The Ground-Dwelling Ant Fauna from a Cerrado Reserve in Southeastern Brazil: Vegetation Heterogeneity as a Promoter of Ant Diversity.

Ants represent one of the most diverse and ecologically important group of insects in tropical ecosystems, including in highly threatened ones such as the Brazilian Cerrado. Yet, a detailed understanding of the species diversity and composition of local Cerrado ant assemblages is lacking in many cases. Here we present the results of a comprehensive ant inventory performed within a region of the Cerrado (in São Paulo state) where most of the original vegetation has already been lost and where few conservation units exist. We performed consecutive surveys of the ant fauna that forage on the ground in replicated plots established in open savanna (campo sujo), dense savanna (cerrado sensu stricto), and forest (cerradão). Our surveys, with an estimated sample coverage of 99.4%, revealed a total of 219 species of ants from 60 genera, of which 36.1% were found in all the three vegetation types and 29.7% in just one. Rarefied species richness did not differ between vegetation types, but species composition differed markedly, especially between the two savannas in one hand and the forest in the other. Several species (60.1% of the 128 species analyzed) were significant "indicator" species due to their strong association with a given vegetation type. Overall, our findings reinforce the idea that habitat heterogeneity enhances ant diversity and that the mosaic of vegetation types that characterizes the Cerrado biome is one of the main factors explaining the elevated number of species that can be found at relatively small scales.

Revisiting florivory: an integrative review and global patterns of a neglected interaction.

Florivory is an ancient interaction which has rarely been quantified due to a lack of standardized protocols, thus impairing biogeographical and phylogenetic comparisons. We created a global, continuously updated, open-access database comprising 180 species and 64 families to compare floral damage between tropical and temperate plants, to examine the effects of plant traits on floral damage, and to explore the eco-evolutionary dynamics of flower-florivore interactions. Flower damage is widespread across angiosperms, but was two-fold higher in tropical vs temperate species, suggesting stronger fitness impacts in the tropics. Flowers were mostly damaged by chewers, but neither flower color nor symmetry explained differences in florivory. Herbivory and florivory levels were positively correlated within species, even though the richness of the florivore community does not affect florivory levels. We show that florivory impacts plant fitness via multiple pathways and that ignoring this interaction makes it more difficult to obtain a broad understanding of the ecology and evolution of angiosperms. Finally, we propose a standardized protocol for florivory measurements, and identify key research avenues that will help fill persistent knowledge gaps. Florivory is expected to be a central research topic in an epoch characterized by widespread decreases in insect populations that comprise both pollinators and florivores.

Patterns of Ant Diversity in the Natural Grasslands of Southern Brazil.

In the south of Brazil, grasslands are naturally widespread over two different biomes, the Pampa in the southernmost region and within the Atlantic Forest in the northern portions. The natural grasslands of the state of Paraná comprise a very particular physiognomy composed of two distinct formations: the Campos Gerais and the grasslands of the southwest. The first is located in the edge of the second plateau of Paraná state, comprising a great diversity of environments. The grasslands of the southwest are more homogeneous, with a continuous herbaceous stratum dominating the landscape. In this context, the aim of this study was to evaluate the patterns of species richness and composition of ants, an ecologically prominent group, along the natural grasslands of Paraná. We also intended to compare the faunal similarity between the two different grassland formations. For that, four different Conservation Unities were sampled along a latitudinal gradient. A remarkable total of 245 ant species was recorded, and the results indicate that species richness decreases as latitude increases along the grasslands of Paraná. There were clear differences in species composition between these two grasslands formations, given the significative number of endemic species in each of these two grassland formations. Ten species were recorded for the first time in the state of Paraná, of which three also for the first time in the Southern Region of Brazil. Overall, our study contributes to a better understanding about the diversity and composition of ant communities in subtropical grasslands.

Ecosystem engineering in the arboreal realm: heterogeneity of wood-boring beetle cavities and their use by cavity-nesting ants.

Wood-boring beetle larvae act as ecosystem engineers by creating stem cavities that are used secondarily as nests by many arboreal ant species. Understanding the heterogeneity and distribution of available cavities and their use by ants is therefore key to understanding arboreal ant community assembly and diversity. Our goals were to quantify the abundance and diversity of beetle-produced cavity resources in a tropical canopy, reveal how ants use these resources, and determine which characteristics of the cavity resource contribute to ant use. We dissected branches from six common tree species in the Brazilian Cerrado savanna, measuring cavity characteristics and identifying the occupants. We sampled 2310 individual cavities, 576 of which were used as nests by 25 arboreal ant species. We found significant differences among tree species in the proportion of stem length bored by beetles, the number of cavities per stem length, average entrance-hole size, and the distribution of cavity volumes. The likelihood that a cavity was occupied was greater for cavities with larger entrance-hole sizes and larger volumes. In particular, there was a strong positive correlation between mean head diameters of ant species and the mean entrance-hole diameter of the cavities occupied by those ant species. Wood-boring beetles contribute to the structuring of the Cerrado ant community by differentially attacking the available tree species. In so doing, the beetles provide a wide range of entrance-hole sizes which ant species partition based on their body size, and large volume cavities that ants appear to prefer.

The effects of high-severity fires on the arboreal ant community of a Neotropical savanna.

Fire-suppression is of concern in fire-prone ecosystems because it can result in the loss of endemic species. Suppressing fires also causes a build-up of flammable biomass, increasing the risk of severe fires. Using a Before-After, Control-Impacted design, we assessed the consequences of high-severity fires on Neotropical savanna arboreal ant communities. Over a 9-year period, we sampled the ant fauna of the same trees before and after two severe fires that hit a savanna reserve in Brazil and the trees from an unburned savanna site that served as a temporal control. The ant community associated with the unburned trees was relatively stable, with no significant temporal variation in species richness and only a few species changing in abundance over time. In contrast, we found a strong decline in species richness and marked changes in species composition in the burned trees, with some species becoming more prevalent and many becoming rare or locally extinct. The dissimilarity in species richness and composition was significantly smaller between the two pre-fire surveys than between the pre- and post-fire surveys. Fire-induced changes were much more marked among species with strictly arboreal nesting habits, and therefore more susceptible to the direct effects of fire. The decline of some of the ecologically dominant arboreal ant species may be particularly important, as it opens substantial ecological space for cascading community-wide changes. In particular, severe fires appear to disrupt the typical vertical stratification between the arboreal and ground-dwelling faunas, which might lead to homogenization of the overall ant community.

How much leaf area do insects eat? A data set of insect herbivory sampled globally with a standardized protocol.

Herbivory is ubiquitous. Despite being a potential driver of plant distribution and performance, herbivory remains largely undocumented. Some early attempts have been made to review, globally, how much leaf area is removed through insect feeding. Kozlov et al., in one of the most comprehensive reviews regarding global patterns of herbivory, have compiled published studies regarding foliar removal and sampled data on global herbivory levels using a standardized protocol. However, in the review by Kozlov et al., only 15 sampling sites, comprising 33 plant species, were evaluated in tropical areas around the globe. In Brazil, which ranks first in terms of plant biodiversity, with a total of 46,097 species, almost half (43%) being endemic, a single data point was sampled, covering only two plant species. In an attempt to increase knowledge regarding herbivory in tropical plant species and to provide the raw data needed to test general hypotheses related to plant-herbivore interactions across large spatial scales, we proposed a joint, collaborative network to evaluate tropical herbivory. This network allowed us to update and expand the data on insect herbivory in tropical and temperate plant species. Our data set, collected with a standardized protocol, covers 45 sampling sites from nine countries and includes leaf herbivory measurements of 57,239 leaves from 209 species of vascular plants belonging to 65 families from tropical and temperate regions. They expand previous data sets by including a total of 32 sampling sites from tropical areas around the globe, comprising 152 species, 146 of them being sampled in Brazil. For temperate areas, it includes 13 sampling sites, comprising 59 species. Thus, when compared to the most recent comprehensive review of insect herbivory (Kozlov et al.), our data set has increased the base of available data for the tropical plants more than 460% (from 33 to 152 species) and the Brazilian sampling was increased 7,300% (from 2 to 146 species). Data on precise levels of herbivory are presented for more than 57,000 leaves worldwide. There are no copyright restrictions. Please cite this paper when using the current data in publications; the authors request to be informed how the data is used in the publications.

Revisiting ecological dominance in arboreal ants: how dominant usage of nesting resources shapes community assembly.

Ecologically dominant species can shape the assembly of ecological communities via altering competitive outcomes. Moreover, these effects may be amplified under limited niche differentiation. Nevertheless, the influences of ecological dominance and niche differentiation on assembly are rarely considered together. Here, we provide a novel examination of dominance in a diverse arboreal ant community, defining dominance by the prevalent usage of nesting resources and addressing how it influences community assembly. We first used a series of quantitative observational and experimental studies to address the natural nesting ecology, colony incidence on surveyed trees, and level of dominance over newly available nesting resources by our focal species, Cephalotes pusillus. The experimental studies were then used further to examine whether C. pusillus shapes assembly via an influence on cavity usage by co-occurring species. C. pusillus was confirmed as a dominant user of cavity nesting resources, with highly generalized nesting ecology, occupying about 50% of the trees within the focal system, and accounting for more than a third of new cavity occupation in experiments. Our experiments showed further that the presence of C. pusillus was associated with modest effects on species richness, but significant decreases in cavity-occupation levels and significant shifts in the entrance-size usage by co-occurring species. These results indicate that C. pusillus, as a dominant user of nesting resources, shapes assembly at multiple levels. Broadly, our findings highlight that complex interactions between a dominant species and the resource-usage patterns of other species can underlie species assembly in diverse ecological communities.

Ant diversity in Neotropical savannas: Hierarchical processes acting at multiple spatial scales.

Understanding what creates and maintains macroscale biodiversity gradients is a central focus of ecological and evolutionary research. Spatial patterns in diversity are driven by a hierarchy of factors operating at multiple scales. Historical and climatic factors drive large-scale patterns of diversity by affecting the size of regional species pools, while habitat heterogeneity or microhabitat characteristics further influence species coexistence at small scales. We tested the degree to which the species-energy, historical factors, habitat heterogeneity and local environment hypotheses explain observed patterns of ant diversity across hierarchical spatial scales. We sampled ground-dwelling ants at 29 sites within a Neotropical savanna region, the Brazilian Cerrado. We measured species density - an abundance-dependent diversity metric - and rarefied species richness - an abundance-independent metric - at spatial scales with varying grain sizes. For each hypothesis, two correlates were used to predict ant diversity patterns: (a) species-energy: rainfall and productivity; (b) historical factors: historical variation in rainfall and refugial areas; (c) habitat heterogeneity: heterogeneity in greenness and diversity of land cover; and (d) local factors: contents of sand and coarse fragments in the soil. Ant diversity patterns correlated to net primary productivity and to the proportion of coarse fragments in the soil, corroborating the species-energy and local environment hypotheses, respectively. Soil negatively influenced species density, but not rarefied species richness, which was positively influenced by productivity. We found scale dependencies in the effects of soil/productivity on species density; productivity best predicted species density patterns at large scales, since sampling completeness offset the abundance-driven effects of soil. Considering abundance differences may help to discern the mechanisms underlying the relationship between macroscale diversity patterns and its ecological drivers. Plant productivity affected ant diversity independently of abundance, possibly by limiting the size of regional species pools. On the other hand, soil properties had an abundance-dependent effect on ant diversity, indicating a sampling mechanism. Our findings are consistent with predictions of the hierarchical theory of diversity. Large-scale patterns of productivity limit regional diversity, an effect that cascades down to finer spatial scales, where soil properties influence the number of coexisting species.

Functional richness shows spatial scale dependency in Pheidole ant assemblages from Neotropical savannas.

There is a growing recognition that spatial scale is important for understanding ecological processes shaping community membership, but empirical evidence on this topic is still scarce. Ecological processes such as environmental filtering can decrease functional differences among species and promote functional clustering of species assemblages, whereas interspecific competition can do the opposite. These different ecological processes are expected to take place at different spatial scales, with competition being more likely at finer scales and environmental filtering most likely at coarser scales. We used a comprehensive dataset on species assemblages of a dominant ant genus, Pheidole, in the Cerrado (savanna) biodiversity hotspot to ask how functional richness relates to species richness gradients and whether such relationships vary across spatial scales. Functional richness of Pheidole assemblages decreased with increasing species richness, but such relationship did not vary across different spatial scales. Species were more functionally dissimilar at finer spatial scales, and functional richness increased less than expected with increasing species richness. Our results indicate a tighter packing of the functional volume as richness increases and point out to a primary role for environmental filtering in shaping membership of Pheidole assemblages in Neotropical savannas. OPEN RESEARCH BADGES: This article has been awarded Open Materials, Open Data, Preregistered Research Designs Badges. All materials and data are publicly accessible via the Open Science Framework at https://doi.org/10.5061/dryad.31201jg.

FragSAD: A database of diversity and species abundance distributions from habitat fragments.

Habitat destruction is the single greatest anthropogenic threat to biodiversity. Decades of research on this issue have led to the accumulation of hundreds of data sets comparing species assemblages in larger, intact, habitats to smaller, more fragmented, habitats. Despite this, little synthesis or consensus has been achieved, primarily because of non-standardized sampling methodology and analyses of notoriously scale-dependent response variables (i.e., species richness). To be able to compare and contrast the results of habitat fragmentation on species' assemblages, it is necessary to have the underlying data on species abundances and sampling intensity, so that standardization can be achieved. To accomplish this, we systematically searched the literature for studies where abundances of species in assemblages (of any taxa) were sampled from many habitat patches that varied in size. From these, we extracted data from several studies, and contacted authors of studies where appropriate data were collected but not published, giving us 117 studies that compared species assemblages among habitat fragments that varied in area. Less than one-half (41) of studies came from tropical forests of Central and South America, but there were many studies from temperate forests and grasslands from all continents except Antarctica. Fifty-four of the studies were on invertebrates (mostly insects), but there were several studies on plants (15), birds (16), mammals (19), and reptiles and amphibians (13). We also collected qualitative information on the length of time since fragmentation. With data on total and relative abundances (and identities) of species, sampling effort, and affiliated meta-data about the study sites, these data can be used to more definitively test hypotheses about the role of habitat fragmentation in altering patterns of biodiversity. There are no copyright restrictions. Please cite this data paper and the associated Dryad data set if the data are used in publications.

Do an ecosystem engineer and environmental gradient act independently or in concert to shape juvenile plant communities? Tests with the leaf-cutter ant Atta laevigata in a Neotropical savanna.

BACKGROUND: Ecosystem engineers are species that transform habitats in ways that influence other species.While the impacts of many engineers have been well described, our understanding of how their impact varies along environmental gradients remains limited. Although disentangling the effects of gradients and engineers on biodiversity is complicated-the gradients themselves can be altered by engineers-doing so is necessary to advance conceptual and mathematical models of ecosystem engineering. We used leaf-cutter ants (Atta spp.) to investigate the relative influence of gradients and environmental engineers on the abundance and species richness of woody plants. METHODS: We conducted our research in South America's Cerrado. With a survey of plant recruits along a canopy cover gradient, and data on environmental conditions that influence plant recruitment, we fit statistical models that addressed the following questions: (1) Does A. laevigata modify the gradient in canopy cover found in our Cerrado site? (2) Do environmental conditions that influence woody plant establishment in the Cerrado vary with canopy cover or proximity to A. laevigata nests? (3) Do A. laevigata and canopy cover act independently or in concert to influence recruit abundance and species richness? RESULTS: We found that environmental conditions previously shown to influence plant establishment in the Cerrado varied in concert with canopy cover, but that ants are not modifying the cover gradient or cover over nests. However, ants are modifying other local environmental conditions, and the magnitude and spatial extent of these changes are consistent across the gradient. In contrast to prior studies, we found that ant-related factors (e.g., proximity to nests, ant changes in surface conditions), rather than canopy cover, had the strongest effect on the abundance of plant recruits. However, the diversity of plants was influenced by both the engineer and the canopy cover gradient. DISCUSSION: Atta laevigata in the Cerrado modify local conditions in ways that have strong but spatially restricted consequences for plant communities. We hypothesize that ants indirectly reduce seedling establishment by clearing litter and reducing soil moisture, which leads to seed and seedling desiccation. Altering soil nutrients could also reduce juvenile growth and survivorship; if so these indirect negative effects of engineering could exacerbate their direct effects of harvesting plants. The effects of Atta appear restricted to nest mounds, but they could be long-lasting because mounds persist long after a colony has died or migrated. Our results support the hypothesis that leaf-cutter ants play a dominant role in Cerrado plant demography. We suggest the ecological and economic footprint of these engineers may increase dramatically in coming decades due to the transformation of the Cerrado by human activities.

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.

Parasitism rate of Myzus persicae (Sulzer) by Diaeretiella rapae (McIntosh) in the presence of an alternative, resistant host

ABSTRACT The aphids Lipaphis pseudobrassicae (Davis) and Myzus persicae (Sulzer) (Hemiptera: Aphididae) are important Brassicaceae pests, occurring worldwide and causing significant damage to crops. Interspecific variations in the resistance to natural enemies can potentially impact the interaction among aphid populations. Here we evaluated the hypothesis of associational resistance by determining if the presence of resistant aphids (L. pseudobrassicae) reduces the rate of parasitism by Diaeretiella rapae (McIntosh) on non-resistant aphids (M. persicae). The experiment was conducted using collard green plants infested with M. persicae and L. pseudobrassicae either resistant or susceptible to D. rapae. The percentage of parasitism by D. rapae was greater on L. pseudobrassicae in the susceptible than in the resistant treatment, but parasitism rates on M. persicae did not differ between the treatments. There was no difference in average growth rate between M. persicae and susceptible L. pseudobrassicae populations, but resistant L. pseudobrassicae had greater growth rate than M. persicae. These results suggest that over a short period of time the presence of resistant L. pseudobrassicae does not affect the rate of parasitism by D. rapae on M. persicae.

Global climate change increases risk of crop yield losses and food insecurity in the tropical Andes.

One of the greatest current challenges to human society is ensuring adequate food production and security for a rapidly growing population under changing climatic conditions. Climate change, and specifically rising temperatures, will alter the suitability of areas for specific crops and cultivation systems. In order to maintain yields, farmers may be forced to change cultivation practices, the timing of cultivation, or even the type of crops grown. Alternatively, farmers can change the location where crops are cultivated (e.g., to higher elevations) to track suitable climates (in which case the plants will have to grow in different soils), as cultivated plants will otherwise have to tolerate warmer temperatures and possibly face novel enemies. We simulated these two last possible scenarios (for temperature increases of 1.3°C and 2.6°C) in the Peruvian Andes through a field experiment in which several traditionally grown varieties of potato and maize were planted at different elevations (and thus temperatures) using either the local soil or soil translocated from higher elevations. Maize production declined by 21%-29% in response to new soil conditions. The production of maize and potatoes declined by >87% when plants were grown under warmer temperatures, mainly as a result of the greater incidence of novel pests. Crop quality and value also declined under simulated migration and warming scenarios. We estimated that local farmers may experience severe economic losses of up to 2,300 US$ ha-1  yr-1 . These findings reveal that climate change is a real and imminent threat to agriculture and that there is a pressing need to develop effective management strategies to reduce yield losses and prevent food insecurity. Importantly, such strategies should take into account the influences of non-climatic and/or biotic factors (e.g., novel pests) on plant development.

Biogeography of mutualistic fungi cultivated by leafcutter ants.

Leafcutter ants propagate co-evolving fungi for food. The nearly 50 species of leafcutter ants (Atta, Acromyrmex) range from Argentina to the United States, with the greatest species diversity in southern South America. We elucidate the biogeography of fungi cultivated by leafcutter ants using DNA sequence and microsatellite-marker analyses of 474 cultivars collected across the leafcutter range. Fungal cultivars belong to two clades (Clade-A and Clade-B). The dominant and widespread Clade-A cultivars form three genotype clusters, with their relative prevalence corresponding to southern South America, northern South America, Central and North America. Admixture between Clade-A populations supports genetic exchange within a single species, Leucocoprinus gongylophorus. Some leafcutter species that cut grass as fungicultural substrate are specialized to cultivate Clade-B fungi, whereas leafcutters preferring dicot plants appear specialized on Clade-A fungi. Cultivar sharing between sympatric leafcutter species occurs frequently such that cultivars of Atta are not distinct from those of Acromyrmex. Leafcutters specialized on Clade-B fungi occur only in South America. Diversity of Clade-A fungi is greatest in South America, but minimal in Central and North America. Maximum cultivar diversity in South America is predicted by the Kusnezov-Fowler hypothesis that leafcutter ants originated in subtropical South America and only dicot-specialized leafcutter ants migrated out of South America, but the cultivar diversity becomes also compatible with a recently proposed hypothesis of a Central American origin by postulating that leafcutter ants acquired novel cultivars many times from other nonleafcutter fungus-growing ants during their migrations from Central America across South America. We evaluate these biogeographic hypotheses in the light of estimated dates for the origins of leafcutter ants and their cultivars.

The biodiversity cost of carbon sequestration in tropical savanna.

Tropical savannas have been increasingly viewed as an opportunity for carbon sequestration through fire suppression and afforestation, but insufficient attention has been given to the consequences for biodiversity. To evaluate the biodiversity costs of increasing carbon sequestration, we quantified changes in ecosystem carbon stocks and the associated changes in communities of plants and ants resulting from fire suppression in savannas of the Brazilian Cerrado, a global biodiversity hotspot. Fire suppression resulted in increased carbon stocks of 1.2 Mg ha-1 year-1 since 1986 but was associated with acute species loss. In sites fully encroached by forest, plant species richness declined by 27%, and ant richness declined by 35%. Richness of savanna specialists, the species most at risk of local extinction due to forest encroachment, declined by 67% for plants and 86% for ants. This loss highlights the important role of fire in maintaining biodiversity in tropical savannas, a role that is not reflected in current policies of fire suppression throughout the Brazilian Cerrado. In tropical grasslands and savannas throughout the tropics, carbon mitigation programs that promote forest cover cannot be assumed to provide net benefits for conservation.

Rediscovery of the enigmatic fungus-farming ant "Mycetosoritis" asper Mayr (Hymenoptera: Formicidae): Implications for taxonomy, phylogeny, and the evolution of agriculture in ants.

We report the rediscovery of the exceedingly rarely collected and enigmatic fungus-farming ant species Mycetosoritis asper. Since the description of the type specimen in 1887, only four additional specimens are known to have been added to the world's insect collections. Its biology is entirely unknown and its phylogenetic position within the fungus-farming ants has remained puzzling due to its aberrant morphology. In 2014 we excavated and collected twenty-one colonies of M. asper in the Floresta Nacional de Chapecó in Santa Catarina, Brazil. We describe here for the first time the male and larva of the species and complement the previous descriptions of both the queen and the worker. We describe, also for the first time, M. asper biology, nest architecture, and colony demographics, and identify its fungal cultivar. Molecular phylogenetic analyses indicate that both M. asper and M. clorindae are members of the genus Cyphomyrmex, which we show to be paraphyletic as currently defined. More precisely, M. asper is a member of the Cyphomyrmex strigatus group, which we also show to be paraphyletic with respect to the genus Mycetophylax. Based on these results, and in the interest of taxonomic stability, we transfer the species M. asper, M. clorindae, and all members of the C. strigatus group to the genus Mycetophylax, the oldest available name for this clade. Based on ITS sequence data, Mycetophylax asper practices lower agriculture, cultivating a fungal species that belongs to lower-attine fungal Clade 2, subclade F.

Co-occurrence patterns in a diverse arboreal ant community are explained more by competition than habitat requirements.

A major goal of community ecology is to identify the patterns of species associations and the processes that shape them. Arboreal ants are extremely diverse and abundant, making them an interesting and valuable group for tackling this issue. Numerous studies have used observational data of species co-occurrence patterns to infer underlying assembly processes, but the complexity of these communities has resulted in few solid conclusions. This study takes advantage of an observational dataset that is unusually well-structured with respect to habitat attributes (tree species, tree sizes, and vegetation structure), to disentangle different factors influencing community organization. In particular, this study assesses the potential role of interspecific competition and habitat selection on the distribution patterns of an arboreal ant community by incorporating habitat attributes into the co-occurrence analyses. These findings are then contrasted against species traits, to explore functional explanations for the identified community patterns. We ran a suite of null models, first accounting only for the species incidence in the community and later incorporating habitat attributes in the null models. We performed analyses with all the species in the community and then with only the most common species using both a matrix-level approach and a pairwise-level approach. The co-occurrence patterns did not differ from randomness in the matrix-level approach accounting for all ant species in the community. However, a segregated pattern was detected for the most common ant species. Moreover, with the pairwise approach, we found a significant number of negative and positive pairs of species associations. Most of the segregated associations appear to be explained by competitive interactions between species, not habitat affiliations. This was supported by comparisons of species traits for significantly associated pairs. These results suggest that competition is the most important influence on the distribution patterns of arboreal ants within the focal community. Habitat attributes, in contrast, showed no significant influence on the matrix-wide results and affected only a few associations. In addition, the segregated pairs shared more biological characteristic in common than the aggregated and random ones.

New distribution records of the savanna specialist fungus-farming ant Cyatta Sosa-Calvo et al. (Hymenoptera: Formicidae: Myrmicinae).

BACKGROUND: The fungus-farming ant genus Cyatta (Formicidae: Myrmicinae) is represented by a single species, C. abscondita Sosa-Calvo et al., known from a few localities in Brazil (in the states of Ceará, Minas Gerais, São Paulo and the Distrito Federal), and a single locality in the Misiones province, Argentina. Cyatta is known to occur predominantly in savanna habitats and occasionally in the transition zones between the Atlantic Forest and Cerrado. NEW INFORMATION: The new records reported here significantly expand the previously known distribution of Cyatta abscondita and provide further support for the intimate relation between this species and the savannas of South America. We report the first occurrence of the genus in southern Brazil (Paraná state) and the westernmost occurrence (Bolivia) of Cyatta abscondita, which extend its distribution approximately 1450 km to the west. Finally, we discuss the importance of mapping inconspicuous species in order to develop strategies for protecting endangered areas and to increase our understanding of the evolutionary history of organisms and biomes.

Extrafloral nectaries have a limited effect on the structure of arboreal ant communities in a Neotropical savanna.

How environmental contexts shape the strength of species interactions, and their influence on community structure, remains a key focus for the field of community ecology. In particular, the extent to which local competitive interactions impact community structure, and whether this differs across contexts, persists as a general issue that is unresolved across a broad range of animal systems. Studies of arboreal ants have shown that competitive interactions over carbon-rich exudates from extrafloral nectaries (EFNs) and homopteran aggregations can have positive and negative effects on the local abundances of individual species. Nevertheless, it is still unclear the extent to which these local effects scale to community-level effects. Here we address the role of food from extrafloral nectaries on the structure of arboreal ant communities in a savanna of central Brazil. We did this with a combination of a diversity survey across tree species with and without EFNs, a repeated survey at times of peak EFN activity, and testing of our survey findings with two experimental manipulations of nectar availability that also provided supplementary nesting cavities. Species richness, but not composition, differed significantly between trees with and without EFNs. However, trees with EFNs had, on average, only 9% more species than those without EFNs. Furthermore, ant species richness did not differ significantly between periods of high and low EFN activity. Although nectar supplementation significantly affected nest occupation rates, this difference was seen solely in. the experiment with a massive supply of nectar and there was no effect on total ant richness or identity of the focal assemblages. Our findings suggest that the effects of extrafloral nectar on the abundances of arboreal ants at local scales do not scale to a strong structuring force at the community level. We suggest that this is most likely due to a lack of specificity of community members for EFN tree species, and the diffuse temporal and spatial nature of the availability of active EFNs. These properties mean that observable short-lived activity and competition over particular EFNs does not ultimately drive lasting changes in the associated assemblage of species, and therefore, the community as a whole.