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Redalyc.The Role of Historical and Local Factors in Determining Species Revista de Biología Tropical ISSN: 0034-7744 [email protected] Universidad de Costa Rica Costa Rica Barrantes, Gilbert The role of historical and local factors in determining species composition of the highland avifauna of Costa Rica and Western Panamá Revista de Biología Tropical, vol. 57, núm. 1, noviembre, 2009, pp. 333-349 Universidad de Costa Rica San Pedro de Montes de Oca, Costa Rica Available in: http://www.redalyc.org/articulo.oa?id=44918950029 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative The role of historical and local factors in determining species composition of the highland avifauna of Costa Rica and Western Panamá Gilbert Barrantes Escuela de biología, Universidad de Costa Rica, 11501-2060, San José, Costa Rica; [email protected] Received 30-VI-2007. Corrected 09-X-2008. Accepted 18-XI-2008. Abstract: The formation of the mountain ranges of Costa Rica and western Panamá, as well as the cold climatic conditions that prevailed during the upper Pleistocene, played a crucial role in determining the bird species composition of the highlands in this region. Glacial conditions favored dispersal movements of bird species from the Andes, and from the Neartic region. Subsequent inter-glacial conditions reduced the connectivity between neotropical highlands (e.g., Talamanca-Andes), and between neotropical highlands and Neartic temper- ate region, isolating recently established populations from the ancestral populations, and promoting speciation. Within Costa Rica, the highland vegetation and the birds that occupied this vegetation possibly had a continuous distribution throughout all mountain ranges during glacial periods. This vegetation retreated to the summit of the mountains during inter-glacial periods, fragmenting the original continuous bird populations and forming “sky islands”, which decrease in size from Talamanca mountains towards the northwestern ranges. The sizes of such islands of available habitat determine the number of highland birds present in each mountain range. Rev. Biol. Trop. 57 (Suppl. 1): 333-349. Epub 2009 November 30. Key words: highland birds, biogeography, climate, endemism, Diglossa, Catharus, Piranga, Costa Rica. Species composition in bird communities modified by extinction, a continuos process is influenced by both, historical and ecological that may be compensated by colonization and factors (Vuilleumier & Simberloff 1980, Rick- speciation (Mayr 1963, Wiens 1989). Coloni- lefs 1987, Wiens 1989). The role of history in zation of a given location depends on species- structuring communities has been frequently specific dispersal ability and certain historical acknowledged, although few studies have spe- events (e.g., climatic change, formation of cifically addressed its importance (Ricklefs land bridges), that reduce or eliminate habitat & Cox 1972, Karr 1976a, Ricklefs 1978, discontinuities (Haffer 1970, 1974, 1987, Dia- Vuilleumier & Simberloff 1980, Lovette et mond 1984). Following colonization, specia- al. 1998). More frequently, patterns of spe- tion may later occur if isolation prevents gene cies occurrence have been attributed to past flow between newly established and ancestral and contemporary interspecific interactions, populations (Grant 1986, Hackett 1995). How- such as competition and predation (MacArthur ever, the importance of speciation in determin- 1958, Diamond 1975, Cody 1978, Terborgh ing the species pool of a bird community has 1985, Martin 1992). received little attention by community ecolo- Regional species composition is likely the gists. On the contrary, interspecific interactions result of several factors acting in a more or less have frequently been considered to play a pri- sequential order, e.g., colonization, speciation, mary role in determining species membership and interspecific interactions (Wiens 1989). in a bird assemblage (Lack 1947, Diamond Species composition in a community is further 1975, Pulliam 1983). For example, competitive Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 57 (Suppl. 1): 333-349, November 2009 333 interactions have been proposed as a fac- were described by Hooghiemstra et al. (1992), tor shaping bird communities through niche and Islebe et al. (1995, 1996). Using informa- partitioning (Cody 1974, 1981). It is unlikely, tion from Haffer (1974), Hooghiemstra et al. however, that any single factor fully explains (1992), and Islebe et al. (1995, 1996) on vegeta- species composition in a bird community, and tion patterns, I examined the role of vegetation examination of present day communities is not changes on the current distribution of highland complete without considering the role of his- birds in Costa Rica. Using a digital elevation tory (Schluter & Ricklefs 1993). model (DEM) available at 30 arc-second scale Here, I examine how historical and eco- for Costa Rica from USGS EROS web site logical events have potentially interacted to and Geographic Information System software determine species composition of the highland (Arc/INFO version 7.2, ArcView version 3.1, avifauna of Costa Rica and Western Panamá. ESRITM, University of Missouri-St. Louis), I This avifauna has a complex biogeographic estimated the likely expansion and contraction origin and possesses the highest number of of highland vegetation that occurred during the endemic species in Central America (Sánchez late Pleistocene and beginning of Holocene. et al. 2004, Barrantes 2005). Highland bird assemblages in this region show discontinuous Dispersal events: I used information on distributions due to the present day topography fossil record (Olson 1985, Feduccia 1999), phy- of highland sites and past historical events. I logenetic relationships (Hackett 1995, Burns addressed two main questions in this study: (1) 1998, Outlaw et al. 2003, Winker & Pruett How did geological and vegetational changes 2006), and historical events (e.g., geological, associated with Quaternary climatic fluctua- vegetation, and climatic events) to reconstruct tions influenced the bird species composition past dispersal events on the highlands of south- in this region?; (2) How did local factors, ern Central America (Haffer 1974, Hooghi- specifically the area of highland forests (e.g., emstra et al. 1992, Islebe et al. 1995, 1996). upper montane and subalpine forests), affected Phylogenetic information on some extant taxa, highland species composition and distribution including species present in Costa Rica and among Costa Rican mountain ranges? Panamá highlands, was used to infer the ori- gin and dispersal routes of some bird lineages MATERIALS AND METHODS (Hackett 1995, Burns 1998, Outlaw et al. 2003, Winker & Pruett 2006). Although information Historical data: I used published informa- on fossil record and pylogenetic relationships tion on geology, palaeoclimate, fossil record, is incomplete, it still provides important infor- and evolution of some groups of birds, to mation on the origin and expansion of ancient reconstruct possible dispersal and coloniza- lineages. tion events. These events are hypothesized to have had a major impact in determining spe- Highland avifauna and current distri- cies composition of the highland avifauna of bution: I relied on Stiles & Skutch’s (1989) Costa Rica and Western Panamá. I obtained information to determine the distribution of geological information from Haffer (1974), highland bird species in the main mountain Castillo (1984), Gómez (1986), and Coates & ranges of Costa Rica (Talamanca, Central Obando (1996). Information on palaeoclimate Volcanic, Tilarán, and Guanacaste; Fig. 1). and vegetation changes during the upper Pleis- Highland species were defined by Wolf (1976), tocene were described by Haffer (1974, 1987), as those birds that occur above 2 600 m Gómez (1986), Colinvaux (1996), and Webb & in the Talamanca mountain range (Appendix Rancy (1996). Changes in climate and expan- 1.1). The elevation of Costa Rican mountains sion and contraction of highland forests on decreases to the northwest (e.g, Tilarán and Talamanca mountains during late Quaternary Guanacaste mountain ranges), where highest 334 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 57 (Suppl. 1): 333-349, November 2009 Nicaragua Guanacaste Tilarán Caribbean sea Central 10º N Talamanca Pacic Ocean Elevation 0-1000 m 1000-2000 m Panama 2000-3000 m 3000-3688 m 84º W Fig. 1. Costa Rica; main mountain ranges: Talamanca, Central Volcanic, Tilarán, and Guanacaste. peaks lie between 1 500 and 2 000 m. How- among mountain ranges on species richness of ever, climatic conditions have permitted the highland birds. establishment of small fragments of the same highland vegetation present in the Talamancas RESULTS at lower elevations on these more northerly mountains. Thus, highland habitats for birds Geological events are present in all Costa Rican mountain ranges, although extension of these habitats decreases Early archipelagos: The recent formation to the northwest. of the Costa Rica-Panamá isthmus was the culmination of a long and intensive interaction I described the effect of area of habitat of tectonic plates and volcanism (Haffer 1974, available on species composition and distribu- Castillo 1984, Gómez 1986, Coates & Obando tion of highland, for non-endemic and endemic 1996). The formation of the isthmus likely ini-
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