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A Metacommmunity Approach to Co-Occurrence Patterns and the Core-Satellite Hypothesis in a Community of Tropical Arboreal Ants

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A Metacommmunity Approach to Co-Occurrence Patterns and the Core-Satellite Hypothesis in a Community of Tropical Arboreal Ants Ecol Res (2010) 25: 1129–1140 DOI 10.1007/s11284-010-0738-7 ORIGINAL ARTICLE George F. Livingston • Stacy M. Philpott A metacommmunity approach to co-occurrence patterns and the core-satellite hypothesis in a community of tropical arboreal ants Received: 6 April 2010 / Accepted: 31 May 2010 / Published online: 15 July 2010 Ó The Ecological Society of Japan 2010 Abstract Inferring mechanisms of community assembly included in the co-occurrence matrix, patterns become from co-occurrence patterns is difficult in systems where aggregated for all three species groupings. This suggests assembly processes occur at multiple spatial scales and that P. simplex ‘‘assembles’’ the community by providing a among species with heterogeneous dispersal abilities. core metapopulation that other species track. Analyzing Here, we demonstrate that local scale analysis of co-occurrence patterns among candidate subsets of spe- co-occurrence patterns is inadequate to fully describe cies, at multiple spatial scales, and linking them to species assembly mechanisms and instead utilize a metacommu- traits substantially improves the explanatory power of nity and core-satellite approach. We generated a co-occurrence analyses in complex metacommunities. co-occurrence and life-history data set for a community of twig-nesting ants on coffee plants across 36 sites within a Keywords Assembly Æ Core-satellite hypothesis Æ tropical agroecosystem to test the following three Metacommunity Æ Tropical ecology Æ Co-occurrence hypotheses: (1) twig-nesting ant species compete for nest-sites, (2) they are structured as a metacommunity, and (3) core species show segregated patterns, while satellite species show random patterns of co-occurrence. Species Introduction were divided into four groups: core species that are well distributed regionally and dominant locally, regional do- Species co-occurrence patterns have been used to infer a minants that are well distributed regionally but do not range of mechanisms assembling ecological communi- dominate locally, local dominants that are dominant ties. These mechanisms include interspecific competition locally but are not widely distributed, and satellites that are (Diamond 1975), habitat filtering (Horner-Devine et al. neither widely distributed nor dominant locally. Only the 2007), body size and energy constraints (Gotelli and most abundant species in the community, Pseudomyrmex McCabe 2002), phylogenetic and historical influences simplex, was classified as a core species. Regional domi- (Cavender-Bares et al. 2006), and neutral processes nants, local dominants, and satellite species show random (Kelly et al. 2008), among others. Dispersal plays an patterns of co-occurrence. However, when P. simplex is implicit role in driving many of these mechanisms. Habitat filtering, for example, requires that species are able to disperse to suitable environments. However, a G. F. Livingston general framework incorporating dispersal is lacking. Department of Ecology and Evolutionary Biology, The presence and magnitude of dispersal of interacting University of Michigan, 430 N. University, Ann Arbor, MI 48109, USA species among local communities (forming a metacom- munity, i.e., Leibold et al. 2004) should influence S. M. Philpott co-occurrence patterns given the diverse ways in which Department of Environmental Sciences, dispersal can impact species coexistence (Amarasekare University of Toledo, 2801 W Bancroft St., Mail Stop #604, Toledo, OH 43606, USA et al. 2004). Leibold and Mikkelson (2002) identified a number of metrics to describe co-occurrence patterns Present address: G. F. Livingston (&) within metacommunities, but these are largely indepen- Section of Integrative Biology, dent of assembly processes. Utilizing co-occurrence data University of Texas at Austin, is important given that relative species abundance dis- 1 University Station C0930, Austin, TX 78712, USA E-mail: [email protected] tributions are of limited use in distinguishing among Tel.: +1-208-7211145 mechanisms (Adler et al. 2007). 1130 Many metacommunities are composed of species with species. Specifically, we tested the following three heterogeneous dispersal abilities and that differ in niche hypotheses: (1) species compete for nest-sites, (2) they breadth (Pandit et al. 2009). One way to address dis- exist in local communities linked by dispersal (i.e., the persal heterogeneity is to identify ‘‘core’’ and ‘‘satellite’’ twig-nesting ant community is a metacommunity), and species (Hanski 1982). Core species are dominant in (3) core species show segregated co-occurrence patterns communities and are common at both local and regional due to the effects of interspecific competition, while scales, whereas satellite species are rare both locally and satellite species show random or aggregated patterns regionally. The ‘‘core-satellite’’ distinction can either resulting from stochastic dispersal. Taken together, form ends of a continuum or distinct clusters of species hypotheses one and two test for a competitive meta- depending on the assemblage considered (Hanski and community structure. We tested hypothesis one by Gyllenberg 1993). Interspecific competition among core measuring natural nests to determine if species use species should result in nonrandom segregated patterns similar nest-sites or if local scale microhabitat parti- of co-occurrence (Ulrich and Zalewski 2006). In con- tioning precludes direct interaction. We also experi- trast, satellite species should show random or aggregated mentally tested for nest-site preferences to confirm the co-occurrence patterns resulting from the effects of dis- role of competition in generating field patterns. We persal (Ulrich and Zalewski 2006) or habitat segregation tested hypotheses two and three by dividing the com- (Pandit et al. 2009). munity into four classes of core and satellite species Several hypotheses have been proposed to explain the based on clustering within an abundance-occupancy occurrence of core and satellite species. These include distribution and with respect to life-history traits. incomplete sampling (Selmi and Boulinier 2004) and We then ran co-occurrence simulations for each of these biological mechanisms such as the distribution of het- species groups. Finally, to determine the relative erogeneous habitats, dispersal ability, relative geometry influence of competition and stochastic dispersal on of geographic ranges, and metapopulation dynamics co-occurrence patterns we constructed a behavioral (McGeoch and Gaston 2002). However, few empirical dominance hierarchy and correlated the metacommuni- tests have distinguished among mechanisms driving ty abundance of each species with local community core-satellite patterns (but see Holt et al. 2002). properties. Co-occurrence analyses provide one way to test among possible mechanisms (Ulrich and Zalewski 2006). Studies further linking clusters of core and satellite Methods species to relevant life history traits (e.g., dispersal ability) are needed (Cadotte and Lovett-Doust 2007). We conducted this study during June and July of 2007 in In this study we focus on an arboreal community of a 45-ha plot within Finca Irlanda, a 300-ha traditional neotropical twig-nesting ants that opportunistically uti- organic shaded coffee agroecosystem in the Soconusco lize dry hollow twigs (Carroll 1979) on coffee plants region of Chiapas, Mexico (15°10¢ N and 92°20¢ W, within a tropical agroecosystem. Coffee plants act as 950 m elevation, 4,500 mm rain annually). The agro- uniformly distributed patches between which ants ecosystem is comprised of a shade canopy and an migrate via the dispersal of queens and direct movement understory of coffee plants. The canopy in Finca Irlanda of nests. Coffee is a relatively homogeneous environ- is diverse (>200 species) but dominated by Inga ment, with plants differing in a limited number of micheliana and Alchornea latifolia (Perfecto and Van- characteristics, including the size and number of nest- dermeer 2002). The farm is managed as a commercial sites available (effectively setting local carrying capac- polyculture (Moguel and Toledo 1999). ity), plant height, identity of nearby species of shade We used thirty-six randomly selected sites from across trees, and the extent of disturbance by an aggressive the plot (extending from a focal shade tree) to survey the carton nesting species, Azteca instabilis (Philpott 2010). twig-nesting ant community. To control for possible Importantly, previous research has demonstrated that effects of tree species we established half of the sites be- this community is nest-site limited (Philpott and Foster neath I. micheliana, and the other half beneath A. latifolia 2005). Research focusing on A. instabilis, other arboreal trees. To assess the impact of disturbance on the com- ants in the shade tree canopy, hemipteran assemblages, munity, half of these sites contained nests of A. instabilis predatory beetles, and fungal pathogens has demon- (at least 2 years old), while the remaining half did not and strated the capacity of coffee agroecosystems to eluci- were instead situated at least 20 m from A. instabilis nests. date mechanisms driving intrinsic pattern formation Sites were 1.5 · 10 m transects extending from the base of (e.g., Philpott 2006; Liere and Perfecto 2008; Livingston a shade tree. In addition to random tree selection, the et al. 2008; Vandermeer et al. 2008). Thus, abundance- direction of each transect was chosen at random among occupancy patterns can be interpreted against a back- the eight cardinal and intercardinal
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