Competitive dominance in the organisation of Mediterranean ant communities

1. Competition by dominant species is thought to be key to structuring ant communities. However, recent findings suggest that the effect of dominant species on communities is less pronounced than previously assumed. 2. The aim of the present study was to identify the role of dominant ants in the organisation of Mediterranean communities, particularly the role of competition in invaded and uninvaded communities. The effects on ant assemblages of two dominant ants, the invasive Argentine ant and the native ant, Tapinoma nigerrimum Nylander, were assessed. 3. The abundances of both dominant ants were significantly correlated with a decrease in native ant richness at traps. However, only the invasive ant was associated with a reduction in diversity and abundance of other ant species at site scale. In the presence of T. nigerrimum, species co‐occurrence patterns were segregated or random. Community structure in both the dominant‐free and the Argentine ant sites showed random patterns of species co‐occurrence. 4. The present findings indicate that dominant ants regulate small‐scale diversity by competition. However, at the broader scale of the assemblage, T. nigerrimum may only affect species distribution, having no apparent effect on community composition. Moreover, we find no evidence that inter‐specific competition shapes species distribution in coastal Mediterranean communities free of dominant ants. 5. These results show that dominant species may affect ant assemblages but that the nature and the intensity of such effects are species and scale dependent. This confirms the hypothesis that competitive dominance may be only one of a range of factors that structure ant communities.     

Invasive African big-headed ants, Pheidole megacephala, on coral cays of the southern Great Barrier Reef: distribution and impacts on other ants

Infestation of islands by exotic ants is widespread and increasing due to human activities throughout the world. Exotic ants, particularly the invasive African big-headed ant, Pheidole megacephala (Fabricius), are of great conservation concern for coral cays at the southern end of the Great Barrier Reef, Australia. Little is known, however, about the distribution and ecological impacts of invasive ants in this insular system. We surveyed the ants of 14 vegetated coral cays recording a total of 24 ant species, including at least nine exotics. Pheidole megacephala was by far the most abundant and widespread species, occurring on 11 of 14 islands, often in very large numbers. The inter-island distribution of P. megacephala was best explained by human activities, with frequently visited, and to a lesser degree disturbed islands, more likely to be infested. On large islands (≥?10?ha) P. megacephala exhibited distinct habitat preferences, occurring in significantly lower abundances within heavily-shaded Pisonia grandis forest in the centre of islands, compared to more open, fringing woodland or shrubland. On smaller islands (<10?ha) with less extensive Pisonia stands, P. megacephala penetrated throughout the forest where its abundance was similar to that in open woodland. Despite considerable differences in biotic (floristic composition) and abiotic factors (e.g. island size) as well as the spatial configuration among islands, the severity of infestation by P. megacephala best explained variation in species richness, abundance and assemblage composition of other ants. We suggest a number of strategies to manage P. megacephala infestations on these islands.     

Dominance in a ground‐dwelling ant community of banana agroecosystem

In tropical ecosystems, ants represent a substantial portion of the animal biomass and contribute to various ecosystem services, including pest regulation and pollination. Dominant ant species are known to determine the structure of ant communities by interfering in the foraging of other ant species. Using bait and pitfall trapping experiments, we performed a pattern analysis at a fine spatial scale of an ant community in a very simplified and homogeneous agroecosystem, that is, a single‐crop banana field in Martinique (French West Indies). We found that the community structure was driven by three dominant species (Solenopsis geminata, Nylanderia guatemalensis, and Monomorium ebeninum) and two subdominant species (Pheidole fallax and Brachymyrmex patagonicus). Our results showed that dominant and subdominant species generally maintained numerical dominance at baits across time, although S. geminata, M. ebeninum, and B. patagonicus displayed better abilities to maintain dominance than P. fallax and N. guatemalensis. Almost all interspecific correlations between species abundances, except those between B. patagonicus and N. guatemalensis, were symmetrically negative, suggesting that interference competition prevails in this ground‐dwelling ant community. However, we observed variations in the diurnal and nocturnal foraging activity and in the daily occurrence at baits, which may mitigate the effect of interference competition through the induction of spatial and temporal niche partitioning. This may explain the coexistence of dominant, subdominant, and subordinate species in this very simplified agroecosystem, limited in habitat structure and diversity.     

Plants feed ants: food bodies of myrmecophytic Piper and their significance for the interaction with Pheidole bicornis ants

Several species of Piper (Piperaceae) live in symbiosis with Pheidole bicornis (Formicidae-Myrmicinae) on the southern Pacific slope of Costa Rica. These plants produce small single-celled food bodies (FBs) in leaf domatia, formed by the petiole bases and roofing leaf sheaths. In the present study the dependency of ants on FBs of Piper fimbriulatum as a food source was analysed by comparing the natural abundance of ¹³C and ¹⁵N in ants and FBs. Both '¹³C and '¹⁵N values were very similar between FBs and Pheidole bicornis ants but differed substantially between the plant and other ant species. Therefore we suggest that FBs are a main food source for Pheidole bicornis ants. To strengthen this suggestion, the chemical composition of FBs of four myrmecophytic Piper species was analysed, with special emphasis on the nutritional requirements of inhabiting Pheidole bicornis ants. Standard chemical methods were modified and combined to a novel analysis scheme by which all major FB constituents could be quantified from minute [3-10 mg dry mass (DM)] quantities. Piper FBs mainly consisted of lipids (41-48% of DM) and proteins (17-24% of DM). Soluble carbohydrates and amino acids proved to be quantitatively unimportant. N was predominantly stored as soluble protein and, thus, was easily available to the ants. FBs proved to be a high-energy food source (up to 23 kJ g^-1 DM), with a chemical composition that meets well the nutritional needs of the inhabiting ants.     

Ants as Bioindicators of Natural Succession in Savanna and Riparian Vegetation Impacted by Dredging in the Jequitinhonha River Basin,Brazil

We investigated the response of ant species to landscape and geomorphologic parameters of a long‐term (7–11 years) restoration project in the Jequitinhonha River (Northern State of Minas Gerais, Brazil) margins, previously dredged by a diamond mining company. Geomorphological changes from the dredging were severe and the area is unlikely to be adequately restored, mainly due to the negative effects of flooding. Our hypothesis is that ant species assemblages bioindicate successional stages and soil characteristics. We studied the association of effects from the river's flooding zone, the native vegetation, and sedimentary grain size with that of ant species diversity, abundance, and composition. An ant sampling program was conducted in April 2005, using three methods: baits, pitfall traps, and direct collection. Grain size was measured by sieving. In total, 10,784 ants were sampled, belonging to 7 subfamilies, 24 genera and 45 morphospecies. Ant species richness was greater in the undisturbed savanna area than in the restored habitats, and equivalently greater in the ecotone and intermediate zone habitats than on the river bank, the poorest habitat. Atta sexdens rubropilosa indicated a condition related to small forest remnants having well‐structured soil. On the other hand, ants with a body length of under 0.5 cm (Dorymyrmex pyramicus and Pheidole fallax) predominated in sandy areas, where the majority of the granules were the finest. The lack of organic matter and soil structure for constructing suitable nests may prevent large ants from colonizing such areas, and thus inhibit the advance of natural succession.     

Biodiversity responses to vegetation structure in a fragmented landscape: ant communities in a peri-urban coastal dune system

Habitat fragmentation often results in significant degradation of the structure and composition of remnant natural vegetation, leading to substantial biodiversity decline. Ants are an ecologically dominant faunal group known to be sensitive to vegetation degradation following fragmentation. We examined ant diversity and composition in relation to changes in vegetation structure in remnant coastal vegetation in the global biodiversity hotspot of southwestern Western Australia. The key features of vegetation structure driving the species and functional diversity and composition of ant communities were measures of cover of vegetation and bare ground. However, these effects were highly idiosyncratic at the species level. Cluster analyses based on plant species composition classified plots into two groups corresponding to relatively intact and degraded vegetation respectively. Although systematic changes in plant diversity and vegetation structure were observed between the two groups, key features from an ant perspective (native plant cover and bare ground) remained unchanged. Vegetation degradation consequently had little overall effect on ant species composition and functional diversity. The major disturbance–related impact on ant communities was through invasion by exotic ants, especially Pheidole megacephala; however, this occurred only in close proximity to development. Our results suggest that the priority for conserving ant diversity in our coastal dune system is the prevention of invasion by exotic species.     

Impact of predatory ants on two lepidopteran insect pests in Indian cauliflower agroecosystems

Abstract:  The impact of predatory, agroecosystem-nesting ant species on the larval stages of the two major lepidopteran pests, the diamondback moth (DBM), Plutella xylostella and the black cutworm (BC), Agrotis ipsilon , in cauliflower agroecosystem in Uttar Pradesh, India, was investigated in farmers' fields. Field observations revealed that six ant species, the most important being Pheidole sp., preyed on the crop-damaging stages of DBM and BC. Bait experiments with larvae of DBM and BC, placed randomly on the ground, demonstrated significant active retrieval by Pheidole sp. Investigations on larval density of both the insect pests on cauliflower crops, located within 1–5 m distances from nests of Pheidole sp., revealed a significant increase in the level of infestation of the crop plants with increasing distance from the ant nests. While a significant negative correlation was found between infested plants with low larval densities, a positive correlation was found between plants with high larval densities with regard to the distance of crop plant location from the ant nest. Ant-inclusion and -exclusion experiments under field conditions further confirmed the role of predatory ants, with Pheidole sp. having the major impact in the direct retrieval of the lepidopteran larvae from the crop plants. A significant difference was found in the leaf area damaged by BC larvae between the ant-included and -excluded potted plants during the 6-h observation period. Thus, this study provides direct evidence of the role of ants in actively searching and directly preying on the larval stages of DBM and BC, on cauliflower plants. Moreover, we demonstrate that in ephemeral, annual cropping agroecosystems, predatory ants are important natural enemies of lepidopteran pests, playing a significant role in plant protection.     

Ants in a hospital environment and its importance as vector of bacteria

The external ant community of Hospital Municipal de Morrinhos, in Morrinhos, Goiás State, was characterized by the low rates of richness, diversity, dominance and equity of species abundance. Pheidole sp.1, a polygynic species was numerically dominant in this environment, although it coexists with potentially competitive species. This ant species prevailed within all hospital departments and its space-time distribution was a little aggregated (variance/mean ratio = 1.102, chi2 = 29.38, P < 0.01). Escherichia, Salmonella, Aeromonas, Enterococcus, Staphylococcus and Klebsiella were the bacteria associated to this ant species in nearly all hospital annexes. The unicolonialism of Pheidole sp.1 tends to increase the contamination and dissemination process of infecto-contagious agents. The control and management of this ant species must be followed by practices that reduce the colonization process by other queens and the quantity of site nidification within the hospital.     

Geographic and taxonomic distribution of a positive interaction: ant-tended homopterans indirectly benefit figs across southern Africa

Although species pairs and assemblages often occur across geographic regions, ecologists know very little about the outcome of their interactions on such large spatial scales. Here, we assess the geographic distribution and taxonomic diversity of a positive interaction involving ant-tended homopterans and fig trees in the genus Ficus. Previous experimental studies at a few locations in South Africa indicated that Ficus sur indirectly benefited from the presence of a homopteran (Hilda patruelis) because it attracted ants (primarily Pheidole megacephala) that reduced the effects of both pre-dispersal ovule gallers and parasitoids of pollinating wasps. Based on this work, we evaluated three conditions that must be met in order to support the hypothesis that this indirect interaction involves many fig species and occurs throughout much of southern Africa and Madagascar. Data on 429 trees distributed among five countries indicated that 20 of 38 Ficus species, and 46% of all trees sampled, had ants on their figs. Members of the Sycomorus subgenus were significantly more likely to attract ants than those in the Urostigma subgenus, and ant-colonization levels on these species were significantly greater than for Urostigma species. On average, each ant-occupied F.sur tree had 37% of its fig crop colonized by ants, whereas the value was 24% for other Ficus species. H. patruelis was the most common source for attracting ants, although figs were also attacked by a range of other ant-tended homopterans. P. megacephala was significantly more common on figs than other ant species, being present on 58% of sampled trees. Ant densities commonly exceeded 4.5 per fig, which a field experiment indicated was sufficient to provide protection from ovule gallers and parasitoids of pollinators. Forty-nine percent of all colonized F. sur trees sampled had ant densities equal to or greater than 4.5 per fig, whereas this value was 23% for other Ficus species. We conclude that there is considerable evidence to suggest that this indirect interaction occurs across four southern African countries and Madagascar, and involves many Ficus species. Received: 11 December 1997 / Accepted: 6 April 1998     

Interactions of the ant Crematogaster scutellaris with the fig/fig wasp mutualism

Abstract. 1. A classical example of specialised pollination mutualisms is the relationship between fig trees and their pollinating wasps, in which each partner depends completely on the other for its reproduction; however the fig/fig wasp association is also the target of a great diversity of other species, ranging from specialised parasites to opportunistic foragers, among them ants.
2. The ant community and the sources of ant attraction observed on the Mediterranean fig tree Ficus carica were characterised.
3. A guild of ants attracted by homopterans tended on the plant was distinguished from a second guild composed of two co-dominant ant species ( Crematogaster scutellaris and Pheidole pallidula ) that prey mostly on pollinating wasps, abundant during certain parts of the fig cycle.
4. Foraging workers of C. scutellaris search for prey on the fig inflorescence (syconium), capturing pollinating wasps mostly at the peak of wasp emergence and at a rate estimated to reach 600 prey per day for an entire tree.
5. Detailed study of the predatory sequences displayed under experimental conditions showed that ant workers captured 100% of the pollinating wasps offered, while they captured only 5.5% of the parasitoid wasp specific to the pollinator. The respective impacts of the interaction on ants and on the figs are discussed, as well as several behavioural traits of predation by the dominant ant on pollinators.     

The role of resource dispersion in promoting the co‐occurrence of dominant and subordinate ant species

Dominant competitors govern resource use in many communities, leading to predictions of local exclusion and lower species diversity where dominant species are abundant. However, subordinate and dominant species frequently co‐occur. One mechanism that could facilitate resource sharing and co‐occurrence of dominant and subordinate competitors is fine‐scale resource dispersion. Here, we distributed 6 g of a food resource into 1, 2, 8, 32 or 64 units in small 0.40 m² areas centred on nests of the dominant ant Monomorium sydneyense. We tested three hypotheses. First, we hypothesized that the species richness and abundance of foraging ants would increase with increasing resource dispersion. Accordingly, species richness doubled and total ant abundance was two orders of magnitude higher in high resource dispersion treatments. Secondly, we hypothesized that increasing resource dispersion would reduce competitive interactions such as resource turnover events and lower the probability of food resources being occupied. Substantial support for this hypothesis was observed. Finally, we tested the hypothesis that the foraging time of each species would be proportional to the relative abundance of each species solely in high resource dispersion treatments. Expected and observed foraging times were statistically similar for only the dominant ant M. sydneyense. The subdominant Pheidole rugosula increased its foraging time much more than was expected, while two subordinate ants showed no relationship between observed and expected times. Thus, while increasing resource dispersion significantly increased overall species richness, this increase in co‐occurrence did not correlate with a significant increase in foraging time for the two subordinate species. Rather, changes in resource dispersion appeared to benefit only the subdominant species. Inter‐site variation appeared more important for other subordinate species indetermining co‐occurrence and foraging time. Multiple mechanisms facilitate co‐occurrence and resource sharing in this community, and probably in most other communities.     

Effects of prescribed fire on an ant community in Florida pine savanna

Abstract.  1. The effects of prescribed fire on ant community structure were examined in a regenerating longleaf pine savanna in Florida, U.S.A. The presence of ants on 20, 10 × 10 m plots was determined by baiting every 1–3 months from 18 months before a fire until 6 months afterwards.
2. Expected species richness (based on rarefaction) and species density 6 months post-fire were significantly lower than for the same month (September) 6 months before the fire.
3. Cluster analysis revealed that the effects of fire were far less important predictors of ant community structure than seasonality and unexplained inter-annual variation. Thus, overall, the impacts of fire were relatively minor and short term at the community level.
4. Different functional groups of ants (as defined by Andersen, 1997) responded to fire in strikingly different ways. Generalised Myrmicinae (e.g. Pheidole spp., Monomorium viride ) were affected more severely by fire than were the other functional groups. In contrast, the dominant Dolichoderinae ( Forelius pruinosus ) exhibited a large increase after the fire and seemed to be responsible for the decline in abundance of several species.
5. A strong negative correlation between F. pruinosus and other groups of ants immediately after the fire suggested more intense competition among ants at that time. Six months post-fire, the abundance of F. pruinosus decreased markedly and the abundance of other species rebounded.
6. The rapid post-fire recovery of the ant community probably reflects adaptations of ants to a chronic fire regime.     

Habitat complexity modifies ant–parasitoid interactions: implications for community dynamics and the role of disturbance

Species must balance effective competition with avoidance of mortality imposed by predators or parasites to coexist within a local ecological community. Attributes of the habitat in which species interact, such as structural complexity, have the potential to affect how species balance competition and mortality by providing refuge from predators or parasites. Disturbance events such as fire can drastically alter habitat complexity and may be important modifiers of species interactions in communities. This study investigates whether the presence of habitat complexity in the form of leaf litter can alter interactions between the behaviorally dominant host ants Pheidole diversipilosa and Pheidole bicarinata, their respective specialist dipteran parasitoids (Phoridae: Apocephalus sp. 8 and Apocephalus sp. 25) and a single species of ant competitor (Dorymyrmex insanus). We used a factorial design to manipulate competition (presence/absence of competitors), mortality risk (presence/absence of parasitoids) and habitat complexity (presence/absence of leaf litter). Parasitoid presence reduced soldier caste foraging, but refuge from habitat complexity allowed increased soldier foraging in comparison to treatments in which no refuge was available. Variation in soldier foraging behavior correlated strongly with foraging success, a proxy for colony fitness. Habitat complexity allowed both host species to balance competitive success with mortality avoidance. The effect of fire on habitat complexity was also studied, and demonstrated that the immediate negative impact of fire on habitat complexity can persist for multiple years. Our findings indicate that habitat complexity can increase dominant host competitive success even in the presence of parasitoids, which may have consequences for coexistence of subordinate competitors and community diversity in general.     

Herbivore damage increases avian and ant predation of caterpillars on trees along a complete elevational forest gradient in Papua New Guinea

Signals given off by plants to alert predators to herbivore attack may provide exciting examples of coevolution among organisms from multiple trophic levels. We examined whether signals from mechanically damaged trees (simulating damage by herbivores) attract predators of insects along a complete elevational rainforest gradient in tropical region, where various predators are expected to occur at particular elevational belts. We studied predation of artificial caterpillars on trees with and without ‘herbivorous’ damage; as well as diversity and abundances of potential predators at eight study sites along the elevational gradient (200–3700 m a.s.l.). We focused on attacks by ants and birds, as the main predators of herbivorous insect. The predation rate decreased with elevation from 10% d^?1 at 200 m a.s.l. to 1.8% d^?1 at 3700 m a.s.l. Ants were relatively more important predators in the lowlands, while birds became dominant predators above 1700 m a.s.l. Caterpillars exposed on trees with herbivorous damage were attacked significantly more than caterpillars exposed on trees without damage. Results suggest that relative importance of predators varies along elevational gradient, and that observed predation rates correspond with abundances of predators. Results further show that herbivorous damage attracts both ants and birds, but its effect is stronger for ants.     

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.     

Deterrent effect of a guild of extrafloral nectary-visiting ant species on Raphidopalpa foveicollis, a major insect pest of sponge gourd, Luffa cylindrica

Among the 10 ant species visiting the extrafloral nectaries of sponge gourd plants [ Luffa cylindrica L. (Cucurbitaceae)], Camponotus compressus (Fabricius) (Hymenoptera: Formicidae) appears to be the ecologically dominant species, due to its abundance and the high frequency of deterrent encounters with the major insect herbivore, Raphidopalpa foveicollis Lucas (Coleoptera: Chrysomelidae). A significant positive linear correlation was found between the number of extrafloral nectaries per leaf, calyx, bract, and bracteole and the patrolling time of C. compressus , Camponotus paria (Emery), Pheidole spec., and Tetramorium spec. (all Hymenoptera: Formicidae) at the corresponding plant parts. Many one-to-one deterrent interactions between the ant visitors and adult R. foveicollis were recorded, with C. compressus being involved in most encounters on the calyx and bracts, Pheidole spec. on the leaves, and Pachycondyla tesserinoda (Emery) (Hymenoptera: Formicidae) on the bracteoles. Tapinoma melanocephalum (Fabr.) (Hymenoptera: Formicidae) exhibited significant deterrent effects on adult herbivores on the corolla when in groups larger than 15 ants. Ant exclusion and inclusion experiments revealed a significant reduction in the residence time, as well as dose- and time-dependent reduction in the frequency of visits of the herbivore on crop plants with C. compressus , C. paria , Pheidole spec., and T. melanocephalum ants, compared to the control. Results support the optimal defence hypothesis, as the plant structures most closely linked to plant fitness bear the extrafloral nectaries and are defended by the visiting ant species. This is a valuable, indirect plant-protection strategy of an annual crop plant.     

The role of competitive dominance in the invasive ability of the Argentine ant (Linepithema humile)

To assess the importance of competition in the advance of invasive species, bait stations have been used to determine the dominance hierarchy of a community of native ants in Doñana National Park, southern Spain, and the status of the introduced species Linepithema humile (Argentine ant). Some native species, e.g. Cataglyphis floricola or Camponotus pilicornis, seem to be subordinate, i.e. occupy a low position in the competitive hierarchy; some are dominant (e.g. Pheidole pallidula), and others (e.g. Aphaenogaster senilis) occupy an intermediate position in the hierarchy. The Argentine ant is a competitively dominant species, because of its aggressive behavior and relative abundance. Irrespective of their position in the dominance hierarchy, L. humile and some native species adopt what games theory terms “the bourgeois strategy” during agonistic encounters with other species. Lone workers tend to be submissive in encounters whereas workers in the presence of other colony members are aggressive. L. humile was the only species which aggressively displaced large numbers of ants of other species from the bait. L. humile also expanded its range in the course of the experiment, displacing native species from parts of the study area.     

RETRACTED ARTICLE: Interspecific competition among ants in the boreal forest: Testing predictions from a linear hierarchical competition model

Competitive interactions often play an important role in local community structure, and particularly so in ant communities. We test predictions derived from a competition model proposed for Scandinavian ant communities by comparing ant nest densities inside and outside fifty randomly-selected territories of competitively dominant wood ants within 30 km of Ume?, northern Sweden. As predicted by the model, competitively-intermediate encounter species, as well as other territorial species, showed complementary occurrences with both studied wood ants, Formica aquilonia Yarrow and F. lugubris Zett.. In contrast, complementary abundances, i.e. lower density of competitively-inferior submissive species in the presence of wood ants, as predicted by the model, was not supported for all submissive species. Of the two studied wood ant species, the nest density of submissive species was negatively correlated only with F. aquilonia. Submissives as a group, as well as Myrmica-species, showed complementary abundances with F. aquilonia, but one Leptothorax and two Serviformica species had higher nest densities in the presence of this competitively-dominant species. We propose that, for Leptothorax, these deviations from the model predictions may be because of limited niche overlap with dominant wood ants, a small worker force and a timid behaviour which does not elicit aggression in wood ants. For the two Serviformica species, we propose a combination of protection against social parasites, inter-specific social control performed by F. aquilonia, and dominance relationships between competitively-inferior submissive species as reasons for the higher nest density inside F. aquilonia territories. Monogyny, and thereby smaller nests, lower worker force, smaller territory (as shown by this study) in F. lugubris, as compared to the polygyny in F. aquilonia, may also help explain the differences in their effects on subordinate species. Our analyses indicate that the linear competition hierarchy model proposed for Scandinavian ants accurately predicts the outcome of interaction between and community composition for dominant territorial and encounter species, but that it needs refinement with respect to the relationship between territorial and submissive species and the resulting community composition. Further studies are needed, especially addressing the complex relationships between these latter groups, and the effects of different competitively-dominant wood ants, to determine the mechanisms determining the outcome of these relationships and to more accurately predict community composition. Received 11 June 2007; revised 11 September 2007; accepted 17 September 2007.     

Dominance and species co-occurrence in highly diverse ant communities: a test of the interstitial hypothesis and discovery of a three-tiered competition cascade

The role of competitive exclusion is problematic in highly diverse ant communities where exceptional species richness occurs in the face of exceptionally high levels of behavioural dominance. A possible non-niche–based explanation is that the abundance of behaviourally dominant ants is highly patchy at fine spatial scales, and subordinate species act as insinuators by preferentially occupying these gaps—we refer to this as the interstitial hypothesis. To test this hypothesis, we examined fine-scale patterns of ant abundance and richness according to a three-tiered competition hierarchy (dominants, subdominants and subordinates) in an Australian tropical savanna using pitfall traps spaced at 2 m intervals. Despite the presence of gaps in the fine-scale abundance of individual species, the combined abundance of dominant ants (species of Iridomyrmex, Papyirus and Oecophylla) was relatively uniform. There was therefore little or no opportunity for subordinate species to preferentially occupy gaps in the foraging ranges of dominant species, and we found no relationship between the abundance of dominant ants and nondominant species richness at fine spatial scales. However, we found a negative relationship between subdominant and subordinate ants, a negative relationship between dominant and subdominant ants, and a positive relationship between dominant and subordinate ants. These results suggest that dominant species actually promote species richness by neutralizing the effects of subdominant species on subordinate species. Such indirect interactions have very close parallels with three-tiered trophic cascades in food webs, and we propose a “competition cascade” where the interactions are through a competition rather than trophic hierarchy.