Ecological Differentiation Among Key Plant Mutualists from a Cryptic Ant Guild
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See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/225103359 Ecological differentiation among key plant mutualists from a cryptic ant guild Article in Insectes Sociaux · November 2011 DOI: 10.1007/s00040-011-0174-x CITATIONS READS 14 102 3 authors, including: Robert J Warren Volker Bahn State University of New York Buffalo State College Wright State University 75 PUBLICATIONS 1,536 CITATIONS 40 PUBLICATIONS 825 CITATIONS SEE PROFILE SEE PROFILE All content following this page was uploaded by Volker Bahn on 03 June 2014. The user has requested enhancement of the downloaded file. Insect. Soc. (2011) 58:505–512 DOI 10.1007/s00040-011-0174-x Insectes Sociaux RESEARCH ARTICLE Ecological differentiation among key plant mutualists from a cryptic ant guild R. J. Warren II • P. McAfee • V. Bahn Received: 2 April 2011 / Revised: 1 May 2011 / Accepted: 13 May 2011 / Published online: 3 June 2011 Ó International Union for the Study of Social Insects (IUSSI) 2011 Abstract As key dispersers of herbaceous seeds, Aphae- Such shifts could disrupt current community associations nogaster ants strongly influence the distribution of woodland and biotic interactions with ant-dispersed plants. plants in eastern North America. Ants within this genus are difficult to distinguish and often are identified by subgroup, Keywords Aphaenogaster Á Climate change Á but emerging research suggests they occupy species-specific Deciduous forest Á Facilitation Á Myrmecochory Á ecological niches. As such, distinct climatic requirements Species distribution among Aphaenogaster spp. might result in transient plant interactions with climate change. We examine whether there are ecological and distributional differences among Introduction Aphaenogaster species that coincide with current taxonomic differentiations. We use occurrence records for six Aphae- Ant-mediated seed dispersal (myrmecochory) occurs nogaster spp. that occur in deciduous forests in eastern North worldwide (Gorb and Gorb, 2003; Rico-Gray and Oliveira, America. We associate the geographic patterning of species 2007), and it is a major facilitative interaction in understory occurrence with temperature and precipitation data, and we woodland communities in eastern North America (N.A.) examine whether unique climatic niches characterize each with direct influence on a substantial part of plant diversity species. We then predict habitat suitability throughout (Beattie and Hughes, 2002; Ness et al., 2009). Myrmec- eastern North America using species distribution models. ochorous plants benefit from ant-mediated dispersal through For verification, we test how well the predicted ranges fit seed predator avoidance, decreased intraspecific competi- observed occurrences using novel data sets for each species. tion and placement in nutrient-rich environments (see Gorb We find that Aphaenogaster species within this cryptic genus and Gorb, 2003; Giladi, 2006; Rico-Gray and Oliveira, 2007 demonstrate unique ecological and geographic signatures. and references therein), and the ants benefit by receiving Each species within the subgroup generally responds dif- nutrition for larvae from lipid-rich seed appendages (Carroll ferently to temperature, and somewhat differently to and Janzen, 1973; Marshall et al., 1979; Morales and precipitation and seasonal variance, suggesting unique eco- Heithaus, 1998; Gammans et al., 2005). The benefits of this logical niches for each species. Our results indicate that each interaction are mutual, but the plants depend far more on the ant species may respond uniquely to changes in climate. ants than vice versa, making the interaction facultative for ants and obligate for plants (Pudlo et al., 1980; Ness et al., 2009; Warren et al., 2010). Myrmecochory long was considered a diffuse mutualism R. J. Warren II School of Forestry and Environmental Studies, Yale University, between a host of scavenging ants and spring-flowering New Haven, CT 06511, USA plants (Berg, 1966; Handel, 1976; Beattie et al., 1979; Beattie and Hughes, 2002; Garrido et al., 2002), but & R. J. Warren II ( ) Á P. McAfee Á V. Bahn researchers increasingly identify specific ant genera as Department of Biological Sciences, Wright State University, Dayton, OH 45435, USA dominant specialists in myrmecochorous interactions e-mail: [email protected] worldwide (Anderson, 1988; Espadaler and Gomez, 1996; 123 506 R. J. Warren II et al. Garrido et al., 2002; Gorb and Gorb, 2003; Rico-Gray and the rudis complex, particularly seed-dispersal studies, Oliveira, 2007). Indeed, the timing of propagule release may makes no discrimination among members within this sub- correspond with peak foraging in certain seed-dispersing ant group (also noted by Ness et al., 2009). However, Warren genera and these genera appear far more effective as seed et al. (2011) demonstrated that differences exist in the for- dispersers than other seed-gathering ants (Giladi, 2006; aging phenology of two members of the rudis complex, Boulay et al., 2007; Gove et al., 2007; Zelikova et al., 2008; suggesting species-specific ecological roles in seed dis- Bas et al., 2009; Manzaneda and Rey, 2009; Ness et al., persal. Aphaenogaster picea began foraging early enough in 2009). In eastern deciduous forests (USA), members of the the spring to overlap with seed set by all local myrmec- ant genus Aphaenogaster dominate these mutualist inter- ochorous plants, whereas A. rudis began foraging too late in actions (Ness et al., 2009). the spring to disperse early flowering species (Warren et al., The Aphaenogaster genus is cosmopolitan (Bolton, 2011). 2010) with approximately a dozen species occurring in If individual Aphaenogaster spp. respond dissimilarly to eastern N.A. (Creighton, 1950; Smith, 1979; Umphrey, rapid climate change, Warren et al. (2011)’s results indicate 1996). Aphaenogaster spp. colonies typically are found that the ant–plant mutualism may become disrupted. Myr- below rocks and logs in mesic deciduous forests in the mecochores comprise a large portion of understory plant eastern USA (Talbot, 1934; Beattie, 1978; Lynch et al., diversity, and a potential disruption of the ant mutualism 1980; Smallwood, 1982a). Given the high diversity of ant would entail great consequence for forest diversity. It is species worldwide, there are many challenges in species unfortunate, then, that little is known about how climate identification and taxonomic revisions are common. These impacts the current distributions of individual Aphaeno- considerations make genus-based ant classification a rela- gaster spp. We explore the climatic niches of individual tively useful way to approach ant research (Fisher and Aphaenogaster ant species to examine ecological and dis- Cover, 2007; Ward, 2007; Bolton, 2010; Gue´nard et al., tributional differences among Aphaenogaster species within 2010). In addition, the taxonomic status of several Aphae- the rudis complex that coincide with their current taxonomic nogaster species in the eastern USA remains unsettled, and status. We associate the geographic patterning of species members of this complex can be difficult to distinguish occurrence (from collection localities) with temperature and (Creighton, 1950; Umphrey, 1996). For these reasons, precipitation data, and we examine whether unique climate workers studying ant-mediated seed dispersal generally associations occur for each species. We then use these identify Aphaenogaster spp. to genus, particularly members climatic niche models to predict habitat suitability through- of a species/sub-species known as the fulva-rudis-texana out eastern N.A. using species distribution models (SDMs). complex (hereafter ‘rudis complex’), which also contains For verification, we test how well the predicted ranges fit the majority of seed-dispersing species (see Ness et al., 2009 observed occurrences using novel occurrence data sets. and references therein). In doing so, we generate probability distribution maps for The rudis complex includes six identified species in each species and, more importantly, assess whether mem- eastern N.A. (Umphrey, 1996). For Aphaenogaster species, bers of the rudis complex are ecologically equivalent or and ants in general, habitat partitioning appears to be driven require greater effort for species-specific ecological inves- by temperature (Brian, 1956; Bernstein, 1979; Lynch et al., tigations. These insights will help elucidate the interactions 1980; Smallwood, 1982a; Cerda` et al., 1997; Retana and between the ants and the plants, the seeds of which they Cerda, 2000; Dunn et al., 2007; Sanders et al., 2007), but disperse, and how these relationships may vary under climate moisture also plays a role (Cerda` et al., 1997; Warren et al., change. 2010). Whereas spatial and temporal habitat partitioning is common among ant species (see Parr and Gibb, 2010), little is known about species partitioning within genera (e.g., Methods Brian, 1956). Workers have demonstrated segregation between Aphaenogaster spp. and sympatric ant genera (e.g., Records of Aphaenogaster species and climate variables Lynch et al., 1980; Fellers, 1987), but only limited, mostly were obtained from published literature (Umphrey, 1996) anecdotal, information exists surrounding habitat segrega- and online databases to investigate the climatic niche and tion by species within the rudis complex (Talbot, 1934; create species distribution models (SDMs) for each member Crozier, 1977; Mitchell et al., 2002). Previous taxonomic of the rudis complex. Climate layers were