Arthropod decline in grasslands and forests is associated with landscape-level drivers.

Recent reports of local extinctions of arthropod species1, and of massive declines in arthropod biomass2, point to land-use intensification as a major driver of decreasing biodiversity. However, to our knowledge, there are no multisite time series of arthropod occurrences across gradients of land-use intensity with which to confirm causal relationships. Moreover, it remains unclear which land-use types and arthropod groups are affected, and whether the observed declines in biomass and diversity are linked to one another. Here we analyse data from more than 1 million individual arthropods (about 2,700 species), from standardized inventories taken between 2008 and 2017 at 150 grassland and 140 forest sites in 3 regions of Germany. Overall gamma diversity in grasslands and forests decreased over time, indicating loss of species across sites and regions. In annually sampled grasslands, biomass, abundance and number of species declined by 67%, 78% and 34%, respectively. The decline was consistent across trophic levels and mainly affected rare species; its magnitude was independent of local land-use intensity. However, sites embedded in landscapes with a higher cover of agricultural land showed a stronger temporal decline. In 30 forest sites with annual inventories, biomass and species number-but not abundance-decreased by 41% and 36%, respectively. This was supported by analyses of all forest sites sampled in three-year intervals. The decline affected rare and abundant species, and trends differed across trophic levels. Our results show that there are widespread declines in arthropod biomass, abundance and the number of species across trophic levels. Arthropod declines in forests demonstrate that loss is not restricted to open habitats. Our results suggest that major drivers of arthropod decline act at larger spatial scales, and are (at least for grasslands) associated with agriculture at the landscape level. This implies that policies need to address the landscape scale to mitigate the negative effects of land-use practices.

Stratification and recovery time jointly shape ant functional reassembly in a neotropical forest.

Microhabitat differentiation of species communities such as vertical stratification in tropical forests contributes to species coexistence and thus biodiversity. However, little is known about how the extent of stratification changes during forest recovery and influences community reassembly. Environmental filtering determines community reassembly in time (succession) and in space (stratification), hence functional and phylogenetic composition of species communities are highly dynamic. It is poorly understood if and how these two concurrent filters-forest recovery and stratification-interact. In a tropical forest chronosequence in Ecuador spanning 34 years of natural recovery, we investigated the recovery trajectory of ant communities in three overlapping strata (ground, leaf litter, lower tree trunk) by quantifying 13 traits, as well as the functional and phylogenetic diversity of the ants. We expected that functional and phylogenetic diversity would increase with recovery time and that each ant community within each stratum would show a distinct functional reassembly. We predicted that traits related to ant diet would show divergent trajectories reflecting an increase in niche differentiation with recovery time. On the other hand, traits related to the abiotic environment were predicted to show convergent trajectories due to a more similar microclimate across strata with increasing recovery age. Most of the functional traits and the phylogenetic diversity of the ants were clearly stratified, confirming previous findings. However, neither functional nor phylogenetic diversity increased with recovery time. Community-weighted trait means had complex relationships to recovery time and the majority were shaped by a statistical interaction between recovery time and stratum, confirming our expectations. However, most trait trajectories converged among strata with increasing recovery time regardless of whether they were related to ant diet or environmental conditions. We confirm the hypothesized interaction among environmental filters during the functional reassembly in tropical forests. Communities in individual strata respond differently to recovery, and possible filter mechanisms likely arise from both abiotic (e.g. microclimate) and biotic (e.g. diet) conditions. Since vertical stratification is prevalent across animal and plant taxa, our results highlight the importance of stratum-specific analysis in dynamic ecosystems and may generalize beyond ants.

Experiments are needed to quantify the main causes of insect decline.

Sparked by reports of insect declines of unexpected extent, there has been a surge in the compilation and analysis of insect time series data. While this effort has led to valuable databases, disagreement remains as to whether, where and why insects are declining. The 'why' question is particularly important because successful insect conservation will need to address the most important drivers of decline. Despite repeated calls for more long-term data, new time series will have to run for decades to quantitatively surpass those currently available. Here we argue that experimentation in addition to quantitative analysis of existing data is needed to identify the most important drivers of insect decline. While most potential drivers of insect population change are likely to have already been identified, their relative importance is largely unknown. Researchers should thus unite and use statistical insight to set up suitable experiments to be able to rank drivers by their importance. Such a coordinated effort is needed to produce the knowledge necessary for conservation action and will also result in increased monitoring and new time series.

Forest gaps increase true bug diversity by recruiting open land species.

Forests canopy gaps play an important role in forest ecology by driving the forest mosaic cycle and creating conditions for rapid plant reproduction and growth. The availability of young plants, which represent resources for herbivores, and modified environmental conditions with greater availability of light and higher temperatures, promote the colonization of animals. Remarkably, the role of gaps on insect communities has received little attention and the source of insects colonizing gaps has not been studied comprehensively. Using a replicated full-factorial forest experiment (treatments: Gap; Gap + Deadwood; Deadwood; Control), we show that following gap creation, there is a rapid change in the true bug (Heteroptera) community structure, with an increase in species that are mainly recruited from open lands. Compared with closed-canopy treatments (Deadwood and Control), open canopy treatments (Gap and Gap + Deadwood) promoted an overall increase in species (+ 59.4%, estimated as number of species per plot) and individuals (+ 76.3%) of true bugs, mainly herbivores and species associated to herbaceous vegetation. Community composition also differed among treatments, and all 17 significant indicator species (out of 117 species in total) were associated with the open canopy treatments. Based on insect data collected in grasslands and forests over an 11-year period, we found that the species colonizing experimental gaps had greater body size and a greater preference for open vegetation. Our results indicate that animal communities that assemble following gap creation contain a high proportion of habitat generalists that not occurred in closed forests, contributing significantly to overall diversity in forest mosaics.

Ecosystem restoration strengthens pollination network resilience and function.

Land degradation results in declining biodiversity and the disruption of ecosystem functioning worldwide, particularly in the tropics. Vegetation restoration is a common tool used to mitigate these impacts and increasingly aims to restore ecosystem functions rather than species diversity. However, evidence from community experiments on the effect of restoration practices on ecosystem functions is scarce. Pollination is an important ecosystem function and the global decline in pollinators attenuates the resistance of natural areas and agro-environments to disturbances. Thus, the ability of pollination functions to resist or recover from disturbance (that is, the functional resilience) may be critical for ensuring a successful restoration process. Here we report the use of a community field experiment to investigate the effects of vegetation restoration, specifically the removal of exotic shrubs, on pollination. We analyse 64 plant-pollinator networks and the reproductive performance of the ten most abundant plant species across four restored and four unrestored, disturbed mountaintop communities. Ecosystem restoration resulted in a marked increase in pollinator species, visits to flowers and interaction diversity. Interactions in restored networks were more generalized than in unrestored networks, indicating a higher functional redundancy in restored communities. Shifts in interaction patterns had direct and positive effects on pollination, especially on the relative and total fruit production of native plants. Pollinator limitation was prevalent at unrestored sites only, where the proportion of flowers producing fruit increased with pollinator visitation, approaching the higher levels seen in restored plant communities. Our results show that vegetation restoration can improve pollination, suggesting that the degradation of ecosystem functions is at least partially reversible. The degree of recovery may depend on the state of degradation before restoration intervention and the proximity to pollinator source populations in the surrounding landscape. We demonstrate that network structure is a suitable indicator for pollination quality, highlighting the usefulness of interaction networks in environmental management.

Stochasticity Leads to Coexistence of Generalists and Specialists in Assembling Mutualistic Communities.

AbstractSpecies interaction networks are subject to natural and anthropogenic disturbances that lead to their disassembly, while natural regeneration or restoration efforts facilitate their reassembly. Previous models for assembling ecological networks did not include stochasticity at the level of population dynamics (e.g., demographic noise, environmental noise) and focused mainly on food webs. Here, we present a model for the assembly of mutualistic bipartite networks, such as plant-pollinator networks, and examine the influence of demographic noise on the trajectory of species and strategy diversity, that is, the range of present strategies from specialism to generalism. We find that assembled communities show at intermediate assembly stages a maximum of species diversity and of average generalization. Our model thus provides a mechanism for nonlinear, hump-shaped diversity trajectories at intermediate succession, consistent with the intermediate disturbance hypothesis. Long-term coexistence of specialists and generalists emerges only in the presence of demographic noise and is due to a persistent species turnover. These findings highlight the importance of stochasticity for maintaining long-term diversity.

Rapid ant community reassembly in a Neotropical forest: Recovery dynamics and land-use legacy.

Regrowing secondary forests dominate tropical regions today, and a mechanistic understanding of their recovery dynamics provides important insights for conservation. In particular, land-use legacy effects on the fauna have rarely been investigated. One of the most ecologically dominant and functionally important animal groups in tropical forests are the ants. Here, we investigated the recovery of ant communities in a forest-agricultural habitat mosaic in the Ecuadorian Chocó region. We used a replicated chronosequence of previously used cacao plantations and pastures with 1-34 years of regeneration time to study the recovery dynamics of species communities and functional diversity across the two land-use legacies. We compared two independent components of responses on these community properties: resistance, which is measured as the proportion of an initial property that remains following the disturbance; and resilience, which is the rate of recovery relative to its loss. We found that compositional and trait structure similarity to old-growth forest communities increased with regeneration age, whereas ant species richness remained always at a high level along the chronosequence. Land-use legacies influenced species composition, with former cacao plantations showing higher resemblance to old-growth forests than former pastures along the chronosequence. While resistance was low for species composition and high for species richness and traits, all community properties had similarly high resilience. In essence, our results show that ant communities of the Chocó recovery rapidly, with former cacao reaching predicted old-growth forest community levels after 21 years and pastures after 29 years. Recovery in this community was faster than reported from other ecosystems and was likely facilitated by the low-intensity farming in agricultural sites and their proximity to old-growth forest remnants. Our study indicates the great recovery potential for this otherwise highly threatened biodiversity hotspot.

Negative effects of forest gaps on dung removal in a full-factorial experiment.

Ecosystem functioning may directly or indirectly-via change in biodiversity-respond to land use. Dung removal is an important ecosystem function central for the decomposition of mammal faeces, including secondary seed dispersal and improved soil quality. Removal usually increases with dung beetle diversity and biomass. In forests, dung removal can vary with structural variables that are, however, often interrelated, making experiments necessary to understand the role of single variables on ecosystem functions. How gaps and deadwood, two main outcomes of forest management influence dung removal, is unknown. We tested if dung removal responds to gap creation and deadwood provisioning or if treatment effects are mediated via responses of dung beetles. We expected lower removal rates in gaps due to lower dung beetle biomass and diversity. We sampled dung beetles and measured dung removal in a highly-replicated full-factorial forest experiment established at 29 sites in three regions of Germany (treatments: Gap, Gap + Deadwood, Deadwood, Control). All gaps were experimentally created and had a diameter of around 30 m. Dung beetle diversity, biomass and dung removal were each lower in gaps than in controls. Dung removal decreased from 61.9% in controls to 48.5% in gaps, irrespective of whether or not the gap had deadwood. This treatment effect was primarily driven by dung beetle biomass but not diversity. Furthermore, dung removal was reduced to 56.9% in the deadwood treatment. Our findings are not consistent with complementarity effects of different dung beetle species linked to biodiversity-ecosystem functioning relationships that have been shown in several ecosystems. In contrast, identity effects can be pronounced: gaps reduced the abundance of a large-bodied key forest species (Anoplotrupes stercorosus), without compensatory recruitment of open land species. While gaps and deadwood are important for many forest organisms, dung beetles and dung removal respond negatively. Our results exemplify how experiments can contribute to test hypotheses on the interrelation between land use, biodiversity and ecosystem functioning.

Unravelling insect declines: can space replace time?

Temporal trends in insect numbers vary across studies and habitats, but drivers are poorly understood. Suitable long-term data are scant and biased, and interpretations of trends remain controversial. By contrast, there is substantial quantitative evidence for drivers of spatial variation. From observational and experimental studies, we have gained a profound understanding of where insect abundance and diversity is higher-and identified underlying environmental conditions, resource change and disturbances. We thus propose an increased consideration of spatial evidence in studying the causes of insect decline. This is because for most time series available today, the number of sites and thus statistical power strongly exceed the number of years studied. Comparisons across sites allow quantifying insect population risks, impacts of land use, habitat destruction, restoration or management, and stressors such as chemical and light pollution, pesticides, mowing or harvesting, climatic extremes or biological invasions. Notably, drivers may not have to change in intensity to have long-term effects on populations, e.g. annually repeated disturbances or mortality risks such as those arising from agricultural practices. Space-for-time substitution has been controversially debated. However, evidence from well-replicated spatial data can inform on urgent actions required to halt or reverse declines-to be implemented in space.

Land-use intensification causes multitrophic homogenization of grassland communities.

Land-use intensification is a major driver of biodiversity loss. Alongside reductions in local species diversity, biotic homogenization at larger spatial scales is of great concern for conservation. Biotic homogenization means a decrease in ß-diversity (the compositional dissimilarity between sites). Most studies have investigated losses in local (α)-diversity and neglected biodiversity loss at larger spatial scales. Studies addressing ß-diversity have focused on single or a few organism groups (for example, ref. 4), and it is thus unknown whether land-use intensification homogenizes communities at different trophic levels, above- and belowground. Here we show that even moderate increases in local land-use intensity (LUI) cause biotic homogenization across microbial, plant and animal groups, both above- and belowground, and that this is largely independent of changes in α-diversity. We analysed a unique grassland biodiversity dataset, with abundances of more than 4,000 species belonging to 12 trophic groups. LUI, and, in particular, high mowing intensity, had consistent effects on ß-diversity across groups, causing a homogenization of soil microbial, fungal pathogen, plant and arthropod communities. These effects were nonlinear and the strongest declines in ß-diversity occurred in the transition from extensively managed to intermediate intensity grassland. LUI tended to reduce local α-diversity in aboveground groups, whereas the α-diversity increased in belowground groups. Correlations between the ß-diversity of different groups, particularly between plants and their consumers, became weaker at high LUI. This suggests a loss of specialist species and is further evidence for biotic homogenization. The consistently negative effects of LUI on landscape-scale biodiversity underscore the high value of extensively managed grasslands for conserving multitrophic biodiversity and ecosystem service provision. Indeed, biotic homogenization rather than local diversity loss could prove to be the most substantial consequence of land-use intensification.

Biodiversity at multiple trophic levels is needed for ecosystem multifunctionality.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Seeing through the static: the temporal dimension of plant-animal mutualistic interactions.

Most studies of plant-animal mutualistic networks have come from a temporally static perspective. This approach has revealed general patterns in network structure, but limits our ability to understand the ecological and evolutionary processes that shape these networks and to predict the consequences of natural and human-driven disturbance on species interactions. We review the growing literature on temporal dynamics of plant-animal mutualistic networks including pollination, seed dispersal and ant defence mutualisms. We then discuss potential mechanisms underlying such variation in interactions, ranging from behavioural and physiological processes at the finest temporal scales to ecological and evolutionary processes at the broadest. We find that at the finest temporal scales (days, weeks, months) mutualistic interactions are highly dynamic, with considerable variation in network structure. At intermediate scales (years, decades), networks still exhibit high levels of temporal variation, but such variation appears to influence network properties only weakly. At the broadest temporal scales (many decades, centuries and beyond), continued shifts in interactions appear to reshape network structure, leading to dramatic community changes, including loss of species and function. Our review highlights the importance of considering the temporal dimension for understanding the ecology and evolution of complex webs of mutualistic interactions.

Landscape complexity promotes resilience of biological pest control to climate change.

Increased climate variability as a result of anthropogenic climate change can threaten the functioning of ecosystem services. However, diverse responses to climate change among species (response diversity) can provide ecosystems with resilience to this growing threat. Measuring and managing response diversity and resilience to global change are key ecological challenges. Here, we develop a novel index of climate resilience of ecosystem services, exemplified by the thermal resilience of predator communities providing biological pest control. Field assays revealed substantial differences in the temperature-dependent activity of predator species and indices of thermal resilience varied among predator communities occupying different fields. Predator assemblages with higher thermal resilience provided more stable pest control in microcosms where the temperature was experimentally varied, confirming that the index of thermal resilience developed here is linked to predator function. Importantly, complex landscapes containing a high number of non-crop habitat patches were more likely to contain predator communities with high thermal resilience. Thus, the conservation and restoration of non-crop habitats in agricultural landscapes-practices known to strengthen natural pest suppression under current conditions-will also confer resilience in ecosystem service provisioning to climate change.

A remarkable legion of guests: Diversity and host specificity of army ant symbionts.

Tropical rainforests are among the most diverse biomes on Earth. While species inventories are far from complete for any tropical rainforest, even less is known about the intricate species interactions that form the basis of these ecological communities. One fascinating but poorly studied example are the symbiotic associations between army ants and their rich assemblages of parasitic arthropod guests. Hundreds of these guests, or myrmecophiles, have been taxonomically described. However, because previous work has mainly been based on haphazard collections from disjunct populations, it remains challenging to define species boundaries. We therefore know little about the species richness, abundance and host specificity of most guests in any given population, which is crucial to understand co-evolutionary and ecological dynamics. Here, we report a quantitative community survey of myrmecophiles parasitizing the six sympatric Eciton army ant species in a Costa Rican rainforest. Combining DNA barcoding with morphological identification of over 2,000 specimens, we discovered 62 species, including 49 beetles, 11 flies, one millipede and one silverfish. At least 14 of these species were new to science. Ecological network analysis revealed a clear signal of host partitioning, and each Eciton species was host to both specialists and generalists. These varying degrees in host specificities translated into a moderate level of network specificity, highlighting the system's level of biotic pluralism in terms of biodiversity and interaction diversity. By providing vouchered DNA barcodes for army ant guest species, this study provides a baseline for future work on co-evolutionary and ecological dynamics in these species-rich host-symbiont networks across the Neotropical realm.

Multiple phenotypic traits as triggers of host attacks towards ant symbionts: body size, morphological gestalt, and chemical mimicry accuracy.

BACKGROUND: Ant colonies are plagued by a diversity of arthropod guests, which adopt various strategies to avoid or to withstand host attacks. Chemical mimicry of host recognition cues is, for example, a common integration strategy of ant guests. The morphological gestalt and body size of ant guests have long been argued to also affect host hostility, but quantitative studies testing these predictions are largely missing. We here evaluated three guest traits as triggers of host aggression-body size, morphological gestalt, and accuracy in chemical mimicry-in a community of six Eciton army ant species and 29 guest species. We quantified ant aggression towards 314 guests in behavioral assays and, for the same individuals, determined their body size and their accuracy in mimicking ant cuticular hydrocarbon (CHC) profiles. We classified guests into the following gestalts: protective, myrmecoid, staphylinid-like, phorid-like, and larval-shaped. We expected that (1) guests with lower CHC mimicry accuracy are more frequently attacked; (2) larger guests are more frequently attacked; (3) guests of different morphological gestalt receive differing host aggression levels. RESULTS: Army ant species had distinct CHC profiles and accuracy of mimicking these profiles was variable among guests, with many species showing high mimicry accuracy. Unexpectedly, we did not find a clear relationship between chemical host similarity and host aggression, suggesting that other symbiont traits need to be considered. We detected a relationship between the guests' body size and the received host aggression, in that diminutive forms were rarely attacked. Our data also indicated that morphological gestalt might be a valuable predictor of host aggression. While most ant-guest encounters remained peaceful, host behavior still differed towards guests in that ant aggression was primarily directed towards those guests possessing a protective or a staphylinid-like gestalt. CONCLUSION: We demonstrate that CHC mimicry accuracy does not necessarily predict host aggression towards ant symbionts. Exploitation mechanisms are diverse, and we conclude that, besides chemical mimicry, other factors such as the guests' morphological gestalt and especially their body size might be important, yet underrated traits shaping the level of host hostility against social insect symbionts.

How land-use intensity affects sexual and parthenogenetic oribatid mites in temperate forests and grasslands in Germany.

Intensive land use has been shown to alter the composition and functioning of soil communities. Due to their low dispersal ability, oribatid mites are particularly vulnerable to land-use intensification and species which are not adjusted to management-related disturbances become less abundant. We investigated how different land-use parameters in forests and grasslands affect oribatid mite diversity and abundance, with a focus on: (1) species-level impacts, by classifying species as increasing ('winners') or decreasing ('losers') in abundance with higher land-use intensity, and (2) reproductive impact, by investigating whether sexual and parthenogenetic species react differently. We collected 32,542 adult oribatid mites in 60 forests and grasslands of known land-use intensity in two regions of Germany. Diversity and total abundance as well as the proportion of sexual species were higher in forests than in grasslands. Diversity declined with higher land-use intensity in forests, but increased with higher mowing and fertilization in grasslands. Depending on land-use parameter and region, abundance either declined or remained unaffected by increasing intensity. Gravidity was higher in sexual than in parthenogenetic species and sexuals had 1.6× more eggs per gravid female. Proportions of sexual species and gravid females decreased with land-use intensity in forests, but increased with mowing in grasslands. At the species level, 75% of sexuals and 87.5% of parthenogens were 'losers' of higher percentages of dead wood originating from management-related disturbances. Across land-use parameters and habitats, a similar proportion of sexual and parthenogenetic oribatid mite species were 'losers' of high land-use intensity. However, 'winner' species were more common among sexuals.

Resilience of ecosystem processes: a new approach shows that functional redundancy of biological control services is reduced by landscape simplification.

Functional redundancy can increase the resilience of ecosystem processes by providing insurance against species loss and the effects of abundance fluctuations. However, due to the difficulty of assessing individual species' contributions and the lack of a metric allowing for a quantification of redundancy within communities, few attempts have been made to estimate redundancy for individual ecosystem processes. We present a new method linking interaction metrics with metabolic theory that allows for a quantification of redundancy at the level of ecosystem processes. Using this approach, redundancy in the predation on aphids and other prey by natural enemies across a landscape heterogeneity gradient was estimated. Functional redundancy of predators was high in heterogeneous landscapes, low in homogeneous landscapes and scaled with predator specialisation. Our approach allows quantifying functional redundancy within communities and can be used to assess the role of functional redundancy across a wide variety of ecosystem processes and environmental factors.

Tree diversity increases robustness of multi-trophic interactions.

Multi-trophic interactions maintain critical ecosystem functions. Biodiversity is declining globally, while responses of trophic interactions to biodiversity change are largely unclear. Thus, studying responses of multi-trophic interaction robustness to biodiversity change is crucial for understanding ecosystem functioning and persistence. We investigate plant-Hemiptera (antagonism) and Hemiptera-ant (mutualism) interaction networks in response to experimental manipulation of tree diversity. We show increased diversity at both higher trophic levels (Hemiptera and ants) and increased robustness through redundancy of lower level species of multi-trophic interactions when tree diversity increased. Hemiptera and ant diversity increased with tree diversity through non-additive diversity effects. Network analyses identified that tree diversity also increased the number of tree and Hemiptera species used by Hemiptera and ant species, and decreased the specialization on lower trophic level species in both mutualistic and antagonist interactions. Our results demonstrate that bottom-up effects of tree diversity ascend through trophic levels regardless of interaction type. Thus, local tree diversity is a key driver of multi-trophic community diversity and interaction robustness in forests.

Species-level predation network uncovers high prey specificity in a Neotropical army ant community.

Army ants are among the top arthropod predators and considered keystone species in tropical ecosystems. During daily mass raids with many thousand workers, army ants hunt live prey, likely exerting strong top-down control on prey species. Many tropical sites exhibit a high army ant species diversity (>20 species), suggesting that sympatric species partition the available prey niches. However, whether and to what extent this is achieved has not been intensively studied yet. We therefore conducted a large-scale diet survey of a community of surface-raiding army ants at La Selva Biological Station in Costa Rica. We systematically collected 3,262 prey items from eleven army ant species (genera Eciton, Nomamyrmex and Neivamyrmex). Prey items were classified as ant prey or non-ant prey. The prey nearly exclusively consisted of other ants (98%), and most booty was ant brood (87%). Using morphological characters and DNA barcoding, we identified a total of 1,103 ant prey specimens to the species level. One hundred twenty-nine ant species were detected among the army ant prey, representing about 30% of the known local ant diversity. Using weighted bipartite network analyses, we show that prey specialization in army ants is unexpectedly high and prey niche overlap very small. Besides food niche differentiation, we uncovered a spatiotemporal niche differentiation in army ant raid activity. We discuss competition-driven multidimensional niche differentiation and predator-prey arms races as possible mechanisms underlying prey specialization in army ants. By combining systematic prey sampling with species-level prey identification and network analyses, our integrative approach can guide future research by portraying how predator-prey interactions in complex communities can be reliably studied, even in cases where morphological prey identification is infeasible.

Under pressure: force resistance measurements in box mites (Actinotrichida, Oribatida).

BACKGROUND: Mechanical defenses are very common and diverse in prey species, for example in oribatid mites. Here, the probably most complex form of morphological defense is known as ptychoidy, that enables the animals to completely retract the appendages into a secondary cavity and encapsulate themselves. The two groups of ptychoid mites constituting the Ptyctima, i.e. Euphthiracaroidea and Phthiracaroidea, have a hardened cuticle and are well protected against similar sized predators. Euphthiracaroidea additionally feature predator-repelling secretions. Since both taxa evolved within the glandulate group of Oribatida, the question remains why Phthiracaroidea lost this additional protection. In earlier predation bioassays, chemically disarmed specimens of Euphthiracaroidea were cracked by the staphylinid beetle Othius punctulatus, whereas equally sized specimens of Phthiracaroidea survived. We thus hypothesized that Phthiracaroidea can withstand significantly more force than Euphthiracaroidea and that the specific body form in each group is key in understanding the loss of chemical defense in Phthiracaroidea. To measure force resistance, we adapted the principle of machines applying compressive forces for very small animals and tested the two ptyctimous taxa as well as the soft-bodied mite Archegozetes longisetosus. RESULTS: Some Phthiracaroidea individuals sustained about 560,000 times their body weight. Their mean resistance was about three times higher, and their mean breaking point in relation to body weight nearly two times higher than Euphthiracaroidea individuals. The breaking point increased with body weight and differed significantly between the two taxa. Across taxa, the absolute force resistance increased sublinearly (with a 0.781 power term) with the animal's body weight. Force resistance of A. longisetosus was inferior in all tests (about half that of Euphthiracaroidea after accounting for body weight). As an important determinant of mechanical resistance in ptychoid mites, the individuals' cuticle thickness increased sublinearly with body diameter and body mass as well and did not differ significantly between the taxa. CONCLUSION: We showed the feasibility of the force resistance measurement method, and our results were consistent with the hypothesis that Phthiracaroidea compensated its lack of chemical secretions by a heavier mechanical resistance based on a different body form and associated build-up of hemolymph pressure (defensive trade-off).