Abstract:
In generalized, multispecies mutualisms, competition among members of one guild can influence the net benefits that each species in the other guild receives. Hence seasonal factors that affect the dynamics of competition can also affect net benefits, especially if the benefit or cost of mutualism also varies seasonally. In the Sonoran Desert, two common species of generalist ants compete for access to extrafloral nectaries on the fishhook barrel cactus Ferocactus wislizeni, but their relative abundances vary seasonally; one ant dominates more cacti in winter/spring, and the other in summer/autumn. Recently, a third ant species, which also varies in abundance seasonally, has appeared at our study sites and is competing with resident ants for access to cactus nectaries. This empirical system motivated us to examine a metapopulation model of competition for patches in an open system with periodic forcing. We use the model to: (1) illustrate three ways in which competing species may differ in their sensitivities to environmental conditions that are consistent with seasonally displaced patterns of abundance; (2) ask under what conditions the invasion of a third competitor into a two-species system could alter the sensitivity of the system to environmental forcing at low vs. high frequency; and (3) show how differences among competitors in the pattern of seasonal forcing alone can dramatically alter the outcome of a competitor invasion. We discuss implications of these results for the ways in which seasonal variation among competing mutualists, and the benefits and costs of mutualism, may affect the functioning of generalized, facultative mutualisms. © 2005 by the Ecological Society of America.
Abstract:
Traits that are attractive to mutualists may also attract antagonists, resulting in conflicting selection pressures. Here we develop the idea that increased floral nectar production can, in some cases, increase herbivory. In these situations, selection for increased nectar production to attract pollinators may be constrained by a linked cost of herbivore attraction. In support of this hypothesis, we report that experimentally supplementing nectar rewards in Datura stramonium led to increased oviposition by Manduca sexta, a sphingid moth that pollinates flowers, but whose larvae feed on leaf tissue. We speculate that nectar composition may provide information about plant nutritional status or defense that floral visitors could use to make oviposition decisions. Thus, selection by floral visitors and leaf herbivores may be inextricably intertwined, and herbivores may represent a relatively unexplored agent of selection on nectar traits.
Abstract:
Comparing populations that differ in access to mutualists can suggest how traits associated with these interactions have evolved. I discuss geographical and seasonal variation in the success of a primarily tropical mutualism (the fig/pollinator interaction), and evaluate some possible adaptations allowing it to persist at the edge of its range. Pollinators probably have difficulty in seasonal sites because 1) fig trees flower rarely in winter and 2) trees that do flower are less detectable and more difficult to reach. Fig biologists believe that seasonality must have selected for adaptations allowing pollinators to survive winter. However, geographical comparisons do not support two current ideas, the synchrony-breakdown hypothesis and the specificity-breakdown hypothesis. I pose two alternatives: plasticity of fruit and wasp developmental time, and adaptations of free-living fig wasps. I also distinguish between the impact of seasonality on monoecious versus dioecious figs; the latter group appear better adapted to reproduce in cool climates. A combination of comparative, observational, and experimental approaches has great potential for advancing our understanding of mutualisms. © 1989 Birkhäuser Verlag Basel.
PMID: 17257099;Abstract:
Mutualisms are ubiquitous in nature, as is their exploitation by both conspecific and heterospecific cheaters. Yet, evolutionary theory predicts that cheating should be favoured by natural selection. Here, we show theoretically that asymmetrical competition for partners generally determines the evolutionary fate of obligate mutualisms facing exploitation by third-species invaders. When asymmetry in partner competition is relatively weak, mutualists may either exclude exploiters or coexist with them, in which case their co-evolutionary response to exploitation is usually benign. When asymmetry is strong, the mutualists evolve towards evolutionary attractors where they become extremely vulnerable to exploiter invasion. However, exploiter invasion at an early stage of the mutualism's history can deflect mutualists' co-evolutionary trajectories towards slightly different attractors that confer long-term stability against further exploitation. Thus, coexistence of mutualists and exploiters may often involve an historical effect whereby exploiters are co-opted early in mutualism history and provide lasting 'evolutionary immunization' against further invasion. © 2007 Blackwell Publishing Ltd/CNRS.
