J. H. Ness

The Effects of Invasive Ants on Prospective Ant Mutualists

Ants are recognized for their abilities both to engage in mutualistic interactions with diverse taxa, and to invade and dominate habitats outside their native geographic range. Here, we review the effects of invasive ants on three guilds of mutualists: ant-dispersed plants, ant-tended arthropods, and ant-tended plants. We contrast how those three guilds are affected by invasions, how invasive ants differ from native ants in their interactions with those guilds, and how the seven most invasive ant species differ amongst themselves in those interactions. Ant-dispersed plants typically suffer from interactions with invasive ants, a result we attribute to the small size of those ants relative to native seed-dispersing ants. Effects on the ant-tended arthropods and plants were more frequently positive or non-significant, although it is unclear how often these interactions are reciprocally beneficial. For example, invasive ants frequently attack the natural enemies of these prospective mutualists even in the absence of rewards, and may attack those prospective mutualists. Many studies address whether invasive ants provide some benefit to the partner, but few have asked how invasives rank within a hierarchy of prospective mutualists that includes other ant species. Because ant invasions typically result in the extirpation of native ants, this distinction is highly relevant to predicting and managing the effects of such invasions. Interspecific comparisons suggest that invasive ants are poorer partners of ant-dispersed plants than are most other ants, equally effective partners of ant-tended arthropods, and perhaps better partners of ant-tended plants. Last, we note that the invasive ant taxa differ amongst themselves in how they affect these three mutualist guilds, and in how frequently their interactions with prospective mutualists have been studied. The red imported fire ant, Solenopsis invicta, appears particularly likely to disrupt all three mutualistic interactions, relative to the other six invasive species included in this review.

Ant body size predicts dispersal distance of ant-adapted seeds: Implications of small-ant invasions

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 si...

Cheaters must prosper: reconciling theoretical and empirical perspectives on cheating in mutualism

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.

ENVIRONMENTAL FORCING AND THE COMPETITIVE DYNAMICS OF A GUILD OF CACTUS-TENDING ANT MUTUALISTS

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.

Abiotic influences on the behaviour of rodents, ants, and plants affect an ant-seed mutualism

ABSTRACT Deserts are open environments characterized by striking shifts in temperature and light regimes. We hypothesized that the abiotic environment mediates the interaction between an ant-dispersed plant, Datura wrightii (Solanaceae), ant mutualists, and rodent seed predators in the Sonoran Desert. Field experiments contrasting diurnal and nocturnal seed collection rates in the presence of ants only, vertebrates only, and in the presence of both groups indicated that 85%of seed collection by mammalian seed predators occurred at night (between 1900 and 0700). Seed collection by ants, in contrast, was similar between day and night, although seed collection decreased during very hot days and very bright nights. The total number of seeds collected by both groups foraging separately exceeded the number removed when both groups shared access to seed depots, suggesting that ants and rodents compete for seeds. However, D. wrightii plants dehisced 86%of their fruits between 0700 and 1900, increasing the likelihood of seed collection by ant mutualists rather than rodent seed predators. Dehiscence was sensitive to environmental cues: greenhouse plants kept at constant temperature and humidity dehisced 47%of their fruits between 0700 and 1900. Additional field experiments demonstrated that seed-collecting ants transported seeds considerable distances to their nests, microsites that can be rich in nutrients. The mean (±SE) seed dispersal distance was 6.1 ±0.5 m, the longest mean dispersal distance yet reported for an ant-dispersed seed.

The demographic consequences of mutualism: ants increase host-plant fruit production but not population growth

The impact of mutualists on a partner’s demography depends on how they affect the partner’s multiple vital rates and how those vital rates, in turn, affect population growth. However, mutualism studies rarely measure effects on multiple vital rates or integrate them to assess the ultimate impact on population growth. We used vital rate data, population models and simulations of long-term population dynamics to quantify the demographic impact of a guild of ant species on the plant Ferocactus wislizeni. The ants feed at the plant’s extrafloral nectaries and attack herbivores attempting to consume reproductive organs. Ant-guarded plants produced significantly more fruit, but ants had no significant effect on individual growth or survival. After integrating ant effects across these vital rates, we found that projected population growth was not significantly different between unguarded and ant-guarded plants because population growth was only weakly influenced by differences in fruit production (though strongly influenced by differences in individual growth and survival). However, simulations showed that ants could positively affect long-term plant population dynamics through services provided during rare but important events (herbivore outbreaks that reduce survival or years of high seedling recruitment associated with abundant precipitation). Thus, in this seemingly clear example of mutualism, the interaction may actually yield no clear benefit to plant population growth, or if it does, may only do so through the actions of the ants during rare events. These insights demonstrate the value of taking a demographic approach to studying the consequences of mutualism.