, D. W. GIBBONS, AND J. M. OLESEN. 2007. Im- UK. pacts of a volcanic eruption on the forest bird BURNS, K. J., S. J. HACKETT, AND N. K. KLEIN. 2002. community of Montserrat, Lesser Antilles. Ibis 149: Phylogenetic relationships and morphological diver- 298–312. Vol. 83, No. 2 Avian Research in the Caribbean 117

———, A. M. MARTÍN GONZALEZ´ ,J.M.OLESEN,A. and conservation (A. KEAST AND E. S. MORTON, TIMMERMANN,L.H.ANDERSEN, AND J. OLLERTON. eds.), pp. 157–163. Smithsonian Institution Press, 2008. Pollination networks and functional specializa- Washington, D.C. tion: a test using Lesser Antillean plant-hummingbird ———, W. J. ARENDT, AND K. DUGGER. 2000. The assemblages. Oikos 117: 789–793. Guanica,´ Puerto Rico, bird monitoring project. Bird ———, A. M. MARTÍN GONZALEZ´ ,J.M.OLESEN,J. Populations 5: 102–111. OLLERTON,A.TIMMERMANN,L.H.ANDERSEN, AND ———, K. M. DUGGER, AND W. J. A RENDT. 2007. Long- A. G. TOSSAS. 2009. Plant-hummingbird interac- term variation in the winter resident bird community tions in the West Indies: floral specialization gradients of Guanica´ Forest, Puerto Rico: lessons for measuring associated with environment and hummingbird size. and monitoring species richness. Journal of Field Oecologia 159: 757–766. Ornithology 78: 270–278. DIAMOND, A. W. 1974. Annual cycles in Jamaican forest ———, R. T. HOLMES,A.D.ANDERS,K.L.BILDSTEIN, birds. Journal of Zoology 173: 277–301. K. M. DUGGER,S.A.GAUTHREAUX,Jr.,P.HEGLUND, DIAMOND, J. M. 1977. Continental and insular speciation K. A. HOBSON,A.E.JAHN,D.H.JOHNSON,S.C. in Pacific land birds. Systematic Zoology 26: 263– LATTA,D.J.LEVEY,P.P.MARRA,C.L.MERKORD,E. 268. NOL,S.I.ROTHSTEIN,T.W.SHERRY,T.S.SILLETT, DIGGS,N.E.,P.P.MARRA, AND R. J. COOPER. 2011. Re- F. R . T HOMPSON III, AND N. WARNOCK. 2010a. source limitation drives patterns of habitat occupancy Managing migratory landbirds in the New World: during the nonbreeding season for an omnivorous do we know enough? Ecological Applications 20: songbird. Condor 113: 646–654. 398–418. DUGGER,K.M.,J.FAABORG,W.J.ARENDT, AND K. A. ———, R. T. HOLMES,A.D.ANDERS,K.L.BILDSTEIN, HOBSON. 2004. Understanding survival and abun- K. M. DUGGER,S.A.GAUTHREAUX Jr., P.HEGLUND, dance of overwintering warblers: does rainfall matter? K. A. HOBSON,A.E.JAHN,D.H.JOHNSON,S.C. Condor 106: 744–760. LATTA,D.J.LEVEY,P.P.MARRA,C.L.MERKORD,E. DUPUIS,A.P.,P.P.MARRA, AND L. D. KRAMER. 2003. NOL,S.I.ROTHSTEIN,T.W.SHERRY,T.S.SILLETT, Serologic evidence of West Nile virus transmission, F. R . T HOMPSON III, AND N. WARNOCK. 2010b. Jamaica, West Indies. Emerging Infectious Diseases Recent advances in understanding migration systems 9: 860–863. of New World land birds. Ecological Monographs EMLEN, J. T. 1977. Land bird communities of Grand 80: 3–48. Bahama Island: the structure and dynamics of an FALLON, S. M., R. E. RICKLEFS,S.C.LATTA, AND E. avifauna. Ornithological Monograph 24, American BERMINGHAM. 2004. Temporal stability of insular Ornithologists’ Union, Washington, D.C. avian malarial parasite communities. Proceedings of EWERT,D.N.,J.M.WUNDERLE,Jr.,C.W.EBERLY, the Royal Society B 271: 493–500. P. W. H UBER, AND E. CAREY. 2009. The Kirtland’s ———, E. BERMINGHAM, AND R. E. RICKLEFS. 2005. Warbler: interagency and international cooperation Host specialization and geographic localization of works. University of Michigan School of Natural avian malaria parasites: a regional analysis in the Resources, Endangered Species Update 26: 29–38. Lesser Antilles. American Naturalist 165: 466– FAO (FOOD AND AGRICULTURAL ORGANIZATION). 1991. 480. Forest resources assessment 1990. Food and Agri- GNAM, R. S. 1991. Breeding biology of the Bahama cultural Organization of the United Nations, Rome, Parrot (Amazona leucocephala bahamensis). Ph.D. Italy. dissertation, City University of New York, New York, FAABORG, J. 1980. Further observation on ecological NY. release in Mona Island birds. Auk 97: 624–627. GOCHFELD, M. 1974. Current status of and threats ———. 1982. Trophic and size structure of West In- to some parrots of the Lesser Antilles. Biological dian bird communities. Proceedings of the National Conservation 6: 184–197. Academy of Sciences USA 79: 1563–1567. GREENBERG, R. 1995. Insectivorous migratory birds in ———. 1985. Ecological constraints on West Indian tropical ecosystems: the breeding currency hypothe- bird distributions. In: Neotropical ornithology (P. sis. Journal of Avian Biology 26: 260–264. A. BUCKLEY,M.S.FOSTER,E.S.MORTON,R.S. HAYES,W.K.,R.X.BARRY,Z.MCKENZIE, AND P. B ARRY. RIDGELY, AND F. G . B UCKLEY, eds.), pp. 621–653. 2004. Grand Bahama’s Brown-headed Nuthatch: a Ornithological Monograph 36, American Ornithol- distinct and endangered species. Bahamas Journal of ogists’ Union, Washington, D.C. Science 12: 21–28. ———. 2000. Avian monitoring systems. In: Results of HEARTSILL-SCALLEY,T.,F.N.SCATENA,C.ESTRADA,W. the national planning workshop for avian conserva- H. MCDOWELL, AND A. E. LUGO. 2007. Disturbance tion in the Dominican Republic (S. C. LATTA AND and long-term patterns of rainfall and throughfall R. LORENZO, eds.), pp. 89–98. Direccion´ Nacional nutrient fluxes in a subtropical wet forest in Puerto de Parques, Santo Domingo, Dominican Republic. Rico. Journal of Hydrology 333: 472–485. ———. 2002. Saving migrant birds. University of Texas HILTON,G.M.,P.W.ATKINSON,G.A.L.GRAY,W.J. Press, Austin, TX. ARENDT, AND D. W. GIBBONS. 2003. Rapid decline ———, AND W. J. A RENDT. 1989. Long-term declines in of the volcanically threatened Montserrat Oriole. winter resident warblers in a Puerto Rican dry forest. Biological Conservation 111: 79–89. American Birds 43: 1226–1230. HOLMES, R. T. 2007. Understanding population change ———, AND J. W. TERBORGH. 1980. Patterns of in migratory songbirds: long-term and experimental migration in the West Indies. In: Migrant birds studies of Neotropical migrants in breeding and in the Neotropics: ecology, behavior, distribution wintering areas. Ibis 149 (Suppl. 2): 2–13. 118 S. C. Latta J. Field Ornithol.

———. 2011. Avian population and community pro- Club 16, Nuttall Ornithological Club, Cambridge, cesses in forest ecosystems: long-term research in the MA. Hubbard Brook Experimental Forest. Forest Ecology KEPLER,C.B.,AND A. K. KEPLER. 1973. The distri- and Management 262: 20–32. bution and ecology of the Puerto Rican Whip- ———, T. W. SHERRY, AND L. REITSMA. 1989. Popula- poor-will, an endangered species. Living Bird 11: tion structure, territoriality and overwinter survival 207–239. of two migrant warbler species in Jamaica. Condor KERCHNER, C., M. HONZAK´ ,R.KEMKES,A. 91: 545–561. RICHARDSON,J.TOWNSEND, AND C. C. RIMMER. ———, P. P. MARRA, AND T. W. S HERRY. 1996. Habitat- 2009. Designing spatially explicit incentive programs specific demography of breeding Black-throated Blue for habitat conservation: a case study of the Bicknell’s Warblers (Dendroica caerulescens): implications for Thrush wintering grounds. Ecological Economics population dynamics. Journal of Animal Ecology 65: 69: 2108–2115. 183–195. KISSLING, W. D., C. F. DORMANN,J.GROENEVELD,T. JOHNSON, M. D. 2000. Effects of shade-tree species and HICKLER,I.KHN,G.J.MCINERNY,J.M.MONTOYA, crop structure on the winter arthropod and bird C. ROMERMANN¨ ,K.SCHIFFERS,F.M.SCHURR,A. communities in a Jamaican shade coffee plantation. SINGER,J.C.SVENNING,N.E.ZIMMERMANN, AND Biotropica 32: 133–145. R. B. O’HARA. 2012. Towards novel approaches to ———, AND T. W. S HERRY. 2001. Effects of food avail- modeling biotic interactions in multispecies assem- ability on the distribution of migratory warblers blages at large spatial extents. Journal of Biogeogra- among habitats in Jamaica. Journal of Animal Ecol- phy, in press. ogy 70: 546–560. KOENIG,S.E.,J.M.WUNDERLE,Jr.,AND E. C. ENKERLIN- ———, ———, A. M. STRONG, AND A. MEDORI. HOEFLICH. 2007. Vines and canopy contact: a route 2005. Migrants in Neotropical bird communities: for predation on parrot nests. Bird Conservation an assessment of the breeding currency hypothesis. International 17: 79–91. Journal of Animal Ecology 74: 333–341. KOMAR, O., M. B. ROBBINS,K.KLENK,B.J.BLITVICH, ———, ———, R. T. HOLMES, AND P. P. M ARRA. 2006. N. L. MARLENEE,K.L.BURKHALTER,D.J.GUBLER, Assessing habitat quality for a migratory songbird G. GONZALVEZ´ ,C.J.PENA˜ ,A.T.PETERSON, AND wintering in natural and agricultural habitats. Con- N. KOMAR. 2003. West Nile Virus transmission servation Biology 20: 1433–1444. in resident birds, Dominican Republic. Emerging ———, N. J. LEVY,J.L.KELLERMANN, AND D. E. Infectious Diseases 9: 1299–1302. ROBINSON. 2009. Effects of shade and bird predation LACK, D. 1976. Island biology as illustrated by the land on arthropods and leaf damage on coffee farms in the birds of Jamaica. Blackwell Scientific Publications, Blue Mountains. Agroforestry Systems 76: 139–148. Oxford, UK. ———, J. L. KELLERMANN, AND A. M. STERCHO. 2010. LATTA, S. C. 2003. Effects of scaley-leg mite infestations Pest reduction services by birds in shade and sun on body condition and site fidelity of migratory coffee in Jamaica. Animal Conservation 13: 140– warblers. Auk 120: 730–743. 147. ———, AND M. BALTZ. 1997. Population limitation in KEAST,A.,AND E. S. MORTON (eds.). 1980. Migrant Neotropical migratory birds: comments on Rappole birds in the Neotropics: ecology, behavior, distribu- and McDonald (1994). Auk 114: 754–762. tion and conservation. Smithsonian Institution Press, ———, AND C. BROWN. 1999. Autumn stopover ecology Washington, D.C. of the Blackpoll Warbler (Dendroica striata)inthorn KEITH, A. R. 1997. The birds of St. Lucia. British scrub forest of the Dominican Republic. Canadian Ornithologists’ Union, Tring, UK. Journal of Zoology 77: 1147–1156. ———, J. W. WILEY,S.C.LATTA, AND J. A. ———, AND J. FAABORG. 2001. Winter site fidelity of OTTENWALDER. 2003. The birds of Hispaniola: Haiti Prairie Warblers in the Dominican Republic. Condor and the Dominican Republic. British Ornithologists’ 103: 455–468. Union, Tring, UK. ———, AND ———. 2002. Demographic and popula- KELLERMANN,J.L.,M.D.JOHNSON,A.M.STERCHO, tion responses of Cape May Warblers wintering in AND S. HACKETT. 2008. Ecological and economic multiple habitats. Ecology 83: 2502–2515. services provided by birds on Jamaican Blue Moun- ———, AND ———. 2009. Migratory birds in the tain coffee farms. Conservation Biology 22: 1177– Caribbean: benefits of studies of overwintering birds 1185. for understanding resident bird ecology and promot- KENNEDY, C . M . , P. P. M ARRA,W.F.FAGAN, AND M. ing critical development of conservation capacity. C. NEEL. 2010. Landscape matrix and species traits Conservation Biology 23: 286–293. mediate responses of Neotropical resident birds to ———, AND B. M. O’CONNOR. 2001. Patterns of forest fragmentation in Jamaica. Ecological Mono- Knemidokoptes jamaicensis (Acari: Knemidokoptidae) graphs 80: 651–669. infestations among eight new avian hosts in the Do- ———, E. H. CAMPBELL GRANT,M.C.NEEL,W.F. minican Republic. Journal of Medical Entomology FAGAN, AND P. P. M ARRA. 2011. Landscape matrix 38: 437–440. mediates occupancy dynamics of Neotropical avian ———, AND R. E. RICKLEFS. 2010. Prevalence patterns insectivores. Ecological Applications 21: 1837–1850. of avian haemosporidia on Hispaniola. Journal of KEPLER, A. K. 1977. Comparative study of todies (Todi- Avian Biology 41: 25–33. dae): with emphasis on the Puerto Rican Tody, Todus ———, AND J. M. WUNDERLE, Jr. 1998. The assemblage mexicanus. Publications of the Nuttall Ornithological of birds foraging in native West Indian pine (Pinus Vol. 83, No. 2 Avian Research in the Caribbean 119

occidentalis) forests of the Dominican Republic dur- 2006. Aves acuaticas´ en los humedales de Cuba. ing the nonbreeding season. Biotropica 30: 645–656. Editorial Cient´ıfico-Tecnica,´ La Habana, Cuba. ———, M. L. SONDREAL, AND C. R. BROWN. 2000. A MYERS, N. 1988. Threatened biotas: hotspots in tropical hierarchical analysis of nesting and foraging habitat forests. Environmentalist 8: 178–208. for the conservation of the Hispaniolan White- NEELIN,J.D.,M.MUNNICH,H.SU,J.E.MEYERSON, winged Crossbill (Loxia leucoptera megaplaga). Bio- AND C. E. HOLLOWAY. 2006. Tropical drying trends logical Conservation 96: 139–150. in global warming models and observations. Proceed- ———, A. TOSSAS,A.SUTTON,H.GONZALEZ,P.B. ings of the National Academy of Sciences USA 103: HAMEL, AND D. DESANTE. 2003a. Research, mon- 6110–6115. itoring, and conservation of Neotropical migratory NORRIS,D.R.,P.P.MARRA,T.K.KYSER,T.W.SHERRY, land birds in the West Indies: a report to the AND L. M. RATCLIFFE. 2004. Tropical winter habitat Society for the Conservation and Study of Caribbean limits reproduction success in a migratory bird. Birds (SCSCB) by the Neotropical Migratory Bird Proceedings of the Royal Society B 271: 59–64. Working Group of the SCSCB. Journal of Caribbean PARRISH,J.D.,AND T. W. S HERRY. 1994. Sexual habi- Ornithology 16: 1–19. tat segregation by American Redstarts wintering in ———, C. C. RIMMER, AND K. P. MCFARLAND. 2003b. Jamaica: importance of resource seasonality. Auk 111: Winter bird communities in four habitats along an 38–49. elevational gradient on Hispaniola. Condor 105: PAULINO, M. M., D. A. MEJÍA, AND S. C. LATTA. 2010. A 179–197. new review of the status of the Caribbean Flamingo ———, C. J. RALPH, AND G. GEUPEL. 2005. Strate- (Phoenicopterus ruber) in the Dominican Republic gies for the conservation monitoring of permanent and Haiti. Bulletin of the IUCN-SSC/Wetlands resident landbirds and wintering Neotropical mi- International Flamingo Specialist Group; Flamingo grants in the Americas. Ornitolog´ıa Neotropical 16: 18: 62–66. 163–174. POST, W. 1981. Biology of the Yellow-shouldered Black- LOSOS,J.B.,AND R. E. RICKLEFS. 2009. Adaptation and bird – Agelaius on a tropical island. Bulletin of the diversification on islands. Nature 457: 830–836. Florida State Museum Biological Science 26: 125– MACARTHUR,R.H.,AND E. O. WILSON. 1967. The 202. theory of island biogeography. Princeton University ———, AND J. W. WILEY. 1977. Reproductive in- Press, Princeton, NJ. teractions of the Shiny Cowbird and the Yellow- ———, H. RECHER, AND M. CODY. 1966. On the rela- shouldered Blackbird. Condor 79: 176–184. tion between habitat selection and species diversity. ———, A. CRUZ, AND D. B. MCNAIR. 1993. The North American Naturalist 100: 319–332. American invasion pattern of the Shiny Cowbird. MARRA, P. P. 2000. The role of behavioral dominance Journal of Field Ornithology 64: 32–41. in structuring patterns of habitat occupancy in a REUDINK, M . W. , P. P. M ARRA,T.K.KYSER,P.T.BOAG, migrant bird during the non-breeding period. Be- K. M. LANGIN, AND L. M. RATCLIFFE. 2009. Non- havioral Ecology 11: 299–308. breeding season events influence sexual selection in ———, AND R. L. HOLBERTON. 1998. Corticosterone a long-distance migratory bird. Proceedings of the levels as indicators of habitat quality: effects of habitat Royal Society B 276: 1619–1626. segregation in a migratory bird during the non- RICKLEFS, R. E. 1992. The megapopulation: a model of breeding season. Oecologia 116: 284–292. demographic coupling between migrant and resident ———, AND R. T. HOLMES. 2001. Consequences of landbird populations. In: Ecology and conservation dominance-mediated habitat segregation in a mi- of Neotropical migrant landbirds (J. M. HAGAN grant passerine bird during the non-breeding season. III AND D. W. JOHNSTON, eds.), pp. 537–548. Auk 118: 92–104. Smithsonian Institution Press, Washington, D.C. ———, K. A. HOBSON, AND R. T. HOLMES. 1998. ———. 2011. A biogeographical perspective on ecolog- Linking winter and summer events in a migratory ical systems: some personal reflections. Journal of bird by using stable-carbon isotopes. Science 282: Biogeography 38: 2045–2056. 1884–1886. ———, AND E. BERMINGHAM. 2001. Nonequilibrium MARTIN,H.C.,AND P. S . W EECH. 2001. Climate change diversity dynamics of the Lesser Antillean avifauna. in the Bahamas? Evidence in the meteorological Science 294: 1522–1524. records. Bahamas Journal of Science 5: 22–32. ———, AND ———. 2002. The concept of the taxon MCNAIR, D. B., E. B. MASSIAH, AND M. D. FROST. cycle in biogeography. Global Ecology and Biogeog- 2002. Ground-based autumn migration of Blackpoll raphy 11: 353–361. Warblers at Harrison Point, Barbados. Caribbean ———, AND ———. 2004. History and the species- Journal of Science 38: 239–248. area relationship in Lesser Antillean birds. American MITTERMEIER,R.A.,N.MYERS,P.GIL ROBLES, AND C. G. Naturalist 163: 227–239. MITTERMEIER. 1999. Hotspots: Earth’s biologically ———, AND ———. 2007. Evolutionary radiations of richest and most threatened ecosystems. Cemex, passerine birds in archipelagoes. American Naturalist Mexico, D.F. 169: 285–297. MØLLER,A.P.,W.FIELDER, AND P. B ERTHOLD (eds.). ———, AND ———. 2008. The West Indies as a labo- 2010. Effects of climate change on birds. Oxford ratory of biogeography and evolution. Philosophical University Press, New York, NY. Transactions of the Royal Society B 363: 2393–2413. MUGICA VALDES´ ,L.,D.DENIS A´ VILA,M.ACOSTA CRUZ, ———, AND G. W. COX. 1972. Taxon cycles in the West A. JIMENEZ´ REYES, AND A. RODRÍGUEZ SUAREZ´ . Indian avifauna. American Naturalist 106: 195–219. 120 S. C. Latta J. Field Ornithol.

———, AND S. M. FALLON. 2002. Diversification and SLY,N.D.,A.K.TOWNSEND,C.C.RIMMER,J.M. host switching in avian malaria parasites. Proceedings TOWNSEND,S.C.LATTA, AND I. J. LOVETTE. 2010. of the Royal Society B 269: 885–892. Phylogeography and conservation of the endemic ———, ———, AND E. BERMINGHAM. 2004. Evolution- Hispaniolan Palm-Tanagers (Aves: Phaenicophilus). ary relationships, cospeciation, and host switching in Conservation Genetics 11: 2121–2129. avian malaria parasites. Systematic Biology 53: 111– ———, ———, ———, ———, ———, AND 119. ———. 2011. Ancient islands and modern in- RIMMER, C. C., K. P. MCFARLAND,P.L.JOHNSON, AND vasions: disparate phylogeographic histories among J. M. TOWNSEND. 2011. Changes in overwintering Hispaniola’s endemic birds. Molecular Ecology 20: Bicknell’s Thrush population in Sierra de Bahoruco, 5012–5024. Dominican Republic. Vermont Center for Ecostud- SMITH,J.A.,L.R.REITSMA, AND P. P. M ARRA. 2010. ies, Norwich, VT. Moisture as a determinant of habitat quality for a RIVERA-MILAN´ , F. F. 1992. Distribution and relative non-breeding Neotropical migratory songbird. Ecol- abundance patterns of columbids in Puerto Rico. ogy 91: 2874–2882. Condor 94: 224–238. SNYDER,N.F.R.,J.W.WILEY, AND C. B. KEPLER. ———. 1995a. Spatial and temporal variation in the 1987. The parrots of Luquillo: natural history and detectability and density of columbids in Puerto Rico conservation of the Puerto Rican Parrot. Western and on Vieques Island. Ornitologia Neotropical 6: Foundation of Vertebrate Zoology, Los Angeles, CA. 1–17. STATTERSFIELD, A. J., M. J. CROSBY,A.J.LONG, AND D. ———. 1995b. Detectability and population density C. WEGE. 1998. Endemic bird areas of the world: of Scaley-naped Pigeons before and after Hurricane priorities for biodiversity conservation. Birdlife Con- Hugo in Puerto Rico and Vieques Island. Wilson servation Series No. 7, BirdLife International, Bulletin 107: 727–733. Cambridge, UK. ———. 1997. Seasonal and annual changes in the popu- STAUS, N. L. 1998. Habitat use and home range of lation density of Zenaida Doves in the xerophytic West Indian Whistling-Ducks. Journal of Wildlife forest of Guanica,´ Puerto Rico. Journal of Field Management 62: 171–178. Ornithology 68: 259–272. STRONG,A.M.,AND T. W. S HERRY. 2000. Habitat RODRÍGUEZ,D.,AND B. SANCHEZ´ . 1995. Avifauna del specific effects of food abundance on the condition of matorral xeromorfo en la region´ oriental de Cuba Ovenbirds wintering in Jamaica. Journal of Animal durante la migracion´ otonal˜ (Octubre de 1989, 1990, Ecology 69: 883–895. 1991). Poeyana 447: 1–12. STUDDS,C.E.,AND P. P. M ARRA. 2005. Nonbreeding ———, ———, A. TORRES, AND A. RAMS. 1994. Com- habitat occupancy and population processes: an up- posicion´ y abundancia de las aves durante la mi- grade experiment with a migratory bird. Ecology 86: gracion´ otonal˜ en Gibara, Cuba. Avicennia 1: 101– 2380–2385. 109. ———, AND ———. 2007. Linking fluctuations in RUNGE,M.C.,AND P. P. M ARRA. 2005. Modeling seasonal rainfall to nonbreeding season performance in a long- interactions in the population dynamics of migratory distance migratory bird, Setophaga ruticilla. Climate birds. In: Birds of two worlds: the ecology and evolu- Research 35: 115–122. tion of migration (R. GREENBERG AND P. P. M ARRA, ———, AND ———. 2011. Rainfall-induced changes eds.), pp. 375–389. Johns Hopkins University Press, in food availability modify the spring departure Baltimore, MD. programme of a migratory bird. Proceedings of the SHERRY,T.W.,AND R. T. HOLMES. 1995. Summer versus Royal Society B 278: 3437–3443. winter limitation of populations: what are the issues ———, T. K. KYSER, AND P. P. M ARRA. 2008. Natal and what is the evidence? In: Ecology and manage- dispersal driven by environmental conditions inter- ment of Neotropical migratory birds: a synthesis and acting across the annual cycle of a migratory songbird. review of critical issues (T. E. MARTIN AND D. M. Proceedings of the National Academy of Sciences FINCH, eds.), pp. 85–120. Oxford University Press, USA 105: 2929–2933. New York, NY. SUTTON,A.H.,L.G.SORENSON, AND M. A. KEELEY. ———, AND ———. 1996. Winter habitat quality, pop- 2004. Wondrous West Indian wetlands: teachers’ re- ulation limitation, and conservation of Neotropical- source book. West Indian Whistling-Duck Working Nearctic migrant birds. Ecology 77: 36–48. Group of the Society for the Conservation and Study ———, M. D. JOHNSON, AND A. M. STRONG. 2005. of Caribbean Birds, Boston, MA. Does winter food limit populations of migratory TERBORGH, J., AND J. FAABORG. 1973. Turnover and birds? In: Birds of two worlds: the ecology and evolu- ecological release in the avifauna of Mona Island, tion of migration (R. GREENBERG AND P. P. M ARRA, Puerto Rico. Auk 90: 759–779. eds.), pp. 414–424. Johns Hopkins University Press, ———, ———, AND H. J. BROCKMAN. 1978. Island Baltimore, MD. colonization by Lesser Antillean birds. Auk 95: 59– SHORT, L. L. 1974. Habits of three endemic West Indian 72. woodpeckers (Aves, Picidae). American Museum of TOSSAS, A. 2002. Source and sink dynamics of Puerto Natural History Novitates 2549, New York, NY. Rican Vireo in a regional scale. Ph.D. dissertation, SILLETT,T.S.,R.T.HOLMES, AND T. W. S HERRY. 2000. University of Puerto Rico, Rio Piedras, PR. Impacts of a global climate cycle on the population TOWNSEND, A. K., C. C. RIMMER,S.C.LATTA, AND dynamics of a migratory songbird. Science 288: I. J. LOVETTE. 2007. Phylogeographic concordance 2040–2042. of nuclear and mitochondrial gene genealogies in Vol. 83, No. 2 Avian Research in the Caribbean 121

the single-island avian radiation of Hispaniolan ———, AND ——— . 1979b. The biology of the White- Chat-tanagers (Aves: Calyptophilus). Molecular Ecol- crowned Pigeon. Wildlife Monograph No. 64, The ogy 16: 3634–3642. Wildlife Society, Washington, D.C. TOWNSEND, J. M., C. C. RIMMER, AND K. P.MCFARLAND. ———, AND ——— . 1981. Breeding season ecology 2010. Winter territoriality and spatial behavior of and behavior of Ridgway’s Hawk (Buteo ridgwayi). Bicknell’s Thrush (Catharus bicknelli) at two ecolog- Condor 83: 132–151. ically distinct sites in the Dominican Republic. Auk ———, AND J. M. WUNDERLE, Jr. 1993. The effects 127: 514–522. of hurricanes on birds, with special reference to VILELLA, F. J. 2008. Nest habitat use of the Puerto Caribbean islands. Bird Conservation International Rican Nightjar Caprimulgus noctitherus in Guanica´ 3: 319–349. Biosphere Reserve. Bird Conservation International WILSON, E. O. 1961. The nature of the taxon cycle in 18: 307–317. the Melanesian ant fauna. American Naturalist 95: ———, AND P. J . Z WANK. 1993. Geographic distribution 169–193. and abundance of the Puerto Rican Nightjar. Journal WILSON, S., S. L. LADEAU,A.P.TØTTRUP, AND P. P. of Field Ornithology 64: 223–238. MARRA. 2011. Range-wide effects of breeding and World Commission on Protected Areas (WCPA) non-breeding season climate on the abundance of a Caribbean. 2003. Insular Caribbean WCPA report Neotropical migrant songbird. Ecology 92: 1789– to the World Parks Congress. World Commission on 1798. Protected Areas, Durban, South Africa. WOODWORTH, B. L. 1997. Brood parasitism, nest WAIDE, R. B. 1984. The avifaunal component of a predation, and season-long reproductive success simplified tropical food web. In: Tropical rain forest: of a tropical island endemic. Condor 99: 605– ecology and management, vol. A (A. C. CHADWICK 621. AND S. L. SUTTON, eds.), p. 324. Leeds Philosophical ———, J. FAABORG, AND W. J. A RENDT. 1998. Breeding and Literary Society, Leeds, UK. and natal dispersal in the Puerto Rican Vireo. Journal ———. 1991. The effect of Hurricane Hugo on bird of Field Ornithology 69: 1–7. populations in the Luquillo Experimental Forest. WUNDERLE, J. M., Jr. 1995a. Population characteristics Biotropica 23: 475–480. of Black-throated Blue Warblers wintering in three WEAVER,P.L.,AND K. A. GONZALEZ. 2005, Wild-land sites on Puerto Rico. Auk 112: 931–946. fire management and restoration. In: Twelfth meeting ———. 1995b. Response of bird populations in a Puerto of Caribbean foresters (P. L. WEAVER AND K. A. Rican forest to Hurricane Hugo: the first 18 months. GONZALEZ´ , eds.), p. 85. United States Department Condor 97: 879–896. of Agriculture, Forest Service, International Institute ———. 1999. Pre- and post-hurricane fruit availability: of Tropical Forestry, R´ıo Piedras, PR. implications for Puerto Rican Parrots in the Luquillo WHITE, T. H., Jr., AND F. J . V ILELLA. 2004. Nest Mountains. Caribbean Journal of Science 35: 249– management for the Puerto Rican Parrot (Amazona 264. vittata): gaining the technological edge. Ornitologia ———. 2005. Hurricanes and the fate of Caribbean birds Neotropical 15 (Suppl.): 467–476. – What do we know, what do we need to know, who ———, J. A. COLLAZO, AND F. J . V ILELLA. 2005. Survival is vulnerable, how can we prepare, what can we do, of captive-reared Puerto Rican Parrots released in the and what are the management solutions? Journal of Caribbean National Forest. Condor 107: 424–432. Caribbean Ornithology 18: 94–96. ———, A. J. CAMACHO,T.BLOOM,P.L.DIEGUEZ´ , AND ———, D. CURRIE,E.H.HELMER,D.N.EWERT,J.D. R. SELLARES. 2011. Human perceptions regarding WHITE,T.S.RUZYCKI,B.PARRESOL, AND C. KWIT. endangered species conservation: a case study of 2010. Kirtland’s Warblers in anthropogenically dis- Saona Island, Dominican Republic. Latin American turbed early-successional habitats on Eleuthera, The Journal of Conservation 2: 18–29. Bahamas. Condor 112: 123–137. WILEY, J. W.1985. Shiny Cowbird parasitism in two avian ———, AND S. C. LATTA. 1996. Avian abundance in sun communities in Puerto Rico. Condor 87: 165–176. and shade coffee plantations and remnant pine for- ———. 1988. Host selection in the Shiny Cowbird. est in the Cordillera Central, Dominican Republic. Condor 90: 289–303. Ornitologia Neotropical 7: 19–34. ———. 1991. Status and conservation of parrots and ———, AND ———. 1998. Avian resource use in Do- parakeets in the Greater Antilles, Bahama Islands, and minican shade coffee plantations. Wilson Bulletin Cayman Islands. Bird Conservation International 1: 110: 255–265. 187–214. ———, AND ———. 2000. Winter site fidelity of Nearc- ———. 2006. The ecology, behavior, and conservation tic migrant birds in isolated shade coffee plantations of a West Indian corvid, the White-necked Crow of different sizes in the Dominican Republic. Auk (Corvus leucognaphalus). Ornitologia Neotropical 17: 117: 596–614. 105–146. ———, D. J. LODGE, AND R. B. WAIDE. 1992. ———, W. POST, AND A. CRUZ. 1991. Conservation of Short-term effects of Hurricane Gilbert on ter- the Yellow-shouldered Blackbird Agelaius xanthomus, restrial bird populations on Jamaica. Auk 109: an endangered West Indian species. Biological Con- 148–166. servation 55: 119–138. ———, R. B. WAIDE, AND J. FERNANDEZ. 1989. Seasonal ———, AND B. N. WILEY. 1979a. Status of the American abundance of shorebirds in the Jobos Bay Estuary in Flamingo in the Dominican Republic and eastern southern Puerto Rico. Journal of Field Ornithology Haiti. Auk 96: 615–619. 60: 329–339.