Who is the top dog in ant communities? Resources,parasitoids, and multiple competitive hierarchies

A wide variety of animal communities are organized into interspecific dominance hierarchies associated with the control and harvest of food resources. Interspecific dominance relationships are commonly found to be linear. However, dominance relations within communities can form a continuum ranging from intransitive networks to transitive, linear dominance hierarchies. How interference competition affects community structure depends on the configuration of the dominance interactions among the species. This study explores how resource size and the trait-mediated indirect effect (TMIE) specialist phorid fly parasitoids exert on interference competition, affect the transitive nature of competitive interactions in an assemblage of woodland ants. I quantify the linearity of networks of interactions associated with large and small food resources in the presence and absence of phorid parasitoids. Two distinct, significantly linear dominance hierarchies exist within the ant assemblage depending on the size of the disputed resource. However, the presence of phorid fly parasitoids eliminates the linearity of both dominance hierarchies. The hosts phorid defense behaviors reduce the competitive asymmetries between the host and its subdominant competitors, increasing the indeterminacy in the outcome of competitive interactions. Thus, both resource size variation and phorid-induced TMIEs appear to facilitate coexistence in assemblages of scavenging ants.     

Foraging ecology and patterns of diversification in dipteran parasitoids of fire ants in south Brazil

1. At least sixteen species of parasitoid flies in the genus Pseudacteon (family Phoridae) attack fire ants in the Solenopsis saevissima subcomplex in South America. Little is known of behavioural or ecological differences among Pseudacteon parasitoids of fire ants, although their coexistence in multispecies communities would suggest that important differences exist. Seven Pseudacteon species in two separate communities were studied in south-east Brazil. The way in which hosts detect and respond to the presence of parasitoids, attack rates of the parasitoids, and host location behaviour of the parasitoids were examined.
2. Reductions in fire ant recruitment were more closely related to the number of ants attacked along a foraging trail than to the amount of time that a phorid was present.
3. Pseudacteon solenopsidis differed from other phorid species by flying backwards while pursuing ants, by attacking at lower rates than other phorids, and by spending longer around fire ant foraging trails than other phorids before departing. Fire ant recruitment to food often rebounded in the continued presence of P. solenopsidis.
4. In each of the two communities, certain Pseudacteon species appeared frequently at Solenopsis foraging trails, whereas others appeared predominantly at mound disturbances. Two distinct size classes of phorids were present in each community, and the community with the larger ant host species also had a third and larger phorid species. No phorid species from the same community had similar body sizes and similar host location behaviours, although numerous species from different communities shared both of these traits.
5. Heterogeneity in host size and in the ecological circumstances under which hosts are vulnerable to attack appears to have influenced the evolution and perhaps maintenance of diverse Pseudacteon communities.     

Habitat disturbance modifies dominance,coexistence, and competitive interactions in tropical ant communities

1. Interspecific competition is a major structuring principle in ecological communities. Despite their prevalence, the outcome of competitive interactions is hard to predict, highly context-dependent, and multiple factors can modulate such interactions. 2. We tested predictions concerning how competitive interactions are modified by anthropogenic habitat disturbance in ground-foraging ant assemblages inhabiting fragmented Inter-Andean tropical dry forests in southwestern Colombia, and investigated ant assemblages recruiting to baits in 10 forest fragments exposed to varying level of human disturbance. 3. Specifically, we evaluated how different components of competitive interactions (patterns of species co-occurrence, resource partitioning, numerical dominance, and interspecific trade-offs between discovery and dominance competition) varied with level of habitat disturbance in a human-dominated ecosystem. 4. Multiple lines of evidence suggest that the role of competitive interactions in structuring ground-foraging ant communities at baits varied with respect to habitat disturbance. As disturbance increased, community structure was more likely to exhibit random co-occurrence patterns, higher levels of monopolization of food resources by dominant ants, and disproportionate dominance of a single species, the little fire ant (Wasmannia auropunctata). At a regional scale, we found evidence for a trade-off between dominance and discovery abilities of the 15 most common species at baits. 5. Together, these results suggest that human disturbance modifies the outcome of competitive interactions in ground-foraging ant assemblages and may promote dominant species that reduce diversity and coexistence in tropical ecosystems.     

On testing the competition-colonization trade-off in a multispecies assemblage

The competition-colonization trade-off has long been considered an important mechanism explaining species coexistence in spatially structured environments, yet data supporting it remain ambiguous. Most competition-colonization research examines plants and the dispersal-linked traits of their seeds. However, colonization is more than just dispersal because rapid population growth is also an important component of colonization. We tested for the presence of competition-colonization trade-offs with a commonly used artificial assemblage consisting of protozoan and rotifer species, where colonization was the ability of a species to establish populations in patches. By ranking species according to their colonization abilities and their pairwise competitive interactions, we show that these species exhibit competition-colonization trade-offs. These results reveal that the competition-colonization trade-off exists within nonplant assemblages and that even in a laboratory setting, species are constrained to be either good competitors or colonizers but not both.     

Parasite Lost: Chemical and Visual Cues Used by <Emphasis Type="BoldItalic">Pseudacteon</Emphasis> in Search of <Emphasis Type="BoldItalic">Azteca instabilis</Emphasis>

An undescribed species of phorid fly (genus: Pseudacteon) parasitizes the ant Azteca instabilis F Smith, by first locating these ants through the use of both chemical and visual cues. Experiments were performed in Chiapas, Mexico to examine a) the anatomical source of phorid attractants, b) the specific chemicals produced that attract phorids, and c) the nature of the visual cues used by phorids to locate the ants. We determined that phorid-attracting chemicals were present within the dorsal section of the abdomen, the location of the pygidial gland. Further experiments indicate that a pygidial gland compound, 1-acetyl-2-methylcyclopentane, is at least partially responsible for attracting phorid flies to their host. Finally, although visual cues such as movement were important for host location, size and color of objects did not influence the frequency with which phorids attacked moving targets.     

Are Trade-Offs Among Species’ Ecological Interactions Scale Dependent? A Test Using Pitcher-Plant Inquiline Species

Trade-offs among species' ecological interactions is a pervasive explanation for species coexistence. The traits associated with trade-offs are typically measured to mechanistically explain species coexistence at a single spatial scale. However, species potentially interact at multiple scales and this may be reflected in the traits among coexisting species. I quantified species' ecological traits associated with the trade-offs expected at both local (competitive ability and predator tolerance) and regional (competitive ability and colonization rate) community scales. The most common species (four protozoa and a rotifer) from the middle trophic level of a pitcher plant (Sarracenia purpurea) inquiline community were used to link species traits to previously observed patterns of species diversity and abundance. Traits associated with trade-offs (competitive ability, predator tolerance, and colonization rate) and other ecological traits (size, growth rate, and carrying capacity) were measured for each of the focal species. Traits were correlated with one another with a negative relationship indicative of a trade-off. Protozoan and rotifer species exhibited a negative relationship between competitive ability and predator tolerance, indicative of coexistence at the local community scale. There was no relationship between competitive ability and colonization rate. Size, growth rate, and carrying capacity were correlated with each other and the trade-off traits: Size was related to both competitive ability and predator tolerance, but growth rate and carrying capacity were correlated with predator tolerance. When partial correlations were conducted controlling for size, growth rate and carrying capacity, the trade-offs largely disappeared. These results imply that body size is the trait that provides the basis for ecological interactions and trade-offs. Altogether, this study showed that the examination of species' traits in the context of coexistence at different scales can contribute to our understanding of the mechanisms underlying community structure.     

The discovery-dominance trade-off is the exception, rather than the rule

1. Interspecific trade-offs are thought to facilitate coexistence between species at small spatial scales. The discovery-dominance trade-off, analogous to a competition-colonisation trade-off, is considered an important structuring mechanism in ant ecology. A trade-off between species' ability to discover food resources and to dominate them may explain how so many species apparently dependent on similar resources can coexist. 2. The discovery-dominance trade-off is thought to be broken by invasive species in enemy-free space or territorial species whose activity is fuelled by domination of carbohydrate resources. It may also be mediated by factors such as temperature and habitat structure. 3. We investigate the generality and form of the discovery-dominance relationship in an experiment using habitats of contrasting complexity across three continents. In addition, to assess how widespread the discovery-dominance trade-off is, we conducted a systematic review combining all empirical studies (published and from our experiment). 4. From our own fieldwork and meta-analyses of available studies, we find surprisingly little empirical support for the trade-off, with results indicating that mean effect sizes were either not significantly different from 0 or significantly positive. The trade-off was only detected in studies with parasitoids present. Additionally, experimental data from simple and complex habitats within each continent suggest that simple habitats may facilitate both food resource discovery and dominance. 5. We conclude that the discovery-dominance trade-off is the exception, rather than the rule. Instead, these abilities were commonly correlated. Real food resources provide many axes along which partitioning may occur, and discovery-dominance trade-offs are not a prerequisite for coexistence.     

Effects of parasitoids on a mycophagous drosophilid community in northern Japan and an evaluation of the disproportionate parasitism hypothesis

In a host–parasitoid system comprising mycophagous drosophilids and their parasitoids, the drosophilid and parasitoid species assemblages, host use, and the prevalence of parasitism were assessed, and the “disproportionate parasitism hypothesis” was examined with consideration given to yearly variations. The mycophagous drosophilids, their fungal food resources and parasitoids were studied by carrying out an intensive census throughout the activity seasons of 4 years (2000–2003) in Hokkaido, northern Japan. Five hymenopterous parasitoid species, four braconids and one eucoilid, were found. Parasitoids of mycophagous drosophilids are reported for the first time from Asia. Most parasitism (99.2%) was by braconids, in contrast to the dominance of eucoilids in Europe. Parasitism was restricted to the summer, and the rate was high from early July to early August every year. There was considerable yearly variation in the composition of abundant fungus, drosophilid and parasitoid species, especially between 2000 and 2001. The alternation of dominant host species was coupled with the alternation of dominant parasitoid species that differed in host use. Despite the yearly variation in the system, the most dominant host species suffered disproportionately heavy parasitism by the correspondingly dominant parasitoid species every year. The parasitism rate was positively correlated with the relative host abundance. This thus indicates that the disproportionate parasitism mechanism may operate, via which species coexistence is promoted by a higher rate of parasitism of the dominant species.     

Trade-offs and coexistence in microbial microcosms

Trade-offs among the abilities of organisms to respond to different environmental factors are often assumed to play a major role in the coexistence of species. There has been extensive theoretical study of the role of such trade-offs in ecological communities but it has proven difficult to study such trade-offs experimentally. Microorganisms are ideal model systems with which to experimentally study the causes and consequences of ecological trade-offs. In model communities of E. coli B and T-type bacteriophage, a trade-off in E. coli between resistance to bacteriophage and competitive ability is often observed. This trade-off can allow the coexistence of different ecological types of E. coli. The magnitude of this trade-off affects, in predictable ways, the structure, dynamics and response to environmental change of these communities. Genetic factors, environmental factors, and gene-by-environment interactions determine the magnitude of this trade-off. Environmental control of the magnitude of trade-offs represents one avenue by which environmental change can alter community properties such as invasability, stability and coexistence. This revised version was published online in August 2006 with corrections to the Cover Date.     

Life-history trade-offs and ecological dynamics in the evolution of longevity

Longevity is a life-history trait that is shaped by natural selection. An unexplored consequence is how selection on this trait affects diversity and diversification in species assemblages. Motivated by the diverse rockfish (Sebastes) assemblage in the North Pacific, the effects of trade-offs in longevity against competitive ability are explored. A competition model is developed and used to explore the potential for species diversification and coexistence. Invasion analyses highlight that life-history trait trade-offs in longevity can mitigate the effects of competitive ability and favour the coexistence of a finite number of species. Our results have implications for niche differentiation, limiting similarity and assembly dynamics in multispecies interactions.     

Interactions between ant species increase arrival rates of an ant parasitoid

Behaviours or traits associated with aggression and communication may increase an animal's conspicuousness to predators or parasitoids. Most examples of this come from instances of aggression or communication within a species. We tested whether interspecific encounters between ants enhance the host location success of a parasitoid (Diptera: Phoridae) that attacks ants in the genus Linepithema. At food resources recruited to by Linepithema, parasitoid discovery rates were lower when Linepithema was alone than when other ant species were present. In experimentally controlled encounters, parasitoid discovery rates were elevated when Linepithema confronted an ant species that elicited use of chemicals, but not when it confronted an ant species that primarily elicited physical aggression. These results indicate that phorid parasitoids of Linepithema use the ant's chemicals as host location cues. Because Linepithema is known to abandon food resources in the presence of its phorid parasitoids, its use of chemicals during interspecific encounters may diminish its competitive success when phorids are nearby. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Comparing electroantennogram and behavioral responses of two Pseudacteon phorid fly species to body extracts of Black, Red and Hybrid imported fire ants, Solenopsis spp

Several phorid fly species were introduced to the southern United States for biological control of the invasive imported fire ants, Solenopsis richteri (Black), Solenopsis invicta (Red), and their Hybrid S. richteri×S. invicta (Hybrid). It has been previously reported that the Jaguariuna biotype of Pseudacteon tricuspis and the Formosan biotype of Pseudacteon curvatus could distinguish among the three fire ant species with greater preference for Hybrid and Red fire ants. We hypothesized that phorid flies might use host derived chemical cues to differentiate ant species. To determine possible differential olfactory sensitivity of phorid fly species to different fire ant species, we compared electroantennogram (EAG) and behavioral responses of both sexes of P. tricuspis and P. curvatus to body extracts of Black, Red and Hybrid fire ants. As worker sizes of Black and Hybrid fire ants used in this study were much larger than that of Red fire ant (the average weight for Black, Red and Hybrid workers was 1.707, 0.863, 1.223mg per ants, respectively), at doses of 0.01, 0.1, 1 worker equivalent, body extracts of Black and Hybrid fire ant elicited significantly greater EAG response in both sexes of P. tricuspis than that of Red fire ant. Similarly, the EAG response in female P. curvatus to body extract of Black fire ant was significantly greater than to body extract of Red fire ant. To eliminate worker size influence on EAG response in phorid flies, we conducted a second EAG study using a dose of 1mg ant equivalent (body extract from 1mg of worker). No difference in EAG responses was recorded to body extract obtained from the same amount of workers among the three fire ant species (we consider viable Hybrid fire ant as a species in this paper), suggesting that worker size differences contributed to difference in EAG response in the first EAG study. In both EAG studies, male P. tricuspis showed significantly greater EAG responses than male P. curvatus to all three fire ant species. In four-way olfactometer bioassay, worker body extracts of all three fire ant species were equally attractive to P. tricuspis and P. curvatus (i.e. both phorid fly species did not show any preferences among the three fire ant species). Together, the results of the EAG and behavior studies suggest that parasitic phorid flies utilize host derived non-polar compounds from worker ants extracted out by hexane for host location but not for host preference, since both fly species are not able to distinguish among the body extracts of the three fire ant species. Future study will investigate possible involvement of polar compounds and/or non-chemical cues in mediating host preference by phorid flies.     

Dominant ants can control assemblage species richness in a South African savanna

1. Competition is considered a key factor structuring many communities, and has been described as the 'hallmark' of ant ecology. Dominant species are thought to play a key role structuring local ant assemblages through competitive exclusion. 2. However, while there have been many studies demonstrating competitive exclusion and consequently reduced richness at baits, it is not clear whether such regulation of 'momentary' diversity at clumped food resources can scale up to the regulation of richness at the site or assemblage level. 3. In this study, ant assemblages were sampled in three different savanna habitats in South Africa using both baiting and pitfall trapping. 4. As has been found in previous studies, there was a unimodal relationship between dominant ants and species richness at baits, with high abundances of dominant ants regulating species richness through competitive exclusion. Analysis of pitfall samples revealed strong convergence in pattern, and results from null model co-occurrence analyses supported the findings. 5. The importance of competition in structuring local ant assemblages was, however, only apparent at one of the three savanna habitats suggesting that a full range of extreme environments is needed to produce the full unimodal relationship at the assemblage level. 6. Although the relative importance of competition varied with habitat type, the study demonstrated that in some habitats, dominant ants can control species richness at the assemblage level.     

Parasitoid–host relationships when host size varies: the case of Pseudacteon flies and Solenopsis fire ants

Abstract. 1. Phorid flies in the genus Pseudacteon are parasitoids of ants. Variation in host size preferences of four South American and two North American Pseudacteon species on monogyne and polygyne forms of their host Solenopsis species [ S. invicta Buren and S. geminata (F.), respectively] was documented.
2. Monogyne Solenopsis workers were, on average, significantly larger than polygyne workers, and the average size monogyne worker attacked was significantly larger than the average size polygyne worker attacked in four of the six Pseudacteon species.
3. Three South American Pseudacteon species attacked larger than average size workers, whereas one attacked smaller than average size workers, in both monogyne and polygyne forms. Both North American Pseudacteon species attacked larger than average size polygyne workers and smaller than average size monogyne workers.
4. Three Pseudacteon species were reared from eggs to adults in infected ants in the laboratory. The size of the emergent phorid fly was related positively to the size of the host worker ant, with females emerging from larger hosts. Similar patterns were documented for both monogyne and polygyne forms.
5. The mean size of worker host from which phorids emerged did not differ significantly between the monogyne and polygyne forms in the subsample of phorids reared to adults.
6. The observed patterns elucidate factors that may cause variation in Pseudacteon sex ratios, and have implications for biological control efforts of pest Solenopsis species.     

Environmental context alters ecological trade‐offs controlling ant coexistence in a spatially heterogeneous region

1. Ecological trade‐offs in ant (Hymenoptera: Formicidae) assemblages and their implications for coexistence boast a rich history in entomology. Yet investigations of trade‐offs have largely been limited to homogeneous environments. We examined how environmental context modifies trade‐off expression in an ant assemblage spanning a heterogeneous region in central Florida, U.S.A. 2. We examined how trade‐off expression is altered among two contrasting habitat types: open shrub and forest. We tested for the presence of the dominance‐discovery trade‐off and two dominance‐thermal tolerance trade‐offs by estimating behavioral dominance, discovery ability, and thermal tolerance (foraging thermal limit, lethal temperature, and maximal abundance temperature) for a wide range of interacting ant species. 3. We found significantly linear dominance hierarchies in both shrub and forest habitats, showing dominant species out‐compete subordinates for food resources. In thermally stressful shrub habitats, subordinates exhibit higher thermal tolerances, take greater thermal risks, and reach maximum forager abundances at higher temperatures than do dominant species. This suggests temperature mediated trade‐offs control coexistence in shrub habitat. In thermally moderate forest habitat, we found limited evidence for trade‐offs between competitive dominance and resource discovery or between dominance and thermal traits, implying other processes control coexistence. These results demonstrate that trade‐offs controlling ant coexistence may be contingent on environmental context.     

Assembly rules for functional groups of North American shrews: effects of geographic range and habitat partitioning

Trophic cascades exist in numerous terrestrial systems, including many systems with ants as the top predator. Many studies show how behavioral modifications of herbivores are especially important in mediating species interactions and trophic cascades. Although most studies of trophic cascades focus on predator-herbivore-plant links, the trophic cascades concept could be applied to almost any level of trophic interactions. Especially considering the importance of parasites we consider here the interactions between the parasitic phorid fly, Pseudacteon sp. (Diptera: Phoridae), its ant host, Azteca instabilis (F. Smith) (Hymenoptera: Formicidae), and the herbivore, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) in the coffee agroecosystem. We investigated the effects of phorid flies on ant behavior by monitoring ant recruitment to tuna baits over a 30-min period in the presence or absence of phorid flies. To study the indirect effects of phorids on larvae, we placed baits on coffee plants to elevate ant foraging levels to levels near to ant nests, placed larvae near baits, and recorded the effects of ants on larvae either in the presence or absence of phorid flies. We found that phorid fly presence significantly reduced ant ability to recruit to baits through behavioral modifications and also significantly lessened ant ability to attack, carry away, or force herbivores off plants. We conclude there is a behaviorally-modified species-level trophic cascade in the coffee agroecosystem, with potentially important effects in ant and herbivore communities as well as for coffee production.     

Host specialist clownfishes are environmental niche generalists

Why generalist and specialist species coexist in nature is a question that has interested evolutionary biologists for a long time. While the coexistence of specialists and generalists exploiting resources on a single ecological dimension has been theoretically and empirically explored, biological systems with multiple resource dimensions (e.g. trophic, ecological) are less well understood. Yet, such systems may provide an alternative to the classical theory of stable evolutionary coexistence of generalist and specialist species on a single resource dimension. We explore such systems and the potential trade-offs between different resource dimensions in clownfishes. All species of this iconic clade are obligate mutualists with sea anemones yet show interspecific variation in anemone host specificity. Moreover, clownfishes developed variable environmental specialization across their distribution. In this study, we test for the existence of a relationship between host-specificity (number of anemones associated with a clownfish species) and environmental-specificity (expressed as the size of the ecological niche breadth across climatic gradients). We find a negative correlation between host range and environmental specificities in temperature, salinity and pH, probably indicating a trade-off between both types of specialization forcing species to specialize only in a single direction. Trade-offs in a multi-dimensional resource space could be a novel way of explaining the coexistence of generalist and specialists.     

Exploitative competition and coexistence in a parasitoid assemblage

Most insect populations are exploited by a complex of different parasitoid species, providing ample opportunities for competitive interactions among the latter. Despite this, resource-mediated competition (i.e., exploitative competition) among insect parasitoids remains poorly documented in natural systems. Here we propose a novel way to infer the presence of competitive interactions from covariance patterns in parasitism levels, and illustrate the use of this approach on a relatively well-defined and simple host–parasitoid system. The parasitism levels caused by three parasitoid species on a shared host showed a highly consistent negative covariance among samples. With the levels of parasitism by one species increasing, the levels of parasitism attributable to the two others decreased. Importantly, negative covariance between parasitism levels by different species appeared at high abundance, but not at low abundance of the phenologically earlier parasitoid species. This as well as several other lines of evidence indicates the importance of competitive interactions in this system. Feeding biology and phenology of the parasitoids suggest that competition in this parasitoid assemblage is primarily resource-mediated rather than occurring through direct interference. The species attacking earlier stages of the host are competitively superior to those attacking their host later in the season. Better dispersal ability and use of alternative host species by the inferior species could contribute to the coexistence of these competing parasitoids.     

Spatial and temporal variation in host–parasitoid interactions: leafcutter ant hosts and their phorid parasitoids

1. Parasitoid–host interactions are important components of ecological communities. Although parasitoid–host interactions are strongly shaped by evolutionary history, the abundance of both the parasitoid and the host may have a role in determining the nature of the interaction once phylogenetic relationships are considered. 2. Leafcutter ants are hosts of phorid parasitoids and represent a well‐defined and specialised module within a larger network of ant–symbiont interactions. A low specificity host taxa and a positive association between host abundance and parasitoid interaction frequency were expected due to the close phylogenetic relatedness of the hosts. 3. The interactions among all species of leafcutter ants and their parasitoids were quantified in two localities with different species richness. This study also characterised the spatial‐temporal variability of these interactions, determined the patterns of parasitoid specificity and host selection, and tested for an association between host abundance and parasitoid interaction frequency. 4. Contrary to expectation, most parasitoid species were highly specialised and interaction frequency for parasitoid species was not related to host abundance. All host ant species were attacked by more than one phorid species. Some phorid species used more than one host species and showed preference for the same species over space and time, suggesting that there are physiological and/or behavioural restrictions on host use. 5. These results show that there is a tendency for specialisation even when hosts are highly similar in their ecology. From a biological control perspective, these parasitoids may be effective candidates, due to the high specificity of some species and little host‐use variation through time.     

Resisting annihilation: relationships between functional trait dissimilarity,assemblage competitive power and allelopathy

Allelopathic species can alter biodiversity. Using simulated assemblages that are characterised by neutrality, lumpy coexistence and intransitivity, we explore relationships between within‐assemblage competitive dissimilarities and resistance to allelopathic species. An emergent behaviour from our models is that assemblages are more resistant to allelopathy when members strongly compete exploitatively (high competitive power). We found that neutral assemblages were the most vulnerable to allelopathic species, followed by lumpy and then by intransitive assemblages. We find support for our modeling in real‐world time‐series data from eight lakes of varied morphometry and trophic state. Our analysis of this data shows that a lake's history of allelopathic phytoplankton species biovolume density and dominance is related to the number of species clusters occurring in the plankton assemblages of those lakes, an emergent trend similar to that of our modeling. We suggest that an assemblage's competitive power determines its allelopathy resistance.