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Friend or foe? A behavioral and stable isotopic investigation of an ant-plant symbiosis.

In ant-plant symbioses, the behavior of ant inhabitants affects the nature of the interaction, ranging from mutualism to parasitism. Mutualistic species confer a benefit to the plant, while parasites reap the benefits of the interaction without reciprocating, potentially resulting in a negative impact on the host plant. Using the ant-plant symbiosis between Cordia alliodora and its ant inhabitants as a model system, I examine the costs and benefits of habitation by the four most common ant inhabitants at La Selva Biological Station, Costa Rica. Costs are measured by counting coccoids associated with each ant species. Benefits include patrolling behavior, effectiveness at locating resources, and recruitment response. I also compare the diets of the four ant species using stable isotope analysis of nitrogen (N) and carbon (C). Ants varied in their rates of association with coccoids, performance of beneficial behaviors, and diet. These differences in cost, benefit, and diet among the ant species suggest differences in the nature of the symbiotic relationship between C. alliodora and its ants. Two of the ant species behave in a mutualistic manner, while the other two ant species appear to be parasites of the mutualism. I determined that the mutualistic ants feed at a higher trophic level than the parasitic ants. Behavioral and dietary evidence indicate the protective role of the mutualists, and suggest that the parasitic ants do not protect the plant by consuming herbivores.

Animals↗

Personal protection measures against fire ant attacks.

BACKGROUND: Fire ant stings lead to significant morbidity and mortality each year. Virtually no information exists in the scientific literature about the effectiveness of personal protection measures against these ants. OBJECTIVES: To quantify the level of protection from ant stings by socks and cotton tights and to evaluate the efficacy of repellents and other chemicals in preventing stings. METHODS: Commercially obtained infant socks were fitted on a plastic doll's foot, which was touched to fire ant mounds in a series of controlled experiments. Socks and cotton tights were stretched over a human finger and placed in a fire ant mound to test the ability of ants to sting through fabric. Assays were developed to screen chemical substances for their ability to stop or prevent fire ant stings. RESULTS: Socks of any type reduced the number of fire ants that reached the skin and delayed the time required for ants to reach the skin above the sock level. Fire ants were unable to sting through all socks tested. Ants successfully stung both paper towels and human skin, regardless of chemical substance or repellent applied to them. CONCLUSIONS: Socks provide some degree of protection from fire ant stings; therefore, children living in fire ant-infested areas should wear them. Fire ants appear unable to sting through many commercially available socks. Cotton tights may be useful in protecting the lower extremities of children. Fire ants are not deterred from stinging by a wide variety of insect repellents and chemical substances.

Animals↗

Ant allergy in Asia and Australia.

PURPOSE OF REVIEW: Anaphylaxis due to ant sting is increasingly being recognized as a significant problem. Severe allergic reactions to ants are well described in the south-eastern United States, but have only been recognized in recent years as being important in other parts of the world. There are many different ant species and their distribution around the world varies. The purpose of this review is to familiarize the reader with the epidemiology and clinical characteristics of ant allergy in Asia and Australia. RECENT FINDINGS: In Korea, allergy to Pachycondyla chinensis (subfamily Ponerinae) has been well described. In an ant-endemic area, sensitization was 23%, with about 1% having anaphylactic reactions. There were at least eight IgE-binding proteins in P. chinensis venom, with 1 major allergen binding 85% of patient sera. P. chinensis venom was also found to be possibly crossreactive with bee venom, but not with imported-fire-ant venom. In Australia, anaphylactic reactions to ant stings are usually caused by the 'jack jumper' ant (Myrmecia pilosula) or the bull ant (Myrmecia pyriformis). A recent study showed promising results for immunotherapy with M. pilosula venom. There have been reports of stings by other ant species in Asia and Australia, but these reports are few and sporadic. SUMMARY: The study of ant allergy in Asia is in its infancy. Clinicians in Asia need to be aware of ant stings as a cause of severe allergic reactions. Certain species that cause allergic reactions are unique to Asia and Australia and deserve further research. The allergens in the venom of the different ant species need to be identified. We should aim for improved understanding of the epidemiology of ant-sting anaphylaxis, formulation of better diagnostic tests and possibly the introduction of immunotherapeutic strategies.

Adrenergic Agonists↗

Behavioural responses of Eurasian treecreepers, Certhia familiaris, to competition with ants

Competition for a specific resource that is essential for the survival of both the competitors may be intense even between very dissimilar taxa. However, the importance of the effects caused by such interspecific competition has seldom been emphasized. These effects can appear as differences in individual foraging behaviour during the breeding season, which can result in critical variation in fitness. In this study we examined the effects of wood ants (Formica rufa group) on the abundance of other invertebrates on tree trunks and on the foraging site selection of breeding Eurasian treecreepers, which use the same habitat as wood ants. Arthropods were scarcer on the trunks with ants present; the treecreepers avoided these trunks and foraged for a shorter time on trunks with ants than on trunks without ants. We also tested experimentally the existence of competition between ants and treecreepers by comparing the foraging behaviour of breeding treecreepers on spruce trunks with ants, without ants and with experimentally reduced numbers of ants. On average arthropods were scarcest on trunks with ants present. Male treecreepers also foraged for a shorter time on spruce trunks with ants. The reduction in ant numbers allowed food resources on trunks to recover over a week and led to longer foraging times of the treecreepers on these trunks than on trunks with ants present. The longest treecreeper visits were on trunks without ants. Our results suggest that competition between two very different taxa may be effective in determining the behaviour of foraging individuals.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour

Journal Article↗

Behavioral mechanisms underlie an ant-plant mutualism.

Predators can reduce herbivory by consuming herbivores (a consumptive effect) and by altering herbivore behavior, life history, physiology or distribution (non-consumptive effects). The non-consumptive, or trait-mediated, effects of predators on prey may have important functions in the dynamics of communities. In a facultative ant-plant mutualism, we investigated whether these non-consumptive effects influenced the host plants of prey. Here, predaceous ants (Forelius pruinosus) consume and disturb a dominant lepidopteran folivore (Bucculatrix thurberiella) of wild cotton plants (Gossypium thurberi). Season-long ant exclusion experiments revealed that ants had a larger proportional effect on damage by B. thurberiella than on caterpillar abundance, a result that suggests ants have a strong non-consumptive effect. Behavioral experiments conducted in two populations over 2 years demonstrated that B. thurberiella caterpillars were substantially less likely to damage wild cotton leaves in the presence of ants due to ant-induced changes in caterpillar behavior. In the absence of ants caterpillars spent more time stationary (potential feeding time) and less time dropping from leaves by a thread of silk than when ants were present. Furthermore, ants altered the spatial distribution of both caterpillars and damage; caterpillars spent relatively more time on the upper surfaces of leaves and caused damage further from the leaf margin in ant exclusion treatments. Both direct encounters with ants and information conveyed when ants walked onto leaves were key events leading to the anti-predator behaviors of caterpillars. This study contributes to a small body of evidence from terrestrial systems demonstrating that the trait-mediated effects of predators can cascade to the host plants of prey.

Animals↗

Intrapopulation differences in ant eating in the mountain gorillas of Bwindi Impenetrable National Park, Uganda.

Variability in ant eating has been observed in several populations of eastern and western gorillas. We investigated the occurrence of ant (Dorylus sp.) eating in two groups of mountain gorillas (Gorilla beringei beringei) with overlapping home ranges within Bwindi Impenetrable National Park, Uganda from September 2001 to August 2002. We calculated the frequency of ant eating by an indirect method of analyzing fecal samples from silverbacks, adult females, and juveniles. One group consumed ants significantly more often than the other (3.3 vs 17.6% of days sampled). Furthermore, the group that consumed ants more often also consumed them on a seasonal basis (September-February monthly range: 0-8%; March-August monthly range: 30-42.9%). Finally, females and juveniles of this group consumed ants significantly more often than did the silverback (total samples containing ants: silverback, 2.1%; adult female, 13.2%; juvenile, 11.2%). Differences in ant eating between groups are likely due to variability in use of habitats where ants occur (particularly secondary forests). Surveys of ant densities in differing habitats, nutritional analysis of ants, and quantification of the amount of ants in their diets are necessary to understand if ant consumption is due to availability, nutritional value, group traditions, or taste preference.

Age Factors↗

Anaphylaxis to venom of the Pachycondyla species ant.

BACKGROUND: In the southeastern United States, imported fire ants have caused systemic reactions with a high incidence. On the contrary, in Korea Pachycondyla species ants (P chinensis and P solitaria), and the family Formicidae, which are in the genus Pachycondyla and the subfamily Ponerinae, have only occasionally caused systemic reactions. OBJECTIVE: We sought to assess whether commercially available imported fire ant extract would be useful in treating patients with anaphylaxis induced by venom from a Pachycondyla species ant. METHODS: Serum samples were collected from 2 women who had anaphylaxis induced by Pachycondyla species ant venom and from 6 volunteers with no history of having been stung. Specific IgE to Pachycondyla species ant extracts was measured by means of ELISA and possible allergenic components by immunoblot. Cross-reactivity between Pachycondyla chinensis, P solitaria, and imported fire ant extracts was also measured by inhibitory ELISA. RESULTS: Skin prick test responses were strongly positive to the extract of P chinensis (1:20 wt/vol) in the patient. Ten healthy volunteers exhibited negative responses. The 2 patients' sera exhibited high ELISA values, with absorbencies of 0.78 and 0.61 for P chinensis and 0.83 and 0.68 for P solitaria, respectively, and negative ELISA values for the extract of imported fire ants (absorbency <0.01). Imported fire ants showed no inhibition of the IgE binding to P chinensis or P solitaria. Possible allergenic components of Pachycondyla species ant extracts are 29- and 27-kd proteins and, less frequently, 16 kd proteins. CONCLUSION: Our data suggest that patients who have had an anaphylactic reaction to a Pachycondyla species ant might not benefit from immunotherapy with an imported fire ant extract. Immunotherapy with the extract of Pachycondyla species ants is expected to be highly effective.

Adult↗

Myrmecophily in Hesperiidae. The case of Vettius tertianus in ant gardens.

The larvae of the hesperiid butterfly Vettius tertianus develop by eating the leaves of Aechmea mertensii, a bromeliad epiphyte restricted to ant gardens. The relationships between ants and V. tertianus larvae highlight the preferential association of the caterpillars with Pachycondyla goeldii (Ponerinae), an ant-garden initiator. The oviposition strategy of V. tertianus may thus imply the identification of the inhabiting ant species and not only the identification of the host plant. The caterpillars neither provide secretions to the ants, nor possess defensive devices (i.e. hairs or appendices) against ants. Their activity rhythm does not isolate them from foraging workers of P. goeldii and their shelters are also attainable by the ants. Moreover, as the cuticular lipid profiles of V. tertianus larvae are clearly different from those of the ants and also from the leaf-surface of A. mertensii, acceptance is not due to mimicry between larvae and plants or ants. However, the caterpillars deposit, on the leaf they eat, silk containing a mixture of substances very similar to those found on their own cuticle. No interaction with ants was recorded during observations, even though the ant gardens were patrolled by numerous P. goeldii individuals during their activity period. But when confronted with the caterpillar, none of the tested ant species reacted aggressively. These results suggest the existence of compounds, other than cuticular lipids, responsible for the absence of aggressiveness in the ants. The case of V. tertianus is relatively new as myrmecophily within Hesperiidae has been described only once. Moreover, it preferentially involves a member of the Ponerinae, a subfamily in which interactions with other arthropods are exceptional.

Animals↗

Extensive exchange of fungal cultivars between sympatric species of fungus-growing ants.

Fungal cultivars of fungus-growing ants (Attini, Formicidae) are carried by dispersing queens from parent to offspring nest. This vertical cultivar transmission between generations is thought to result in long-term ant-fungus coevolution and selection for beneficial cultivar traits that maximize harvests and thus colony productivity. In contrast to this traditional view of vertical cultivar transmission, frequent horizontal cultivar transmission between ant species is implicated by a phylogenetic analysis of 72 cultivars propagated by two fungus-growing ant species coexisting sympatrically in central Panama. The two ant species are specialized on the same group of closely related cultivars, but in six of 12 cultivar clades identifiable within this group, cultivars from both ant species were united in the same clade. Five of these 'mixed' clades were supported by bootstrap values of about 90% or higher. In one instance, colonies from the two ant species cultivated the same, genetically identical, cultivar clone. These phylogenetic patterns indicate that: (i) cultivar exchanges between the two ant species occur routinely throughout ecological time; and that (ii) coevolutionary processes between ants and their fungi are more diffuse than previously assumed. Because the two ant species are specialized on a narrow group of closely related cultivars that they regularly exchange among each other, but not with other sympatric ant species, cultivar exchanges are constrained, most likely, by ant preferences for their own cultivar group or by stringent selection against transitions of ant lineages to distantly related cultivars.

Animals↗

Influence of the hypogaeic army ant Dorylus (Dichthadia) laevigatus on tropical arthropod communities.

The majority of army ant species forage hypogaeically. Due to the difficulties in observing these ants, their potential influence on hypogaeic and epigaeic arthropod communities has not yet been investigated. As the first hypogaeically foraging army ant studied in detail, we attracted Dorylus laevigatus to areas monitored for their arthropod diversity. Here, for the first time, the same sites were sampled before and after an army ant raid. Furthermore, interactions between D. laevigatus and the five most common ground-nesting ant species were noted and their life-history traits compared, allowing first inferences on possible mechanisms of their coexistence. The occurrence of D. laevigatus within a study plot had no evident effect on the number of arthropod taxa or individuals collected with epigaeic and hypogaeic pitfall traps. Likewise, juvenile arthropods, which are less mobile and thus are potentially easier prey for D. laevigatus, showed no differences in their collected numbers before and after the army ant had visited a plot. However, significantly fewer ant species were collected with hypogaeic traps after D. laevigatus had been within the study plots, indicating a possible predation of D. laevigatus especially on two Pseudolasius and one Pheidole species. The five most common ground-foraging ant species demonstrated their ability to avoid, kill, and even prey on the army ant. The reaction of Lophomyrmex bedoti towards D. laevigatus indicated the former to be a potential prey species, while Pachycondyla sp. 2 showed signs of "enemy specification." Odontoponera diversus and O. transversa actively preyed on D. laevigatus, while Pheidologeton affinis fought with D. laevigatus over resources. All ant species could co-occur with D. laevigatus at palm oil baits. Adding to the differences detected in previous studies between D. laevigatus and epigaeically foraging army ant species, the occurrence of this hypogaeic army ant seems to have less devastating effects on arthropod community compositions than those of epigaeically mass raiding species.

Animals↗

A comparative analysis of morphological and ecological characters of European aphids and lycaenids in relation to ant attendance.

Ants are a major environmental factor for many insect species. For example, aphids and lycaenids have evolved an array of associations with ants ranging from obligate myrmecophily to the avoidance of contact. Here we (1) analyze the predictive power of different ecological and morphological traits for explaining the strength of the association between ants and aphids/lycaenids and (2) contrast different taxonomic levels with respect to the variance explained by ant attendance. Data come from a literature survey including 112 species of aphids and 103 species of lycaenids from Europe. For aphids, feeding on woody plant parts is positively associated with ant attendance, while a high degree of mobility, feeding in isolation, and the possession of wings in the adult stage are negatively associated with ant attendance. In lycaenids, feeding on inflorescences and feeding on Fabaceae host plants is closely associated with ant attendance, while living in forests bears a smaller likelihood to establish mutualistic relationships. Body size always appeared to be a poor predictor for the degree of ant attendance. Overall, in both insect groups less than 10% of the variation in the ecological traits recorded is explained by the different modes of ant association. When decomposing the variance in traits explained by ant attendance at different taxonomic levels, aphids and lycaenids show contrasting results. In aphids, most variance in the degree of ant attendance is explained at the subfamily level and least at the species level. The opposite is true for lycaenids, where most variance is explained at the lowest taxonomic level. Possible mechanisms explaining these different patterns of associations with ants are suggested.

Adaptation, Physiological↗

Disentangling a rainforest food web using stable isotopes: dietary diversity in a species-rich ant community.

For diverse communities of omnivorous insects such as ants, the extent of direct consumption of plant-derived resources vs. predation is largely unknown. However, determination of the extent of "herbivory" among ants may be crucial to understand the hyper-dominance of ants in tropical tree crowns, where prey organisms tend to occur scarcely and unpredictably. We therefore examined N and C stable isotope ratios (delta(15)N and delta(13)C) in 50 ant species and associated insects and plants from a tropical rainforest in North Queensland, Australia. Variation between ant species was pronounced (range of species means: 7.1 per thousand in delta(15)N and 6.8 per thousand in delta(13)C). Isotope signatures of the entire ant community overlapped with those of several herbivorous as well as predacious arthropods. Variability in delta(15)N between ants was not correlated with plant delta(15)N from which they were collected. Ant species spread out in a continuum between largely herbivorous and purely predacious taxa, with a high degree of omnivory. Ant species' delta(15)N were consistent with the trophic level predicted by natural feeding observations, but not their delta(13)C. Low delta(15)N levels were recorded for ant species that commonly forage for nectar on understorey or canopy plants, intermediate levels for species with large colonies that were highly abundant on nectar and honeydew sources and were predacious, and the highest levels for predominantly predatory ground-foraging species. Colonies of the dominant weaver-ants (Oecophylla smaragdina) had significantly lower delta(15)N in mature forests (where preferred honeydew and nectar sources are abundant) than in open secondary vegetation. N concentration of ant dry mass showed only very limited variability across species and no correlation with trophic levels. This study demonstrates that stable isotopes provide a powerful tool for quantitative analyses of trophic niche partitioning and plasticity in complex and diverse tropical omnivore communities.

Animals↗

Ant behaviour and seed morphology: a missing link of myrmecochory.

Seed dispersal by ants (myrmecochory) is mediated by the presence of a lipid-rich appendage (elaiosome) on the seed that induces a variety of ants to collect the diaspores. When seeds mature or fall onto the ground, these ant species transport them to their nest. After eating the elaiosome, the seed is discarded in nest galleries or outside, in the midden or farther away, where seeds can potentially germinate. The final location of seeds with their elaiosomes removed was evaluated to assess the importance of possible handles (structures that ants can grasp to carry) in transporting ants during re-dispersal experiments of seeds from nests of six species of ants. The results indicate that seeds remained within the nest because the ants were not able to transport them out of the nest. As a consequence of the elaiosome being removed, small ant species could not take Euphorbia characias seeds out of their nests. Only large ant species could remove E. characias seeds from their nests. Attaching an artificial handle to E. characias seeds allowed small ant species to redistribute the seeds from their nests. On the other hand, Rhamnus alaternus seeds that have a natural handle after the elaiosome removal were removed from the nests by both groups of ant species. If a seed has an element that acts as a handle, it will eventually get taken out of the nest. The ants' size and their mandible gap can determine the outcome of the interaction (i.e. the pattern of the final seed shadow) and as a consequence, could influence the events that take place after the dispersal process.

Animals↗

Habitat complexity facilitates coexistence in a tropical ant community.

The role of habitat complexity in the coexistence of ant species is poorly understood. Here, we examine the influence of habitat complexity on coexistence patterns in ant communities of the remote Pacific atoll of Tokelau. The invasive yellow crazy ant, Anoplolepis gracilipes (Smith), exists in high densities on Tokelau, but still coexists with up to seven other epigeic ant species. The size-grain hypothesis (SGH) proposes that as the size of terrestrial walking organisms decreases, the perceived complexity of the environment increases and predicts that: (1) leg length increases allometrically with body size in ants, and (2) coexistence between ant species is facilitated by differential habitat use according to body size. Analysis of morphological variables revealed variation inconsistent with the morphological prediction of the SGH, as leg length increased allometrically with head length only. We also experimentally tested the ability of epigeic ants in the field to discover and dominate food resources in treatments of differing rugosity. A. gracilipes was consistently the first to discover food baits in low rugosity treatments, while smaller ant species were consistently the first to discover food baits in high rugosity treatments. In addition, A. gracilipes dominated food baits in planar treatments, while smaller ant species dominated baits in rugose treatments. We found that the normally predictable outcomes of exploitative competition between A. gracilipes and other ant species were reversed in the high rugosity treatments. Our results support the hypothesis that differential habitat use according to body size provides a mechanism for coexistence with the yellow crazy ant in Tokelau. The SGH may provide a mechanism for coexistence in other ant communities but also in communities of other terrestrial, walking insects that inhabit a complex landscape.

Animals↗

Evaluating alternative hypotheses for the early evolution and diversification of ants.

Ants are the world's most diverse and ecologically dominant eusocial organisms. Resolving the phylogeny and timescale for major ant lineages is vital to understanding how they achieved this success. Morphological, molecular, and paleontological studies, however, have presented conflicting views on early ant evolution. To address these issues, we generated the largest ant molecular phylogenetic data set published to date, containing approximately 6 kb of DNA sequence from 162 species representing all 20 ant subfamilies and 10 aculeate outgroup families. When these data were analyzed with and without outgroups, which are all distantly related to ants and hence long-branched, we obtained conflicting ingroup topologies for some early ant lineages. This result casts strong doubt on the existence of a poneroid clade as currently defined. We compare alternate attachments of the outgroups to the ingroup tree by using likelihood tests, and find that several alternative rootings cannot be rejected by the data. These alternatives imply fundamentally different scenarios for the early evolution of ant morphology and behavior. Our data strongly support several notable relationships within the more derived formicoid ants, including placement of the enigmatic subfamily Aenictogitoninae as sister to Dorylus army ants. We use the molecular data to estimate divergence times, employing a strategy distinct from previous work by incorporating the extensive fossil record of other aculeate Hymenoptera as well as that of ants. Our age estimates for the most recent common ancestor of extant ants range from approximately 115 to 135 million years ago, indicating that a Jurassic origin is highly unlikely.

Animals↗

Pharaoh ant (Monomorium pharaonis): newly identified important inhalant allergens in bronchial asthma.

The nonstinging house ant, Monomorium pharaonis (pharaoh ant), was recently identified as a cause of respiratory allergy. This study was performed to evaluate the extent of sensitization to pharaoh ant, and its clinical significance in asthmatic patients. We carried out skin prick tests in 318 patients with asthma. Specific IgE (sIgE) to pharaoh ant was measured by ELISA, and cross-reactivity was evaluated by ELISA inhibition tests. Bronchial provocation testing was performed using pharaoh ant extracts. Fifty-eight (18.2%) of 318 patients showed positive skin responses to pharaoh ant, and 25 (7.9%) had an isolated response to pharaoh ant. Positive skin responses to pharaoh ant were significantly higher among patients with non-atopic asthma than among those with atopic asthma (26.0% vs. 14.9%, p<0.05). There was significant correlation between sIgE level and skin responses to pharaoh ant (rho=0.552, p<0.001). The ELISA inhibition tests indicated that pharaoh ant allergens had various pattern of cross-reactivity to house dust mites and cockroaches. Bronchial provocation tests to pharaoh ant were conducted for 9 patients, and eight showed typical asthmatic reactions. In conclusion, pharaoh ant is an important source of aeroallergens, and it should be included in the skin test battery for screening the causative allergens in patients with asthma.

Administration, Inhalation↗

The effects of food presentation and microhabitat upon resource monopoly in a ground-foraging ant (Hymenoptera: Formicidae) community.

In Neotropical wet forests several species of omnivorous, resource-defending ants, live and forage in close proximity to one another. Although the forest floor is heterogeneous in microhabitat and food quantity, little is known about the impact of microhabitat and food variation upon resource monopoly among ants. We investigated how food type and microhabitat influence food monopoly in resource-defending ants in old-growth tropical wet forest in the Caribbean lowlands of Costa Rica. We measured several microhabitat characteristics at 66 points in a 0.5 hectare plot, and baited each point with two categories of tuna bait. These baits were presented in "split" and "clumped" arrangements. We measured the frequency of bait monopoly by a single species, as well as the number of recruited ant foragers at a bait. Out of five common species, two (Wasmannia auropunctata and Pheidole simonsi) more frequently monopolized one bait type over the other, and one (P. simonsi) recruited more ants to the split baits. We then considered the recruitment response by all ant species in the community. We found that the frequency of monopoly, sharing, and the absence of ants at a given point in the rainforest differed with bait type. The frequency of monopoly was associated with microhabitat type in two out of eight microhabitat variables (leaf litter depth and palms); variation in two other types (canopy tree distance and leafcutter ant trails) was associated with changes in forager number. In at least two ant species, food presentation affected monopoly at baits; among all resource-defending ants, the microhabitats where ants foraged for food and the type of food located determined in part the frequency of monopoly and the number of foragers at the food item. These results suggest that the location and presentation of food items determines in part which ant species will utilize the resource.

Animals↗

Ant species confer different partner benefits on two neotropical myrmecophytes.

The dynamics of mutualistic interactions involving more than a single pair of species depend on the relative costs and benefits of interaction among alternative partners. The neotropical myrmecophytes Cordia nodosa and Duroia hirsuta associate with several species of obligately symbiotic ants. I compared the ant partners of Cordia and Duroia with respect to two benefits known to be important in ant-myrmecophyte interactions: protection against herbivores provided by ants, and protection against encroaching vegetation provided by ants. Azteca spp., Myrmelachista schumanni, and Allomerus octoarticulatus demerarae ants all provide the leaves of Cordia and Duroia some protection against herbivores. However, Azteca and Allomerus provide more protection than does Myrmelachista to the leaves of their host plants. Although Allomerus protects the leaves of its hosts, plants occupied by Allomerus suffer more attacks by herbivores to their stems than do plants occupied by other ants. Relative to Azteca or Allomerus, Myrmelachista ants provide better protection against encroaching vegetation, increasing canopy openness over their host plants. These differences in benefits among the ant partners of Cordia and Duroia are reflected in the effect of each ant species on host plant size, growth rate, and reproduction. The results of this study show how mutualistic ant partners can differ with respect to both the magnitude and type of benefits they provide to the same species of myrmecophytic host.

Analysis of Variance↗