Similar seed preferences explain trophic ecology of functionally distinct, but co-occurring and closely related harvester ants.

To understand how food resource use and partitioning by closely related species allows local coexistence, it is key to determine whether a species' diet reflects food availability or food preferences. Here, we analysed the diets, seed selection, and seed preferences of three closely related harvester ants: Messor barbarus, M. bouvieri, and M. capitatus. Sympatric within a Mediterranean shrubland, these species differ in foraging behaviour and worker polymorphism. For 2 years, we studied the ants' diets and seed selection patterns as well as the local availability of seeds. Additionally, we performed a seed-choice experiment using a paired comparison design, offering the ants seeds from eight native plant species. The three ant species had the same general diet, which was primarily granivorous. Although they all consumed a wide variety of seeds, they mostly selected seeds from a small subset of plant species. Despite their morphological and behavioural differences, the ants displayed similar seed preferences that were highly consistent with their diets and seed selection patterns. Our results support the idea that the trophic ecology of these three harvester ants is driven by similar seed preferences rather than by their morphological and behavioural differences. Seed diversity and abundance were high near the ants' nests, suggesting that seed availability is not limiting and could in fact favour local species coexistence.

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.

Evidence for locally adaptive metabolic rates among ant populations along an elevational gradient.

As global temperatures rise, the mechanistic links between temperature, physiology and behaviour will increasingly define predictions of ecological change. However, for many taxa, we currently lack consensus about how thermal performance traits vary within and across populations, and whether and how locally adaptive trait plasticity can buffer warming effects. The metabolic cold adaptation hypothesis posits that cold environments (e.g. high elevations and latitudes) select for high metabolic rates (MR), even after controlling for body size differences, and that this enables high activity levels when an organism is near its cold lower thermal limits. Steep MR reaction norms are further predicted at cold temperatures to enable rapid behavioural activation with rising temperatures needed to exploit brief thermal windows suitable for performing eco-evolutionary tasks. We tested these predictions by performing common garden experiments comparing thermal reaction norms of MR (from 15 to 32°C) and behaviour (from 10 to 40°C) across populations of the ant Aphaenogaster iberica sampled from a 2 km elevation gradient in the Sierra Nevada Mountains of southern Spain. As predicted, high-elevation ants had higher MR and steeper MR-temperature reaction norms. However, higher rates of energy use did not yield the predicted benefits of steeper activity-level reaction norms. The evidence for locally adaptive metabolic physiology only became apparent at intermediate temperatures, highlighting the importance of testing thermal performance hypotheses across thermal gradients, rather than focusing only on performance at thermal limits (i.e. critical thermal values). The partial support for the metabolic cold adaptation hypothesis highlights that while organisms likely show a wealth of unexplored metabolic temperature plasticity, the physiological mechanisms and eco-evolutionary trade-offs underlying such local adaptation remain obscure.

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.

Combined exposure to sublethal concentrations of an insecticide and a fungicide affect feeding, ovary development and longevity in a solitary bee.

Pollinators in agroecosystems are often exposed to pesticide mixtures. Even at low concentrations, the effects of these mixtures on bee populations are difficult to predict due to potential synergistic interactions. In this paper, we orally exposed newly emerged females of the solitary bee Osmia bicornis to environmentally realistic levels of clothianidin (neonicotinoid insecticide) and propiconazole (fungicide), singly and in combination. The amount of feeding solution consumed was highest in bees exposed to the neonicotinoid, and lowest in bees exposed to the pesticide mixture. Ovary maturation and longevity of bees of the neonicotinoid and the fungicide treatments did not differ from those of control bees. By contrast, bees exposed to the pesticide mixture showed slow ovary maturation and decreased longevity. We found a synergistic interaction between the neonicotinoid and the fungicide on survival probability. We also found an interaction between treatment and emergence time (an indicator of physiological condition) on longevity. Longevity was negatively correlated to physiological condition only in the fungicide and the mixture treatments. Delayed ovary maturation and premature death imply a shortened nesting period (highly correlated to fecundity in Osmia). Our findings provide a mechanism to explain the observed dynamics of solitary bee populations exposed to multiple chemical residues in agricultural environments.

Disentangling nutritional pathways linking leafcutter ants and their co-evolved fungal symbionts using stable isotopes.

Leafcutter ants are the ultimate insect superorganisms, with up to millions of physiologically specialized workers cooperating to cut and transport vegetation and then convert it into compost used to cultivate co-evolved fungi, domesticated over millions of years. We tested hypotheses about the nutrient-processing dynamics governing this functional integration, tracing 15 N- and 13 C-enriched substrates through colonies of the leafcutter ant Atta colombica. Our results highlight striking performance efficiencies, including rapid conversion (within 2 d) of harvested nutrients into edible fungal tissue (swollen hyphal tips called gongylidia) in the center of fungus gardens, while also highlighting that much of each colony's foraging effort resulted in substrate placed directly in the trash. We also find nutrient-specific processing dynamics both within and across layers of the fungus garden, and in ant consumers. Larvae exhibited higher overall levels of 15 N and 13 C enrichment than adult workers, supporting that the majority of fungal productivity is allocated to colony growth. Foragers assimilated 13 C-labeled glucose during its ingestion, but required several days to metabolically process ingested 15 N-labeled ammonium nitrate. This processing timeline helps resolve a 40-yr old hypothesis, that foragers (but apparently not gardeners or larvae) bypass their fungal crops to directly assimilate some of the nutrients they ingest outside the nest. Tracing these nutritional pathways with stable isotopes helps visualize how physiological integration within symbiotic networks gives rise to the ecologically dominant herbivory of leafcutter ants in habitats ranging from Argentina to the southern United States.

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.

Effects of chronic anthropogenic disturbance and rainfall on the specialization of ant-plant mutualistic networks in the Caatinga, a Brazilian dry forest.

Anthropogenic disturbance and climate change might negatively affect the ecosystem services provided by mutualistic networks. However, the effects of such forces remain poorly characterized. They may be especially important in dry forests, which (1) experience chronic anthropogenic disturbances (CADs) as human populations exploit forest resources, and (2) are predicted to face a 22% decline in rainfall under climate change. In this study, we investigated the separate and combined effects of CADs and rainfall levels on the specialization of mutualistic networks in the Caatinga, a seasonally dry tropical forest typical of north-eastern Brazil. More specifically, we examined interactions between plants bearing extrafloral nectaries (EFNs) and ants. We analysed whether differences in network specialization could arise from environmentally mediated variation in the species composition, namely via the replacement of specialist by generalist species. We characterized these ant-plant networks in 15 plots (20 × 20 m) that varied in CAD intensity and mean annual rainfall. We quantified CAD intensity by calculating three indices related to the main sources of disturbance in the Caatinga: livestock grazing (LG), wood extraction (WE) and miscellaneous resource use (MU). We determined the degree of ant-plant network specialization using four metrics: generality, vulnerability, interaction evenness and H2 '. Our results indicate that CADs differentially influenced network specialization: we observed positive, negative, and neutral responses along LG, MU and WE gradients, respectively. The pattern was most pronounced with LG. Rainfall also shaped network specialization, markedly increasing it. While LG and rainfall were associated with changes in network species composition, this trend was not related to the degree of species specialization. This result suggests that shifts in network specialization might be related to changes in species behaviour, not species composition. Our study highlights the vulnerability of such dry forest ant-plant networks to climate change. Moreover, dry forests experience highly heterogeneous anthropogenic disturbances, creating a geographic mosaic of selective forces that may shape the co-evolution of interactions between ants and EFN-bearing plants.

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.

Body size phenology in a regional bee fauna: a temporal extension of Bergmann's rule.

Bergmann's rule originally described a positive relationship between body size and latitude in warm-blooded animals. Larger animals, with a smaller surface/volume ratio, are better enabled to conserve heat in cooler climates (thermoregulatory hypothesis). Studies on endothermic vertebrates have provided support for Bergmann's rule, whereas studies on ectotherms have yielded conflicting results. If the thermoregulatory hypothesis is correct, negative relationships between body size and temperature should occur in temporal in addition to geographical gradients. To explore this possibility, we analysed seasonal activity patterns in a bee fauna comprising 245 species. In agreement with our hypothesis of a different relationship for large (endothermic) and small (ectothermic) species, we found that species larger than 27.81 mg (dry weight) followed Bergmann's rule, whereas species below this threshold did not. Our results represent a temporal extension of Bergmann's rule and indicate that body size and thermal physiology play an important role in structuring community phenology.

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.

A multidimensional functional trait analysis of resource exploitation in European ants.

The major factors explaining ecological variation in plants have been widely discussed over the last decade thanks to numerous studies that have examined the covariation that exists between pairs of traits. However, multivariate relationships among traits remain poorly characterized in animals. In this study, we aimed to identify the main multivariate trait dimensions that explain variance in important functional traits related to resource exploitation in ants. To this end, we created a large ant trait database. This database includes information on 11 traits that are important in ant resource exploitation; data were obtained for 150 European species found in different biomes. First, we examined the pairwise correlations between the traits included in the database. Second, we used multivariate analyses to identify potential trait dimensions. Our study shows that, to a great extent, resource exploitation strategies align along two main trait dimensions. The first dimension emerged in both the overall and group-specific analyses, where it accounted for the same pairwise trait correlations. The second dimension was more variable, as species were grouped by levels of taxonomy, habitat, and climate. These two dimensions included most of the significant pairwise trait correlations, thus highlighting that complementarity, but also redundancy, exists among different pairs of traits. The first dimension was associated with behavioral dominance: dominance was associated with large colony size, presence of multiple nests per colony, worker polymorphism, and a collective foraging 'strategy. The second dimension was associated with resource partitioning along dietary and microhabitat lines: it ranged from species that consume liquid foods, engage in group foraging, and mainly nest in the vegetation to species that consume insects and seeds, engage in individual foraging, and demonstrate strictly diurnal activity. Our findings establish a proficient ecological trait-based animal research that minimizes the number of traits to be measured while maximizing the number of relevant trait dimensions. Overall, resource exploitation in animals might be framed by behavioral dominance, foraging strategy, diet, and nesting habitat; the position of animal species within this trait space could provide relevant information about their distribution and abundance, for today as well as under future global change scenarios.

Local and landscape effects in a host-parasitoid interaction network along a forest-cropland gradient.

Land-use driven habitat modification is a major driver of biodiversity loss and impoverishment of interaction diversity. This may affect ecosystem services such as pollination and biological control. Our objective is to analyze the effects of local (nesting environment: farms vs. tree stands) and landscape (forest-cropland gradient) factors on the structure and composition of a cavity-nesting bee-wasp (CNBW) community, their nests associates (henceforth parasitoids), and their interactions. We set up 24 nest-trapping stations in a fragmented, extensively farmed area of ~100 km². We obtained 2035 nests containing 7572 brood cells representing 17 bee and 18 wasp species, attacked by 20 parasitoid species. Community structure and composition, as well as network structure, were much more dependent on local than on landscape factors. Host abundance and richness were higher in farms. In addition, host abundance was positively correlated to cropland cover. We also found highly significant differences between nesting environments in host community composition. Structure and composition of the parasitoid community were conditioned by the structure and composition of the host community. Network structure was affected by nesting environment but not by landscape factors. Interactions tended to be more diverse in farms. This result was mostly explained by differences in network size (greater in farms). However, generality was significantly higher in farms even after controlling for network size, indicating that differences in species' interaction patterns associated to differences in community composition between the two nesting environments are also affecting network structure. In conclusion, open habitats associated with extensively farmed exploitations favor local CNBW diversity (especially bees) and result in more complex host-parasitoid interaction networks in comparison to forested areas. The conservation value of this kind of open habitat is important in view of the progressive abandonment of extensively cultivated farmland taking place in Europe at the expense of agricultural intensification and reforestation.

Ant functional responses along environmental gradients.

Understanding species distributions and diversity gradients is a central challenge in ecology and requires prior knowledge of the functional traits mediating species' survival under particular environmental conditions. While the functional ecology of plants has been reasonably well explored, much less is known about that of animals. Ants are among the most diverse, abundant and ecologically significant organisms on earth, and they perform a great variety of ecological functions. In this study, we analyse how the functional species traits present in ant communities vary along broad gradients in climate, productivity and vegetation type in the south-western Mediterranean. To this end, we compiled one of the largest animal databases to date: it contains information on 211 local ant communities (including eight climate variables, productivity, and vegetation type) and 124 ant species, for which 10 functional traits are described. We used traits that characterize different dimensions of the ant functional niche with respect to morphology, life history and behaviour at both individual and colony level. We calculated two complementary functional trait community indices ('trait average' and 'trait dissimilarity') for each trait, and we analysed how they varied along the three different gradients using generalized least squares models that accounted for spatial autocorrelation. Our results show that productivity, vegetation type and, to a lesser extent, each climate variable per se might play an important role in shaping the occurrence of functional species traits in ant communities. Among the climate variables, temperature and precipitation seasonality had a much higher influence on functional responses than their mean values, whose effects were almost lacking. Our results suggest that strong relationships might exist between the abiotic environment and the distribution of functional traits among south-western Mediterranean ant communities. This finding indicates that functional traits may modulate the responses of ant species to the environment. Since these traits act as the link between species distributions and the environment, they could potentially be used to predict community changes under future global change scenarios.

Distinctive life traits and distribution along environmental gradients of dominant and subordinate Mediterranean ant species.

For most animal and plant species, life traits strongly affect their species-specific role, function or position within ecological communities. Previous studies on ant communities have mostly focused on the role of dominant species and the outcome of interspecific interactions. However, life traits of ant species have seldom been considered within a community framework. This study (1) analyses life traits related to ecological and behavioural characteristics of dominant and subordinate ant species from 13 sites distributed throughout the Iberian Peninsula, (2) determines how similar the ant species are within each of the two levels of the dominance hierarchy, and (3) establishes the distribution patterns of these different groups of species along environmental gradients. Our results showed that the differences between dominants and subordinates fall into two main categories: resource exploitation and thermal tolerance. Dominant species have more populated colonies and defend food resources more fiercely than subordinates, but they display low tolerance to high temperatures. We have identified different assemblages of species included within each of these two levels in the dominance hierarchy. The distribution of these assemblages varied along the environmental gradient, shifting from dominant Dolichoderinae and cryptic species in moist areas, to dominant Myrmicinae and hot climate specialists mainly in open and hot sites. We have been able to identify a set of life traits of the most common Iberian ant species that has enabled us to characterise groups of dominant and subordinate species. Although certain common features within the groups of both dominants and subordinates always emerge, other different features allow for differentiating subgroups within each of these groups. These different traits allow the different subgroups coping with particular conditions across environmental gradients.

Critical thermal limits in ants and their implications under climate change.

Critical thermal limits (CTLs) constrain the performance of organisms, shaping their abundance, current distributions, and future distributions. Consequently, CTLs may also determine the quality of ecosystem services as well as organismal and ecosystem vulnerability to climate change. As some of the most ubiquitous animals in terrestrial ecosystems, ants are important members of ecological communities. In recent years, an increasing body of research has explored ant physiological thermal limits. However, these CTL data tend to centre on a few species and biogeographical regions. To encourage an expansion of perspectives, we herein review the factors that determine ant CTLs and examine their effects on present and future species distributions and ecosystem processes. Special emphasis is placed on the implications of CTLs for safeguarding ant diversity and ant-mediated ecosystem services in the future. First, we compile, quantify, and categorise studies on ant CTLs based on study taxon, biogeographical region, methodology, and study question. Second, we use this comprehensive database to analyse the abiotic and biotic factors shaping ant CTLs. Our results highlight how CTLs may affect future distribution patterns and ecological performance in ants. Additionally, we identify the greatest remaining gaps in knowledge and create a research roadmap to promote rapid advances in this field of study.

What are the consequences of ant-seed interactions on the abundance of two dry-fruited shrubs in a Mediterranean scrub?

Strong interactions between dry-fruited shrubs and seed-harvesting ants are expected in early successional scrubs, where both groups have a major presence. We have analysed the implications of the seed characteristics of two dry-fruited shrub species (Coronilla minima and Dorycnium pentaphyllum) on seed predation and dispersal mediated by harvester ants and the consequences of these processes on spatio-temporal patterns of plant abundance in a heterogeneous environment. We found that large C. minima seeds were collected much more (39%) than small D. pentaphyllum seeds (2%). However, not all of the removed seeds of these plant species were consumed, and 12.8% of the seeds were lost along the trails, which increased dispersal distances compared with abiotic dispersal alone. Seed dropping occurred among all microhabitats of the two plant species, but especially in open microhabitats, which are the most suitable ones for plant establishment. The two plant species increased their presence in the study area during the study period: C. minima in open microhabitats and D. pentaphyllum in high vegetation. The large size of C. minima seeds probably limited the primary seed dispersal of this species, but may have allowed strong interaction with ants. Thus, seed dispersal by ants resulted in C. minima seeds reaching more suitable microhabitats by means of increasing dispersal distance and redistribution among microhabitats. In contrast, the smaller size of D. pentaphyllum seeds arguably allows abiotic seed dispersal over longer distances and colonization of all types of microhabitats, although it probably also limits their interaction with ants and, consequently, their redistribution in suitable microhabitats. We suggest that dyszoochory could contribute to the success of plant species with different seed characteristics in scrub habitats where seeds are abundantly collected by seed-harvesting ants.

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.

Introduced ant species occupy empty climatic niches in Europe.

Exploring shifts in the climatic niches of introduced species can provide significant insight into the mechanisms underlying the invasion process and the associated impacts on biodiversity. We aim to test the phylogenetic signal hypothesis in native and introduced species in Europe by examining climatic niche similarity. We examined data from 134 ant species commonly found in western Europe; 130 were native species, and 4 were introduced species. We characterized their distribution patterns using species records from different databases, determined their phylogenetic relatedness, and tested for a phylogenetic signal in their optimal climatic niches. We then compared the introduced species' climatic niches in Europe with their climatic niches in their native ranges and with the climatic niches of their closest relative species in Europe. We found a strong phylogenetic signal in the optimal climatic niches of the most common ant species in Europe; however, this signal was weak for the main climatic variables that affect the distributions of introduced versus native species. Also, introduced species occupied different climatic niches in Europe than in their native ranges; furthermore, their European climatic niches did not resemble those of their closest relative species in Europe. We further discovered that there was not much concordance between the climatic niches of introduced species in their native ranges and climatic conditions in Europe. Our findings suggest that phylogenetics do indeed constrain shifts in the climatic niches of native European ant species. However, introduced species would not face such constraints and seemed to occupy relatively empty climatic niches.

Nutritional niches reveal fundamental domestication trade-offs in fungus-farming ants.

During crop domestication, human farmers traded greater productivity for higher crop vulnerability outside specialized cultivation conditions. We found a similar domestication trade-off across the major co-evolutionary transitions in the farming systems of attine ants. First, the fundamental nutritional niches of cultivars narrowed over ~60 million years of naturally selected domestication, and laboratory experiments showed that ant farmers representing subsequent domestication stages strictly regulate protein harvest relative to cultivar fundamental nutritional niches. Second, ants with different farming systems differed in their abilities to harvest the resources that best matched the nutritional needs of their fungal cultivars. This was assessed by quantifying realized nutritional niches from analyses of items collected from the mandibles of laden ant foragers in the field. Third, extensive field collections suggest that among-colony genetic diversity of cultivars in small-scale farms may offer population-wide resilience benefits that species with large-scale farming colonies achieve by more elaborate and demanding practices to cultivate less diverse crops. Our results underscore that naturally selected farming systems have the potential to shed light on nutritional trade-offs that shaped the course of culturally evolved human farming.