Kumar, A. & S. O'donnell. 2007. Fragmentation and Elevation Effects
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Journal of Tropical Ecology (2007) 23:581–590. Copyright © 2007 Cambridge University Press doi:10.1017/S0266467407004270 Printed in the United Kingdom Fragmentation and elevation effects on bird–army ant interactions in neotropical montane forest of Costa Rica Anjali Kumar1 and Sean O’Donnell Animal Behavior Program, Department of Psychology, Box 351525, University of Washington, Seattle, WA 98195, USA (Accepted 27 May 2007) Abstract: Army ants (Formicidae: Ecitoninae) are top predators in neotropical forests. Army ant raids support a community of diverse organisms, including birds that attend the raids to collect prey. While elevation and forest fragmentation influence army ant and insectivorous bird communities, their effects on the interaction between army ants and bird species is unknown. We studied the size and species composition of bird flocks attending army ant swarms in forest fragments and continuous forest across an elevational gradient (1100–1680 m asl) in a neotropical montane region (Monteverde, Costa Rica). We observed a total of 41 bird species attending army ant swarms. Neither the number of birds, nor the total body mass of birds, nor the number of bird species in attending bird flocks was related to elevation. However, we found a higher bird species richness, larger flock size and greater total body mass of birds attending army ant swarms in continuous forest. Continuous and fragmented forest shared many attending bird species in common, but there was elevational segregation of attending bird species. Some montane endemic birds, and neotropical migrants, attend swarms regularly and use army ant raids as a food source. Key Words: antbird, cloud forest, Eciton burchellii, habitat alteration, Labidus praedator, Monteverde INTRODUCTION raids also show regularity in their spacing among days (Boswell et al. 1998, Franks & Fletcher 1983, Gotwald Army ants (Formicidae: Ecitoninae) are top predators in 1995, Schneirla 1934, 1971). Due to the predictable and neotropical forests (Brady 2003, Franks 1982, Franks & conspicuous swarming behaviour of E. burchellii,some Bossert1983).Recentworksuggeststhatarmyantsoccur neotropical lowland bird species (especially in the families at much higher densities than previously recognized, and Thamnophilidae and Dendrocolaptidae) regularly attend some species of army ant function as ecological keystones swarms in search of prey, and show behavioural in neotropical forests (Boswell et al. 1998, Gotwald 1995, adaptations to foraging at army ant raids (Roberts et al. Kaspari & O’Donnell 2003). Swarms of surface-raiding 2000a,Swartz2001,Willis&Oniki1978,Willson2004). species, such as Eciton burchellii Westwood, 1842, and La- It has been suggested that the abundance of army bidus praedator F. Smith, 1858, host a suite of diverse sym- ant colonies diminishes with increasing elevation, biotic organisms, including mites, beetles and centipedes contributingtotheabsenceofobligatearmyant-following (Gotwald 1995, Rettenmeyer et al. 1983), and swarm bird species in montane forests (Blake & Loiselle 2000, raids are often attended by a diverse array of bird species. Chaves-Campos 2003, Terborgh 1977, Willis & Oniki In order to escape predation by army ants, arthropods 1978). Recent quantitative studies confirm that the flee from the approaching raid. Some of the fleeing density of army ant raids covaries with local net primary arthropods are preyed upon by birds that forage near productivity, which generally declines with elevation, army ant swarm raids (Coates-Estrada & Estrada 1989, and that army ant density declines with elevation at Otis et al. 1986, Swartz 2001, Willis & Oniki 1978). Army a neotropical montane forest site (Kaspari & O’Donnell ant colonies raid almost daily throughout their nomadic 2003, O’Donnell & Kumar 2006). colony cycle (Berghoff et al. 2002, Boswell et al. 1998, In lowland forests, habitat fragmentation affects Schneirla 1934, Topoff & Mirenda 1980). Eciton burchellii both insectivorous birds and army ants independently (Bierregaard et al. 1992, Boswell et al. 1998, Franks & Fletcher 1983, Lovejoy et al. 1984, Partridge et al. 1 Corresponding author. Email: [email protected] 1996, Roberts et al. 2000b, Sekercioglu et al. 2002). 582 ANJALI KUMAR AND SEAN O’DONNELL Insectivorousbirdspeciesarefirsttodisappearfromforests Five Holdridge life zones occur within a 600-m elevation fragments, implying that they are particularly sensitive range at Monteverde (Haber 2000, Holdridge 1966). to habitat change (Bierregaard et al. 1992, Karr 1982, The present study was conducted in continuous forest Sekercioglu et al. 2002, Willis 1974). Similarly, Eciton (11 sites) and forest fragments (7 sites) located between burchelliicoloniesdisappearfromforestfragmentsthatare 1100 m asl and 1680 m asl on the Pacific and Atlantic below a minimum size (Bierregaard et al. 1992, Britton slopes in the vicinity of Monteverde. et al. 1996, 1999, Meisel 2004, Partridge et al. 1996). However, the effects of fragmentation on the army ant- attending bird interaction are unknown. Furthermore, Opportunistic observations of birds at army ant swarms few systematic studies on army ant-bird interactions have been conducted in neotropical montane forest (Roberts et In the dry season of 2005 (12 January to 4 March) al. 2000a). The goal of this study was to estimate the we collected data on army ant density and diversity in combined effects of elevation and forest fragmentation the Monteverde area (O’Donnell & Kumar 2006). Army on interactions between army ants and bird species in ant raids were sampled using standardized trail walks neotropical montane forest by quantifying bird species at four elevationally separated sites, ranging from 1200 richness and abundance at army ant swarms. to 1650 m asl, and army ant activity was observed Recent research shows that the birds attending swarms opportunistically at other locations in the Monteverde are cleptoparasites, reducing prey capture by the army area.Whenaswarmraidfrontwasencountered,wenoted ants (Wrege et al. 2005). Because birds rely on ant swarms whether birds were in attendance. Birds were identified for food to varying degrees, and bird attendance decreases to species and we counted individuals when possible. To the foraging success of the ant raids (Wrege et al. 2005), be counted, birds had to be actively foraging either within fragmentation and elevation effects on the interaction 2 m of the front of the swarm, or in the army ant swarm. could have implications for both the attending birds All dry-season data were collected in continuous forest. and the ants. We tested for elevation and fragmentation effects on the potential cleptoparasitic pressure exerted by montane forest birds. Standardized observations of birds at army ant swarms Wehypothesizedthatarmyant-birdinteractionswould beaffectedbybothelevationandforestfragmentation.We Standardized observations of bird attendance at army made the following predictions: (1) the size and species ant swarms (Eciton burchellii and Labidus praedator)were richness of bird flocks attending army ant raids will made from 29 June to 11 December 2005. We observed a decrease with elevation; (2) the size and species richness of maximum of one swarm per day and alternated sampling bird flocks attending army ant raids will vary with habitat dates among elevations and habitat types to minimize type,withsmallerspecies-poorflocksinfragmentsrelative order effects. Army ant swarms were located by walking to continuous forest; (3) the potential for cleptoparasitism trails, and by referral from other observers to A.K. of army ants by attending birds will be affected by habitat After locating an army ant colony, the foraging trail fragmentation; and (4) bird species will vary in their rate was followed to the swarm front where the attending of attendance at army ant raids. We were particularly birds forage (Swartz 2001, Willis & Oniki 1978). The interested in assessing whether some montane forest following data were collected: (1) Start and end time. resident birds, and neotropical migrants, attend swarms (2) Habitat type (continuous or fragment). Habitat type regularly. Those that do may rely on army ant raids as a was determined by viewing aerial maps, by consulting food source. Guindon (1997), and during site visits. A fragment was defined as any patch of forest separated from another patch by at least 50 m of open habitat. We used fragments METHODS ranging in size from approximately 1 ha to 4 ha (Guindon 1997). (3) Elevation (all elevations were taken to the Study area nearest 1 m with a digital altimeter). The best spot for viewing the swarm front was chosen; The research was conducted in neotropical montane this was usually located off to one side and facing in forest in the Monteverde area, Costa Rica (10◦18N, the direction in which the swarm was moving. The 84◦47W). Monteverde is a well-studied cloud-forest observation period began 5 min after reaching the swarm habitat on the continental divide of the Tilaran´ front to allow the birds to resume their normal activities mountain range. Local habitats include privately (Coates-Estrada&Estrada1989). Eachobservationperiod protected continuous forest and adjacent forest fragments lasted 60 min total, unless the ants traversed over (Guindon 1997, Haber 2000, Harvey 2000). Over small unmanageable terrain or heavy rain began. We recorded changes in elevation, the habitat changes dramatically. bird attendance at the army ant swarm. To be counted Bird-army ant interactions 583 as an attendant,