Ness, J. H. 2006. A mutualism's indirect costs: the most aggressive plant bodyguards also deter pollinators. Á Oikos 113: 506 Á514.Plant defenses against herbivores may be costly if they exclude mutualists. Here, I test the hypothesis that aggressive ant bodyguards of plants deter pollinators, and explore mechanisms by which Ferocactus wislizeni , an extrafloral nectary bearing cactus, limits conflicts between its pollinators and bodyguards. Flower visitation by ants and pollinating bees differed among plants tended by four different ant species. The ant species most rarely found in flowers showed the strongest aversion to F. wislizeni flower petals in laboratory assays, suggesting that those structures may include an ant-deterrent. Species-specific estimates of mean ant abundance within flowers and aggressiveness towards other arthropods were used to distinguish the relative threat of ant attack in flowers on plants tended by each ant species. Pollinator surveys in 2003 and 2004 demonstrated that bee visitation rates and the duration of flower occupation differed among plants with different ant associates, decreasing as the threat of ant attack increased. Flowers on plants tended by Solenopsis xyloni , the best ant bodyguard, were more dangerous than those on plants tended by three milder species, due to that ants' greater aggressiveness and abundance within flowers. These flowers were visited by pollinators least frequently and for less time per visit, and produced fruits with significantly lower total seed mass, fewer seeds, and lighter individual seeds, relative to fruits from similarly-sized plants tended by three other ant species. As a result, the best bodyguard may indirectly constrain plant reproduction in some settings. Conflicts between mutualistic guilds may be particularly common in generalized systems, where there is variation in partner quality and in the relative importance of the protection and pollination mutualisms.

Ant-dispersed herbs (myrmecochores) can account for more than one-third of the stems in the temperate deciduous forests of eastern North America. Because many ant species have been observed collecting the seeds, this interaction is often described as a generalized mutualism. Here, we combine fieldwork and meta-analyses to test this assumption. Our meta-analysis demonstrated that Aphaenogaster ants (predominantly A. rudis) collect approximately 74926% (mean9 SD) of the myrmecochorous seeds in eastern North American forests where any encounters with Aphaenogaster were reported, and approximately 61937% of the seeds in all the eastern forests where any seed collection has been monitored. This remarkable monopolization of seeds is due to at least two factors: 1) Aphaenogaster are significantly more likely to collect the ant-adapted seeds they discover than are ten other ant genera found in these forests and 2) the densities of Aphaenogaster and myrmecochorous plants are positively correlated at three nested spatial scales (within 20 ) 20 m patches, among patches within a forest, and among 41 forests in the eastern United States). Although other ants can collect seeds, our analyses demonstrate that A. rudis is the primary seed dispersal vector for most of this rich temperate ant-dispersed flora. The low levels of plant partner diversity for myrmecochores demonstrated here rivals that of tropical ant-plants (myrmecophytes) and well exceeds that typically observed in temperate plantÁfrugivore and plantÁpollinator mutualisms and myrmecochory in other biomes.

Cheating is a focal concept in the study of mutualism, with the majority of researchers considering cheating to be both prevalent and highly damaging. However, current definitions of cheating do not reliably capture the evolutionary threat that has been a central motivation for the study of cheating. We describe the development of the cheating concept and distill a relative-fitness-based definition of cheating that encapsulates the evolutionary threat posed by cheating, i.e. that cheaters will spread and erode the benefits of mutualism. We then describe experiments required to conclude that cheating is occurring and to quantify fitness conflict more generally. Next, we discuss how our definition and methods can generate comparability and integration of theory and experiments, which are currently divided by their respective prioritisations of fitness consequences and traits. To evaluate the current empirical evidence for cheating, we review the literature on several of the best-studied mutualisms. We find that although there are numerous observations of low-quality partners, there is currently very little support from fitness data that any of these meet our criteria to be considered cheaters. Finally, we highlight future directions for research on conflict in mutualisms, including novel research avenues opened by a relative-fitness-based definition of cheating.

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The services provided within a community can change as the species composition of that community changes. For example, ant–seed dispersal mutualisms can be disrupted in habitats dominated by invasive ants. We propose that this disruption is related to changes in mean ant body size, given that invasive ants are smaller than most native seed‐dispersing ants. We demonstrate that the mean and maximum distances that ants transport seeds adapted for ant dispersal increase with worker body size, and that this relationship is an accelerating power function. This pattern is consistent among three ant subfamilies that include most seed‐dispersing ants as well as most invasive ant species, is generalizable across ant species and communities, and is independent of diaspore mass. Using a case study, we demonstrate that both the mean body size of seed‐collecting ants and seed dispersal distances are decreased in sites invaded by Solenopsis invicta, the imported red fire ant. Furthermore, we demonstrate that the mean size of seed‐collecting ants at a seed depot or within a community is a useful predictor of mean seed dispersal distances at those sites. Last, we show that small seed‐collecting ants and decreased seed dispersal distances are common features of sites occupied by invasive ants. The link between ant body size and seed dispersal distance, combined with the dominance of invaded communities by typically small ants, predicts the disruption of native ant–seed dispersal mutualisms in invaded habitats.