Ecology Letters, (2004) 7: 1024–1028 doi: 10.1111/j.1461-0248.2004.00667.x IDEAS AND PERSPECTIVES Stoichiometric relations in an ant-treehopper mutualism
Abstract Adam D. Kay,1* Sara E. Scott,2 Carbon : nitrogen : phosphorus (C : N : P) stoichiometry can underlie physiological John D. Schade3,4 and and life history characteristics that shape ecological interactions. Despite its potential Sarah E. Hobbie1 importance, there is much to learn about the causes and consequences of stoichiometric 1 Department of Ecology, variation in terrestrial consumers. Here we show that treehoppers (Publilia modesta) Evolution and Behavior, tended by ants (Formica obscuripes) contained lower N concentrations than treehoppers on University of Minnesota, 100 plants from which ants were excluded. Ant presence also affected nutrient concentra- Ecology Building, 1987 Upper tions in host plants: on plants with ants, leaves contained uniformly low concentrations Buford Circle, St Paul, MN of N; on plants without ants, N concentrations were low only in the few leaves fed upon 55108, USA 2Department of Biology, by treehoppers at the time of collection. We suggest treehopper feeding reduces leaf College of St Benedict, St nutrient levels and ants positively affect treehopper abundance, producing a top–down Joseph, MN, USA effect on plant quality. Determining the causes of these stoichiometric changes should 3School of Life Sciences, Arizona help elucidate factors guiding the dynamics of conditional mutualisms between ants and State University, Tempe, AZ, homopterans. USA 4Department of Integrative Keywords Biology, University of California, Ants, conditional, membracid, mutualism, nitrogen, phosphorus, Publilia, Solidago, Berkeley, CA, USA stoichiometry, treehoppers. *Correspondence: E-mail: [email protected] Ecology Letters (2004) 7: 1024–1028
Here, we investigate how ant attendance affects the size INTRODUCTION of aggregations of the treehopper, Publilia modesta, and the Outcomes of conditional mutualisms often depend on carbon : nitrogen : phosphorus (C : N : P) stoichiometry varying ecological factors (Bronstein 1994, 2001; Herre et al. of P. modesta individuals. Stoichiometry can underlie traits 1999). Such conditionality is thought to be widespread in that affect how organisms interact with their environment ant-homopteran mutualisms (Cushman & Addicott 1991), in (Sterner & Elser 2002): one example is the causal which homopterans produce excretions (ÔhoneydewÕ) col- relationship between P concentration and growth rate. lected by ants, and ants provide protection or other services. Although this relationship is taxonomically widespread Ant attendance can positively affect the size of a homop- (Elser et al. 2003), little is known about patterns, causes, teran aggregation (number of individuals on a host plant) or consequences of stoichiometric variation in most depending on several factors including ant abundance terrestrial consumers. (Cushman & Whitham 1991; Morales 2000) and natural Ant attendance could affect treehopper stoichiometry by at enemy levels (Bristow 1984). Ant attendance can also least two mechanisms. First, ant presence could lead to an influence characteristics of individual homopterans such as increase in P in treehoppers if ant protection creates growth rate (Bristow 1984; Stadler & Dixon 1998) by conditions for rapid growth, and growth rate in treehoppers inducing changes in feeding behaviour (Banks & Nixon is tied to body P concentration. When tended by ants, some 1958) and the quantity or quality of honeydew that homopterans increase feeding rates (Banks & Nixon 1958) homopterans produce (Yao & Akimoto 2001). Discovering and may grow faster (El-Ziady 1960; Bristow 1984; Flatt & these individual-level effects, and determining their causes Weisser 2000); however, nothing is known about how such and consequences, may elucidate mechanisms underlying changes are related to homopteran stoichiometry. Second, ant the distribution and strength of ant-homopteran associa- attendance could decrease N and P concentrations in tions. treehoppers if treehoppers produce more nutrient-rich