Mammal dung-dung beetle trophic networks: an improved method based on gut-content DNA.

Background: Dung beetles provide many important ecosystem services, including dung decomposition, pathogen control, soil aeration, and secondary seed dispersal. Yet, the biology of most dung beetles remains unknown. Natural diets are poorly studied, partly because previous research has focused on choice or attraction experiments using few, easily accessible dung types from zoo animals, farm animals, or humans. This way, many links within natural food webs have certainly been missed. In this work, we aimed to establish a protocol to analyze the natural diets of dung beetles using DNA gut barcoding. Methods: First, the feasibility of gut-content DNA extraction and amplification of 12s rDNA from six different mammal dung types was tested in the laboratory. We then applied the method to beetles caught in pitfall traps in Ecuador and Germany by using 12s rDNA primers. For a subset of the dung beetles caught in the Ecuador sampling, we also used 16s rDNA primers to see if these would improve the number of species we could identify. We predicted the likelihood of amplifying DNA using gut fullness, DNA concentration, PCR primer, collection method, and beetle species as predictor variables in a dominance analysis. Based on the gut barcodes, we generated a dung beetle-mammal network for both field sites (Ecuador and Germany) and analyzed the levels of network specificity. Results: We successfully amplified mammal DNA from dung beetle gut contents for 128 specimens, which included such prominent species as Panthera onca (jaguar) and Puma concolor (puma). The overall success rate of DNA amplification was 53%. The best predictors for amplification success were gut fullness and DNA concentration, suggesting the success rate can be increased by focusing on beetles with a full gut. The mammal dung-dung beetle networks differed from purely random network models and showed a moderate degree of network specialization (H2': Ecuador = 0.49; Germany = 0.41). Conclusion: We here present a reliable method of extracting and amplifying gut-content DNA from dung beetles. Identifying mammal dung via DNA reference libraries, we created mammal dung-dung beetle trophic networks. This has benefits over previous methods because we inventoried the natural mammal dung resources of dung beetles instead of using artificial mammal baits. Our results revealed higher levels of specialization than expected and more rodent DNA than expected in Germany, suggesting that the presented method provides more detailed insights into mammal dung-dung beetle networks. In addition, the method could have applications for mammal monitoring in many ecosystems.

Nutrients or resin? - The relationship between resin and food foraging in stingless bees.

Stingless bees are important pollinators in tropical forests. Yet, we know little about their foraging behavior (e.g., their nutritional requirements or their floral sources visited for resource collection). Many stingless bees not only depend vitally on pollen and nectar for food but also on resin for nest building and/or defense. However, it is unclear whether the large effort devoted to collecting resin as a non-food resource by certain stingless bees affects their foraging behavior. Therefore, in this study, we analyzed differences in foraging patterns (i.e., foraging activity, proportion of collected resources, and specialization in plants visited) and resource nutritional composition (i.e., sucrose amount in nectar and amino acids in pollen) of seven different stingless bee species (eleven wild colonies) in north-western Ecuador with a particular focus on the role of resin collection. We found that species with a high resin intake tended to be more active than species with a low resin intake. The foragers per minute invested for pollen collection were similar across all species. Sucrose intake per minute differed between some species but was not affected by increased resin intake. Interestingly, high and low resin collectors partly differed in the plants visited for pollen collection. Pollen amino acid profiles largely, but not completely, overlapped between the two resin collection groups. Our findings show that the foraging patterns and plant choices of stingless bees may vary depending on their resin intake, highlighting the need for more research focusing on resin collection and use by stingless bees.

Las abejas sin aguijón son polinizadores importantes en los bosques tropicales. Sin embargo, sabemos poco acerca de su comportamiento de forrajeo (e.g., sus requisitos nutricionales o las fuentes florales visitadas para la recolección de recursos). Muchas abejas sin aguijón dependen vitalmente no solo de polen y de néctar como alimento, sino también de resinas para la construcción de su nido y/o defensa. Sin embargo, no está claro si el gran esfuerzo dedicado a la recolección de resina como recurso no alimentario de ciertas abejas sin aguijón afecta su comportamiento de forrajeo. Por lo tanto, en este estudio, analizamos las diferencias en los patrones de forrajeo (i.e., actividad de forrajeo, proporción de recursos recolectados y especialización en las plantas visitadas) y la composición nutricional de los recursos recolectados (i.e., cantidad de sacarosa en el néctar y de aminoácidos en el polen) de siete especies diferentes de abejas sin aguijón (once colonias silvestres) en el noroeste de Ecuador, con un enfoque particular en el rol de la recolección de resina. Encontramos que las especies con una recolección alta de resina tienden a ser más activas que las especies con una recolección baja de resina. La cantidad de forrajeadores por minuto dedicada a la recolección de polen fue similar en todas las especies. La ingesta de sacarosa por minuto difirió entre algunas especies, pero no se vio afectada por un aumento en la recolección de resina. Interesantemente, las abejas con una recolección alta y baja de resina difirieron parcialmente en las plantas que visitaron para la recolección de polen. Entre los dos grupos de recolección de resina también hubo diferencias con respecto al perfil de aminoácidos en el polen que recolectaron. El perfil de aminoácidos se sobrelapaba, pero no completamente, entre los dos grupos. Nuestros resultados muestran que los patrones de forrajeo y las elecciones de plantas de las abejas sin aguijón pueden variar según su consumo de resina, destacando la necesidad de hacer más investigaciones centradas en la recolección y el uso de resina por parte de las abejas sin aguijón.

Predicting plant-pollinator interactions: concepts, methods, and challenges.

Plant-pollinator interactions are ecologically and economically important, and, as a result, their prediction is a crucial theoretical and applied goal for ecologists. Although various analytical methods are available, we still have a limited ability to predict plant-pollinator interactions. The predictive ability of different plant-pollinator interaction models depends on the specific definitions used to conceptualize and quantify species attributes (e.g., morphological traits), sampling effects (e.g., detection probabilities), and data resolution and availability. Progress in the study of plant-pollinator interactions requires conceptual and methodological advances concerning the mechanisms and species attributes governing interactions as well as improved modeling approaches to predict interactions. Current methods to predict plant-pollinator interactions present ample opportunities for improvement and spark new horizons for basic and applied research.

Soundscapes and deep learning enable tracking biodiversity recovery in tropical forests.

Tropical forest recovery is fundamental to addressing the intertwined climate and biodiversity loss crises. While regenerating trees sequester carbon relatively quickly, the pace of biodiversity recovery remains contentious. Here, we use bioacoustics and metabarcoding to measure forest recovery post-agriculture in a global biodiversity hotspot in Ecuador. We show that the community composition, and not species richness, of vocalizing vertebrates identified by experts reflects the restoration gradient. Two automated measures - an acoustic index model and a bird community composition derived from an independently developed Convolutional Neural Network - correlated well with restoration (adj-R² = 0.62 and 0.69, respectively). Importantly, both measures reflected composition of non-vocalizing nocturnal insects identified via metabarcoding. We show that such automated monitoring tools, based on new technologies, can effectively monitor the success of forest recovery, using robust and reproducible data.

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.

Dear neighbor: Trees with extrafloral nectaries facilitate defense and growth of adjacent undefended trees.

Plant diversity can increase productivity. One mechanism behind this biodiversity effect is facilitation, which is when one species increases the performance of another species. Plants with extrafloral nectaries (EFNs) establish defense mutualisms with ants. However, whether EFN plants facilitate defense of neighboring non-EFN plants is unknown. Synthesizing data on ants, herbivores, leaf damage, and defense traits from a forest biodiversity experiment, we show that trees growing adjacent to EFN trees had higher ant biomass and species richness and lower caterpillar biomass than conspecific controls without EFN-bearing neighbors. Concurrently, the composition of defense traits in non-EFN trees changed. Thus, when non-EFN trees benefit from lower herbivore loads as a result of ants spilling over from EFN tree neighbors, this may allow relatively reduced resource allocation to defense in the former, potentially explaining the higher growth of those trees. Via this mutualist-mediated facilitation, promoting EFN trees in tropical reforestation could foster carbon capture and multiple other ecosystem functions.

Insect decline in forests depends on species' traits and may be mitigated by management.

Insects are declining, but the underlying drivers and differences in responses between species are still largely unclear. Despite the importance of forests, insect trends therein have received little attention. Using 10 years of standardized data (120,996 individuals; 1,805 species) from 140 sites in Germany, we show that declines occurred in most sites and species across trophic groups. In particular, declines (quantified as the correlation between year and the respective community response) were more consistent in sites with many non-native trees or a large amount of timber harvested before the onset of sampling. Correlations at the species level depended on species' life-history. Larger species, more abundant species, and species of higher trophic level declined most, while herbivores increased. This suggests potential shifts in food webs possibly affecting ecosystem functioning. A targeted management, including promoting more natural tree species composition and partially reduced harvesting, can contribute to mitigating declines.

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.

The supply of multiple ecosystem services requires biodiversity across spatial scales.

The impact of local biodiversity loss on ecosystem functioning is well established, but the role of larger-scale biodiversity dynamics in the delivery of ecosystem services remains poorly understood. Here we address this gap using a comprehensive dataset describing the supply of 16 cultural, regulating and provisioning ecosystem services in 150 European agricultural grassland plots, and detailed multi-scale data on land use and plant diversity. After controlling for land-use and abiotic factors, we show that both plot-level and surrounding plant diversity play an important role in the supply of cultural and aboveground regulating ecosystem services. In contrast, provisioning and belowground regulating ecosystem services are more strongly driven by field-level management and abiotic factors. Structural equation models revealed that surrounding plant diversity promotes ecosystem services both directly, probably by fostering the spill-over of ecosystem service providers from surrounding areas, and indirectly, by maintaining plot-level diversity. By influencing the ecosystem services that local stakeholders prioritized, biodiversity at different scales was also shown to positively influence a wide range of stakeholder groups. These results provide a comprehensive picture of which ecosystem services rely most strongly on biodiversity, and the respective scales of biodiversity that drive these services. This key information is required for the upscaling of biodiversity-ecosystem service relationships, and the informed management of biodiversity within agricultural landscapes.

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.

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.

Differences in prey availability across space and time lead to interaction rewiring and reshape a predator-prey metaweb.

Space and time promote variation in network structure by affecting the likelihood of potential interactions. However, little is known about the relative roles of ecological and biogeographical processes in determining how species interactions vary across space and time. Here we study the spatiotemporal variation in predator-prey interaction networks formed by anurans and arthropods and test for the effects of prey availability in determining interaction patterns, information that is often absent and limits the understanding of the determinants of network structure. We found that network dissimilarity between ecoregions and seasons was high and primarily driven by interaction rewiring.We also found that species turnover was positively related to geographical distance. Using a null model approach to disentangle the effect of prey availability on the spatial and temporal variation, we show that differences in prey availability were important in determining the variation in network structure between seasons and among areas. Our study reveals that fluctuations in prey abundance, alongside the limited dispersal abilities of anurans and their prey, may be responsible for the spatial patterns that emerged in our predator-prey metaweb. These findings contribute to our understanding of the assembly rules that maintain biotic processes in metacommunities and highlight the importance of prey availability to the structure of these systems.

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.

Animal-Mediated Ecosystem Process Rates in Forests and Grasslands are Affected by Climatic Conditions and Land-Use Intensity.

Decomposition, vegetation regeneration, and biological control are essential ecosystem functions, and animals are involved in the underlying processes, such as dung removal, seed removal, herbivory, and predation. Despite evidence for declines of animal diversity and abundance due to climate change and land-use intensification, we poorly understand how animal-mediated processes respond to these global change drivers. We experimentally measured rates of four ecosystem processes in 134 grassland and 149 forest plots in Germany and tested their response to climatic conditions and land-use intensity, that is, grazing, mowing, and fertilization in grasslands and the proportion of harvested wood, non-natural trees, and deadwood origin in forests. For both climate and land use, we distinguished between short-term effects during the survey period and medium-term effects during the preceding years. Forests had significantly higher process rates than grasslands. In grasslands, the climatic effects on the process rates were similar or stronger than land-use effects, except for predation; land-use intensity negatively affected several process rates. In forests, the land-use effects were more pronounced than the climatic effects on all processes except for predation. The proportion of non-natural trees had the greatest impact on the process rates in forests. The proportion of harvested wood had negative effects, whereas the proportion of anthropogenic deadwood had positive effects on some processes. The effects of climatic conditions and land-use intensity on process rates mirror climatic and habitat effects on animal abundance, activity, and resource quality. Our study demonstrates that land-use changes and interventions affecting climatic conditions will have substantial impacts on animal-mediated ecosystem processes.

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.

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.