Individual and interactive effects of chronic anthropogenic disturbance and rainfall on taxonomic, functional and phylogenetic composition and diversity of extrafloral nectary-bearing plants in Brazilian Caatinga.

Chronic anthropogenic disturbance (CAD) and climate change represent two of the major threats to biodiversity globally, but their combined effects are not well understood. Here we investigate the individual and interactive effects of increasing CAD and decreasing rainfall on the composition and taxonomic (TD), functional (FD) and phylogenetic diversity (PD) of plants possessing extrafloral nectaries (EFNs) in semi-arid Brazilian Caatinga. EFNs attract ants that protect plants against insect herbivore attack and are extremely prevalent in the Caatinga flora. EFN-bearing plants were censused along gradients of disturbance and rainfall in Catimbau National Park in north-eastern Brazil. We recorded a total of 2243 individuals belonging to 21 species. Taxonomic and functional composition varied along the rainfall gradient, but not along the disturbance gradient. There was a significant interaction between increasing disturbance and decreasing rainfall, with CAD leading to decreased TD, FD and PD in the most arid areas, and to increased TD, FD and PD in the wettest areas. We found a strong phylogenetic signal in the EFN traits we analysed, which explains the strong matching between patterns of FD and PD along the environmental gradients. The interactive effects of disturbance and rainfall revealed by our study indicate that the decreased rainfall forecast for Caatinga under climate change will increase the sensitivity of EFN-bearing plants to anthropogenic disturbance. This has important implications for the availability of a key food resource, which would likely have cascading effects on higher trophic levels.

Indigenous plants promote insect biodiversity in urban greenspaces.

The contribution of urban greenspaces to support biodiversity and provide benefits for people is increasingly recognized. However, ongoing management practices favor vegetation oversimplification, often limiting greenspaces to lawns and tree canopy rather than multi-layered vegetation that includes under- and midstorey, and the use of nonnative species. These practices hinder the potential of greenspaces to sustain indigenous biodiversity, particularly for taxa like insects that rely on plants for food and habitat. Yet, little is known about which plant species may maximize positive outcomes for taxonomically and functionally diverse insect communities in greenspaces. Additionally, while cities are expected to experience high rates of introductions, quantitative assessments of the relative occupancy of indigenous vs. introduced insect species in greenspace are rare, hindering understanding of how management may promote indigenous biodiversity while limiting the establishment of introduced insects. Using a hierarchically replicated study design across 15 public parks, we recorded occurrence data from 552 insect species on 133 plant species, differing in planting design element (lawn, midstorey, and tree canopy), midstorey growth form (forbs, lilioids, graminoids, and shrubs) and origin (nonnative, native, and indigenous), to assess (1) the relative contributions of indigenous and introduced insect species and (2) which plant species sustained the highest number of indigenous insects. We found that the insect community was overwhelmingly composed of indigenous rather than introduced species. Our findings further highlight the core role of multi-layered vegetation in sustaining high insect biodiversity in urban areas, with indigenous midstorey and canopy representing key elements to maintain rich and functionally diverse indigenous insect communities. Intriguingly, graminoids supported the highest indigenous insect richness across all studied growth forms by plant origin groups. Our work highlights the opportunity presented by indigenous understory and midstorey plants, particularly indigenous graminoids, in our study area to promote indigenous insect biodiversity in urban greenspaces. Our study provides a blueprint and stimulus for architects, engineers, developers, designers, and planners to incorporate into their practice plant species palettes that foster a larger presence of indigenous over regionally native or nonnative plant species, while incorporating a broader mixture of midstorey growth forms.

Responses of ant communities to disturbance: Five principles for understanding the disturbance dynamics of a globally dominant faunal group.

Ecological disturbance is fundamental to the dynamics of biological communities, yet a conceptual framework for understanding the responses of faunal communities to disturbance remains elusive. Here, I propose five principles for understanding the disturbance dynamics of ants-a globally dominant faunal group that is widely used as bioindicators in land management, which appear to have wide applicability to other taxa. These principles are as follows: (1) The most important effects of habitat disturbance on ants are typically indirect, through its effects on habitat structure, microclimate, resource availability and competitive interactions; (2) habitat openness is a key driver of variation in ant communities; (3) ant species responses to disturbance are to a large degree determined by their responses to habitat openness; (4) the same disturbance will have different effects on ants in different habitats, because of different impacts on habitat openness; and (5) ant community responses to the same disturbance will vary according to ant functional composition and biogeographical history in relation to habitat openness. I illustrate these principles using results primarily from studies of ant responses to fire, a dominant agent of disturbance globally, to provide a common disturbance currency for comparative analysis. I argue that many of the principles also apply to other faunal groups and so can be considered as general ecological "laws." As is the case for ants, many impacts of habitat disturbance on other faunal groups are fundamentally related to habitat openness, the effects of disturbance on it and the functional composition of species in relation to it.

Effects of increasing aridity and chronic anthropogenic disturbance on seed dispersal by ants in Brazilian Caatinga.

Anthropogenic disturbance and climate change are the main drivers of biodiversity loss and ecological services around the globe. There is concern that climate change will exacerbate the impacts of disturbance and thereby promote biotic homogenization, but its consequences for ecological services are unknown. We investigated the individual and interactive effects of increasing chronic anthropogenic disturbance (CAD) and aridity on seed dispersal services provided by ants in Caatinga vegetation of north-eastern Brazil. The study was conducted in Catimbau National Park, Pernambuco, Brazil. Within an area of 214 km2 , we established nineteen 50 × 20 m plots that encompassed gradients of both CAD and aridity. We offered diaspores of six plant species, three myrmecochorous diaspores and three fleshy fruits that are secondarily dispersed by ants. We then quantified the number of interactions, seed removal rate and dispersal distances, and noted the identities of interacting ant species. Finally, we used pitfall trap data to quantify the abundances of ant disperser species in each plot. Our results show that overall composition of ant disperser species varied along the gradients of CAD and aridity, but the composition of high-quality dispersers varied only with aridity. The total number of interactions, rates of removal and mean distance of removal all declined with increasing aridity, but they were not related to CAD. These same patterns were found when considering only high-quality disperser species, driven by the responses of the dominant disperser Dinoponera quadriceps. We found little evidence of interactive effects of CAD and aridity on seed dispersal services by ants. Our study indicates that CAD and aridity act independently on ant-mediated seed dispersal services in Caatinga, such that the impacts of anthropogenic disturbance are unlikely to change under the forecast climate of increased aridity. However, our findings highlight the vulnerability of seed dispersal services provided by ants in Caatinga under an increasingly arid climate due to low functional redundancy in high-quality disperser species. Given the large number of plant species dependent on ants for seed dispersal, this has important implications for future plant recruitment and, consequently, for the composition of Caatinga plant communities.

Different trophic groups of arboreal ants show differential responses to resource supplementation in a neotropical savanna.

Resource-ratio theory predicts that consumers should achieve optimal ratios of complementary nutrients. Accordingly, different trophic groups are expected to vary in their N-limitation depending on the extent to which they feed primarily on carbohydrate (CHO) or protein. Among arboreal ants, N-limitation ranges from high (for trophobiont tenders), intermediate (leaf foragers) and low (predators). We report results from a manipulative field experiment in a Brazilian savanna that tests the differential attractiveness of nitrogen and CHO to arboreal ants, as well as experimentally examines changes in broader ant foraging patterns in response to protein and CHO supplementation. Every tree within 32 20 × 20 m plots were supplemented with either protein, CHO; protein + CHO or a water control (n = 8 in each case) for a 7-day period in each of the wet and dry seasons. As predicted, different trophic groups responded differentially to supplementation treatment according to the extent of their N-limitation. The richness and abundance of the most N-limited group (trophobiont tenders) was highest at protein supplements, whereas less N-limited trophic groups showed highest species richness (leaf foragers) or abundance (predators) at CHO supplements. Protein supplementation markedly increased the general foraging abundance of trophobiont tenders, but decreased the abundance of leaf foragers. We attribute the latter to increased competition from behaviorally dominant trophobiont tenders. Our study provides experimental evidence that nutrient availability is a major factor influencing arboreal ant communities, both directly through the provision of different resources, and indirectly through increased competitive pressure.

Dominance-diversity relationships in ant communities differ with invasion.

The relationship between levels of dominance and species richness is highly contentious, especially in ant communities. The dominance-impoverishment rule states that high levels of dominance only occur in species-poor communities, but there appear to be many cases of high levels of dominance in highly diverse communities. The extent to which dominant species limit local richness through competitive exclusion remains unclear, but such exclusion appears more apparent for non-native rather than native dominant species. Here we perform the first global analysis of the relationship between behavioral dominance and species richness. We used data from 1,293 local assemblages of ground-dwelling ants distributed across five continents to document the generality of the dominance-impoverishment rule, and to identify the biotic and abiotic conditions under which it does and does not apply. We found that the behavioral dominance-diversity relationship varies greatly, and depends on whether dominant species are native or non-native, whether dominance is considered as occurrence or relative abundance, and on variation in mean annual temperature. There were declines in diversity with increasing dominance in invaded communities, but diversity increased with increasing dominance in native communities. These patterns occur along the global temperature gradient. However, positive and negative relationships are strongest in the hottest sites. We also found that climate regulates the degree of behavioral dominance, but differently from how it shapes species richness. Our findings imply that, despite strong competitive interactions among ants, competitive exclusion is not a major driver of local richness in native ant communities. Although the dominance-impoverishment rule applies to invaded communities, we propose an alternative dominance-diversification rule for native communities.

Habitat-contingent responses to disturbance: impacts of cattle grazing on ant communities vary with habitat complexity.

Predicting community responses to disturbance is a major challenge for both ecology and ecosystem management. A particularly challenging issue is that the same type and intensity of disturbance can have different impacts in different habitats. We investigate how habitat contingency influences ant community responses to disturbance in arid Australia, testing the hypothesis that disturbance has a greater impact in more complex habitats. We also assess the effectiveness of a highly simplified ant assessment protocol that considers larger species only. We sampled ants at 46 sites from two habitats (Chandler, low chenopod shrubland; and mulga, low woodland) with contrasting complexity, using distance from water as a surrogate for variation in grazing intensity. We assessed variation in habitat structural variables (basal area of perennial grass, and cover of herbs, litter, and bare ground) and ant communities in relation to habitat and distance from water, first using data from the entire ant community and then for larger ants (>4 mm body length) only. Site species richness was almost twice as high in mulga, the more structurally complex habitat, than in Chandler, and ant communities in mulga showed far more variation in relation to distance from water. Litter cover was the key environmental variable associated with the interaction between grazing and habitat: it increased with increasing distance from water in mulga and was virtually absent from Chandler. Analysis of only larger species revealed the same patterns of variation in ant abundance, species richness and composition in relation to habitat and grazing as shown by entire ant communities. Our findings support the hypothesis that disturbance impacts on faunal communities increase with increasing habitat complexity. An appreciation of such habitat contingency is important for a predictive understanding and therefore effective management of disturbances such as rangeland grazing. Our findings also show that simplified assessment can provide robust information on the responses of highly diverse ant communities to disturbance, which enhances their feasibility for use as bio-indicators in land management.

Is thermal limitation the primary driver of elevational distributions? Not for montane rainforest ants in the Australian Wet Tropics.

Terrestrial ectotherms are likely to be especially sensitive to rising temperatures over coming decades. Thermal limits are used to measure climatic tolerances that potentially affect ectotherm distribution. While there is a strong relationship between the critical thermal maximum (CTmax) of insects and their latitudinal ranges, the nature of this relationship across elevation is less clear. Here we investigated the combined relationships between CTmax, elevation and ant body mass, given that CTmax can also be influenced by body mass, in the World Heritage-listed rainforests of the Australian Wet Tropics. We measured the CTmax and body mass of 20 ant species across an elevational gradient from 350 to 1000 m a.s.l. Community CTmax did not vary systematically with increasing elevation and there was no correlation between elevation and elevational ranges of species. However, body mass significantly decreased at higher elevations. Despite the negative correlation between CTmax and body mass at the community level, there was no significant difference in CTmax of different-sized ants within a species. These findings are not consistent with either the climatic variability hypothesis, Rapoport's rule or Bergmann's rule. Models indicated that elevation and body mass had limited influences on CTmax. Our results suggest that the distribution of most montane ants in the region is not strongly driven by thermal limitation, and climate change will likely impact ant species differently. This is likely to occur primarily through changes in rainfall via its effects on vegetation structure and therefore thermal microhabitats, rather than through direct temperature changes.

A global database of ant species abundances.

What forces structure ecological assemblages? A key limitation to general insights about assemblage structure is the availability of data that are collected at a small spatial grain (local assemblages) and a large spatial extent (global coverage). Here, we present published and unpublished data from 51 ,388 ant abundance and occurrence records of more than 2,693 species and 7,953 morphospecies from local assemblages collected at 4,212 locations around the world. Ants were selected because they are diverse and abundant globally, comprise a large fraction of animal biomass in most terrestrial communities, and are key contributors to a range of ecosystem functions. Data were collected between 1949 and 2014, and include, for each geo-referenced sampling site, both the identity of the ants collected and details of sampling design, habitat type, and degree of disturbance. The aim of compiling this data set was to provide comprehensive species abundance data in order to test relationships between assemblage structure and environmental and biogeographic factors. Data were collected using a variety of standardized methods, such as pitfall and Winkler traps, and will be valuable for studies investigating large-scale forces structuring local assemblages. Understanding such relationships is particularly critical under current rates of global change. We encourage authors holding additional data on systematically collected ant assemblages, especially those in dry and cold, and remote areas, to contact us and contribute their data to this growing data set.

Fire in the Amazon: impact of experimental fuel addition on responses of ants and their interactions with myrmecochorous seeds.

The widespread clearing of tropical forests causes lower tree cover, drier microclimate, and higher and drier fuel loads of forest edges, increasing the risk of fire occurrence and its intensity. We used a manipulative field experiment to investigate the influence of fire and fuel loads on ant communities and their interactions with myrmecochorous seeds in the southern Amazon, a region currently undergoing extreme land-use intensification. Experimental fires and fuel addition were applied to 40 × 40-m plots in six replicated blocks, and ants were sampled between 15 and 30 days after fires in four strata: subterranean, litter, epigaeic, and arboreal. Fire had extensive negative effects on ant communities. Highly specialized cryptobiotic and predator species of the litter layer and epigaeic specialist predators were among the most sensitive, but we did not find evidence of overall biotic homogenization following fire. Fire reduced rates of location and transport of myrmecochorous seeds, and therefore the effectiveness of a key ecosystem service provided by ants, which we attribute to lower ant abundance and increased thermal stress. Experimental fuel addition had only minor effects on attributes of fire severity, and limited effects on ant responses to fire. Our findings indicate that enhanced fuel loads will not decrease ant diversity and ecosystem services through increased fire severity, at least in wetter years. However, higher fuel loads can still have a significant effect on ants from Amazonian rainforests because they increase the risk of fire occurrence, which has a detrimental impact on ant communities and a key ecosystem service they provide.

Climate mediates the effects of disturbance on ant assemblage structure.

Many studies have focused on the impacts of climate change on biological assemblages, yet little is known about how climate interacts with other major anthropogenic influences on biodiversity, such as habitat disturbance. Using a unique global database of 1128 local ant assemblages, we examined whether climate mediates the effects of habitat disturbance on assemblage structure at a global scale. Species richness and evenness were associated positively with temperature, and negatively with disturbance. However, the interaction among temperature, precipitation and disturbance shaped species richness and evenness. The effect was manifested through a failure of species richness to increase substantially with temperature in transformed habitats at low precipitation. At low precipitation levels, evenness increased with temperature in undisturbed sites, peaked at medium temperatures in disturbed sites and remained low in transformed sites. In warmer climates with lower rainfall, the effects of increasing disturbance on species richness and evenness were akin to decreases in temperature of up to 9°C. Anthropogenic disturbance and ongoing climate change may interact in complicated ways to shape the structure of assemblages, with hot, arid environments likely to be at greatest risk.

Molecular phylogeny of Indo-Pacific carpenter ants (Hymenoptera: Formicidae, Camponotus) reveals waves of dispersal and colonization from diverse source areas.

Ants that resemble Camponotus maculatus (Fabricius, 1782) present an opportunity to test the hypothesis that the origin of the Pacific island fauna was primarily New Guinea, the Philippines, and the Indo-Malay archipelago (collectively known as Malesia). We sequenced two mitochondrial and four nuclear markers from 146 specimens from Pacific islands, Australia, and Malesia. We also added 211 specimens representing a larger worldwide sample and performed a series of phylogenetic analyses and ancestral area reconstructions. Results indicate that the Pacific members of this group comprise several robust clades that have distinctly different biogeographical histories, and they suggest an important role for Australia as a source of Pacific colonizations. Malesian areas were recovered mostly in derived positions, and one lineage appears to be Neotropical. Phylogenetic hypotheses indicate that the orange, pan-Pacific form commonly identified as C. chloroticus Emery 1897 actually consists of two distantly related lineages. Also, the lineage on Hawai'i, which has been called C. variegatus (Smith, 1858), appears to be closely related to C. tortuganus Emery, 1895 in Florida and other lineages in the New World. In Micronesia and Polynesia the C. chloroticus-like species support predictions of the taxon-cycle hypothesis and could be candidates for human-mediated dispersal.

Anthropogenic disturbance reduces seed-dispersal services for myrmecochorous plants in the Brazilian Caatinga.

Anthropogenic disturbance can have important indirect effects on ecosystems by disrupting species interactions. Here we examine the effects of anthropogenic disturbance on distance dispersal by ants for the diaspores of myrmecochorous Euphorbiaceae in Brazilian Caatinga. Rates of diaspore removal and distances removed of Croton sonderianus and Jatropha mollissima were observed at 24 sites ranging from low to very high disturbance (primarily grazing by livestock, hunting and firewood collection). Despite a large number of seed-disperser ant species, there were only two species providing high-quality distance-dispersal services, Dinoponera quadriceps (40% of all observed seed removals) and Ectatomma muticum (33%). D. quadriceps was responsible for 97% of all removals >2 m, and 100% of all removals >5 m. Removal rates did not vary with disturbance for C. sonderianus (small elaiosome), but declined with increasing disturbance for J. mollissima (large elaiosome). The number of removals by Ectatomma was highest at intermediate levels of disturbance, whereas those by Dinoponera decreased systematically with increasing levels of disturbance. Mean dispersal distance was four times higher at sites experiencing low disturbance, where removals >5 m represented a third of all removal events, compared with very highly disturbed sites, where no removals >5 m were observed. Despite high overall diversity there is very limited functional redundancy in disperser ant species, resulting in low disperser resilience in relation to disturbance. This is likely to have important implications for recruitment by myrmecochorous plants, and therefore on vegetation composition and structure, at sites subject to high anthropogenic disturbance.

Myrmecochores can target high-quality disperser ants: variation in elaiosome traits and ant preferences for myrmecochorous Euphorbiaceae in Brazilian Caatinga.

Recent evidence suggests that the traditional view of myrmecochory as a highly diffuse interaction between diaspores and a wide range of ant species attracted to their elaiosomes may not be correct. The effectiveness of dispersal varies markedly among ant species, and combined with differential attractiveness of diaspores due to elaiosome size and composition, this raises the potential for myrmecochorous plants to target ant species that offer the highest quality dispersal services. We ask the question: Do particular physical and chemical traits of elaiosomes result in disproportionate removal of Euphorbiaceae diaspores by high-quality disperser ants in Caatinga vegetation of north-eastern Brazil? We offered seeds of five euphorb species that varied in morphological and chemical traits of elaiosomes to seed-dispersing ants. High-quality seed-disperser ants (species of Dinoponera, Ectatomma and Camponotus) were identified as those that rapidly collected and transported diaspores to their nests, often over substantial distances, whereas low-quality disperser ants (primarily species of Pheidole and Solenopsis) typically fed on elaiosomes in situ, and only ever transported diaspores very short distances. Low-quality disperser ants were equally attracted to the elaiosomes of all study species. However, high-quality dispersers showed a strong preference for diaspores with the highest elaiosome mass (and especially proportional mass). As far as we are aware, this is the first study to identify a mechanism of diaspore selection by high-quality ant dispersers based on elaiosome traits under field conditions. Our findings suggest that myrmecochorous plants can preferentially target high-quality seed-disperser ants through the evolution of particular elaiosome traits.

Fire influences ant diversity by modifying vegetation structure in an Australian tropical savanna.

Fire is a dominant ecological force shaping many faunal communities globally. Fire affects fauna either directly, such as by killing individuals, or indirectly, such as by modifying vegetation structure. Vegetation structure itself also modulates fire frequency and intensity. As such, faunal responses to fire need to be seen through the lens of variable fire activity and vegetation structure. Here, we incorporate information on fire activity and vegetation structure to enhance an understanding of the response of ants to long-term (17-year) experimental fire treatments in an extremely fire-prone tropical savanna in northern Australia. Previous analysis revealed limited divergence in ant communities after 5 years of experimental fire treatment. Hence, we first investigated the extent to which ant communities diverged over a subsequent 12 years of treatment. We then assessed the relative contribution of fire treatment, cumulative fire intensity (fire activity), and woody cover to responses of ant species frequency of occurrence, richness, and composition. We found that, even after 17 years, fire treatments explained little variation in any ant response variable. In contrast, woody cover was a strong predictor for all of them, while fire activity was a moderate predictor for abundance and richness. Ant species occurrence and richness increased in open habitats receiving higher levels of fire activity, compared with plots with higher vegetation cover experiencing low (or no) fire activity. Moreover, species composition differed between plots with high and low vegetation cover. Our findings provide experimental support to the principle that the effects of fire on fauna are primarily indirect, via its effect on vegetation structure. Furthermore, our results show that a "uniform" fire regime does not have uniform impacts on the ant fauna, because of variability imposed by interactions between vegetation structure and fire activity. This helps to explain why there is often a weak relationship between pyrodiversity and biodiversity, and it lessens the need for active management of pyrodiversity to maintain biodiversity.

Arboreality drives heat tolerance while elevation drives cold tolerance in tropical rainforest ants.

Determining how species thermal limits correlate with climate is important for understanding biogeographic patterns and assessing vulnerability to climate change. Such analyses need to consider thermal gradients at multiple spatial scales. Here we relate thermal traits of rainforest ants to microclimate conditions from ground to canopy (microgeographic scale) along an elevation gradient (mesogeographic scale) and calculate warming tolerance to assess climate change vulnerability in the Australian Wet Tropics Bioregion. We test the thermal adaptation and thermal niche asymmetry hypotheses to explain interspecific patterns of thermal tolerance at these two spatial scales. We tested cold tolerance (CTmin ), heat tolerance (CTmax ), and calculated thermal tolerance range (CTrange ), using ramping assays for 74 colonies of 40 ant species collected from terrestrial and arboreal habitats at lowland and upland elevation sites and recorded microclimatic conditions for one year. Within sites, arboreal ants were exposed to hotter microclimates and on average had a 4.2°C (95% CI: 2.7-5.6°C) higher CTmax and 5.3°C (95% CI: 3.5-7°C) broader CTrange than ground-dwelling ants. This pattern was consistent across the elevation gradient, whether it be the hotter lowlands or the cooler uplands. Across elevation, upland ants could tolerate significantly colder temperatures than lowland ants, whereas the change in CTmax was less pronounced, and CTrange did not change over elevation. Differential exposure to microclimates, due to localized niche preferences, drives divergence in CTmax , while environmental temperatures along the elevation gradient drive divergence in CTmin . Our results suggest that both processes of thermal adaptation and thermal niche asymmetry are at play, depending on the spatial scale of observation, and we discuss potential mechanisms underlying these patterns. Despite the broad thermal tolerance range of arboreal rainforest ants, lowland arboreal ants had the lowest warming tolerance and may be most vulnerable to climate change.

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.

A new species of Epopostruma (Hymenoptera: Formicidae) from monsoonal Australia.

Epopostruma is an uncommon genus of myrmicine ants endemic to relatively mesic regions of southern and eastern Australia. Here we describe a new species recently recorded from the 'Top End of Australias Northern Territory, E. topendi sp. n. from Melville Island and Nitmiluk National Park. The new species is known from one specimen from each of two sites in the Australian monsoonal tropics, the only records of the genus from this region. This addition brings the number of described species of Epopostruma to twenty.

Canopy and litter ant assemblages share similar climate-species density relationships.

Tropical forest canopies house most of the globe's diversity, yet little is known about global patterns and drivers of canopy diversity. Here, we present models of ant species density, using climate, abundance and habitat (i.e. canopy versus litter) as predictors. Ant species density is positively associated with temperature and precipitation, and negatively (or non-significantly) associated with two metrics of seasonality, precipitation seasonality and temperature range. Ant species density was significantly higher in canopy samples, but this difference disappeared once abundance was considered. Thus, apparent differences in species density between canopy and litter samples are probably owing to differences in abundance-diversity relationships, and not differences in climate-diversity relationships. Thus, it appears that canopy and litter ant assemblages share a common abundance-diversity relationship influenced by similar but not identical climatic drivers.

Climatic drivers of hemispheric asymmetry in global patterns of ant species richness.

Although many taxa show a latitudinal gradient in richness, the relationship between latitude and species richness is often asymmetrical between the northern and southern hemispheres. Here we examine the latitudinal pattern of species richness across 1003 local ant assemblages. We find latitudinal asymmetry, with southern hemisphere sites being more diverse than northern hemisphere sites. Most of this asymmetry could be explained statistically by differences in contemporary climate. Local ant species richness was positively associated with temperature, but negatively (although weakly) associated with temperature range and precipitation. After contemporary climate was accounted for, a modest difference in diversity between hemispheres persisted, suggesting that factors other than contemporary climate contributed to the hemispherical asymmetry. The most parsimonious explanation for this remaining asymmetry is that greater climate change since the Eocene in the northern than in the southern hemisphere has led to more extinctions in the northern hemisphere with consequent effects on local ant species richness.