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Forecasting insect responses to environmental variables at local and global spatial scales remains a crucial task in Ecology. However, predicting future responses requires long-term datasets, which are rarely available for insects, especially in the tropics. From 2002 to 2017, we recorded male ant incidence of 155 ant species at ten malaise traps on the 50-ha ForestGEO plot in Barro Colorado Island. In this Panamanian tropical rainforest, traps were deployed for two weeks during the wet and dry seasons. Short-term changes in the timing of male flying activity were pronounced, and compositionally distinct assemblages flew during the wet and dry seasons. Notably, the composition of these distinct flying assemblages oscillated in consistent 4-year cycles but did not change during the 16-year study period. Across time, a Seasonal Auto-Regressive Integrated Moving Average model explained 75% of long-term variability in male ant production (i.e., the summed incidence of male species across traps), which responded negatively to monthly maximum temperature, and positively to sea surface temperature, a surrogate for El Niño Southern Oscillation (ENSO) events. Establishing these relationships allowed us to forecast ant production until 2022 when year-long local climate variables were available. Consistent with the data, the forecast indicated no significant changes in long-term temporal trends of male ant production. However, simulations of different scenarios of climate variables found that strong ENSO events and maximum temperature impacted male ant production positively and negatively, respectively. Our results highlight the dependence of ant male production on both short- and long-term temperature changes, which is critical under current global warming., Competing Interests: Declarations. Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

Understanding the origins of novel, complex phenotypes is a major goal in evolutionary biology. Poison frogs of the family Dendrobatidae have evolved the novel ability to acquire alkaloids from their diet for chemical defense at least three times. However, taxon sampling for alkaloids has been biased towards colorful species, without similar attention paid to inconspicuous ones that are often assumed to be undefended. As a result, our understanding of how chemical defense evolved in this group is incomplete. Here, we provide new data showing that, in contrast to previous studies, species from each undefended poison frog clade have measurable yet low amounts of alkaloids. We confirm that undefended dendrobatids regularly consume mites and ants, which are known sources of alkaloids. Thus, our data suggest that diet is insufficient to explain the defended phenotype. Our data support the existence of a phenotypic intermediate between toxin consumption and sequestration - passive accumulation - that differs from sequestration in that it involves no derived forms of transport and storage mechanisms yet results in low levels of toxin accumulation. We discuss the concept of passive accumulation and its potential role in the origin of chemical defenses in poison frogs and other toxin-sequestering organisms. In light of ideas from pharmacokinetics, we incorporate new and old data from poison frogs into an evolutionary model that could help explain the origins of acquired chemical defenses in animals and provide insight into the molecular processes that govern the fate of ingested toxins., Competing Interests: RT, JC, DD, MB, KL, KG, JS, JS, SR, JS, BS, DC, RF No competing interests declared, (© 2024, Tarvin et al.)

Understanding the origins of novel, complex phenotypes is a major goal in evolutionary biology. Poison frogs of the family Dendrobatidae have evolved the novel ability to acquire alkaloids from their diet for chemical defense at least three times. However, taxon sampling for alkaloids has been biased towards colorful species, without similar attention paid to inconspicuous ones that are often assumed to be undefended. As a result, our understanding of how chemical defense evolved in this group is incomplete. Here we provide new data showing that, in contrast to previous studies, species from each undefended poison frog clade have measurable yet low amounts of alkaloids. We confirm that undefended dendrobatids regularly consume mites and ants, which are known sources of alkaloids. Thus, our data suggest that diet is insufficient to explain the defended phenotype. Our data support the existence of a phenotypic intermediate between toxin consumption and sequestration - passive accumulation - that differs from sequestration in that it involves no derived forms of transport and storage mechanisms yet results in low levels of toxin accumulation. We discuss the concept of passive accumulation and its potential role in the origin of chemical defenses in poison frogs and other toxin-sequestering organisms. In light of ideas from pharmacokinetics we incorporate new and old data from poison frogs into an evolutionary model that could help explain the origins of acquired chemical defenses in animals and provide insight into the molecular processes that govern the fate of ingested toxins.

Over 125 million years of ant-plant interactions have culminated in one of the most intriguing evolutionary outcomes in life history. The myrmecophyte Duroia hirsuta (Rubiaceae) is known for its mutualistic association with the ant Myrmelachista schumanni and several other species, mainly Azteca, in the north-western Amazon. While both ants provide indirect defences to plants, only M. schumanni nests in plant domatia and has the unique behaviour of clearing the surroundings of its host tree from heterospecific plants, potentially increasing resource availability to its host. Using a 12-year survey, we asked how the continuous presence of either only M. schumanni or only Azteca spp. benefits the growth and defence traits of host trees. We found that the continuous presence of M. schumanni improved relative growth rates and leaf shearing resistance of Duroia better than trees with Azteca. However, leaf herbivory, dry matter content, trichome density, and secondary metabolite production were the same in all trees. Survival depended directly on ant association (> 94% of trees died when ants were absent). This study extends our understanding of the long-term effects of strict ant-plant mutualism on host plant traits in the field and reinforces the use of D. hirsuta-M. schumanni as a model system suitable for eco-co-evolutionary research on plant-animal interactions., (© 2024. The Author(s).)

Earth harbours an extraordinary plant phenotypic diversity 1 that is at risk from ongoing global changes 2,3 . However, it remains unknown how increasing aridity and livestock grazing pressure-two major drivers of global change 4-6 -shape the trait covariation that underlies plant phenotypic diversity 1,7 . Here we assessed how covariation among 20 chemical and morphological traits responds to aridity and grazing pressure within global drylands. Our analysis involved 133,769 trait measurements spanning 1,347 observations of 301 perennial plant species surveyed across 326 plots from 6 continents. Crossing an aridity threshold of approximately 0.7 (close to the transition between semi-arid and arid zones) led to an unexpected 88% increase in trait diversity. This threshold appeared in the presence of grazers, and moved toward lower aridity levels with increasing grazing pressure. Moreover, 57% of observed trait diversity occurred only in the most arid and grazed drylands, highlighting the phenotypic uniqueness of these extreme environments. Our work indicates that drylands act as a global reservoir of plant phenotypic diversity and challenge the pervasive view that harsh environmental conditions reduce plant trait diversity 8-10 . They also highlight that many alternative strategies may enable plants to cope with increases in environmental stress induced by climate change and land-use intensification., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Perennial plants create productive and biodiverse hotspots, known as fertile islands, beneath their canopies. These hotspots largely determine the structure and functioning of drylands worldwide. Despite their ubiquity, the factors controlling fertile islands under conditions of contrasting grazing by livestock, the most prevalent land use in drylands, remain virtually unknown. Here we evaluated the relative importance of grazing pressure and herbivore type, climate and plant functional traits on 24 soil physical and chemical attributes that represent proxies of key ecosystem services related to decomposition, soil fertility, and soil and water conservation. To do this, we conducted a standardized global survey of 288 plots at 88 sites in 25 countries worldwide. We show that aridity and plant traits are the major factors associated with the magnitude of plant effects on fertile islands in grazed drylands worldwide. Grazing pressure had little influence on the capacity of plants to support fertile islands. Taller and wider shrubs and grasses supported stronger island effects. Stable and functional soils tended to be linked to species-rich sites with taller plants. Together, our findings dispel the notion that grazing pressure or herbivore type are linked to the formation or intensification of fertile islands in drylands. Rather, our study suggests that changes in aridity, and processes that alter island identity and therefore plant traits, will have marked effects on how perennial plants support and maintain the functioning of drylands in a more arid and grazed world., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

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., Competing Interests: The authors reported no conflict of interest., (© 2024 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

In tropical forests, herbivorous arthropods remove between 7% up to 48% of leaf area, which has forced plants to evolve defense strategies. These strategies influence the palatability of leaves. Palatability, which reflects a syndrome of leaf traits, in turn influences both the abundance and the mean body mass not only of particular arthropod taxa but also of the total communities. In this study, we tested two hypotheses: (H1) The abundance of two important chewer guilds ('leaf chewers' and 'rostrum chewers'), dominant components of arthropod communities, is positively related to the palatability of host trees. (H2) Lower palatability leads to an increased mean body mass of chewers (Jarman-Bell principle). Arthropods were collected by fogging the canopies of 90 tropical trees representing 31 species in three plots at 1000 m and three at 2000 m a.s.l. Palatability was assessed by measuring several 'leaf traits' of each host tree and by conducting a feeding trial with the generalist herbivore Gryllus assimilis (Orthoptera, Gryllidae). Leaf traits provided partial support for H1, as abundance of leaf chewers but not of rostrum chewers was positively affected by the experimentally estimated palatability. There was no support for H2 as neither leaf traits nor experimentally estimated palatability affected the mean body mass of leaf chewers. The mean body mass of rostrum chewers was positively related to palatability. Thus, leaf traits and experimentally estimated palatability influenced the abundance and mean body mass of chewing arthropods on the community level. However, the data were not consistent with the Jarman-Bell principle. Overall, our results suggest that the palatability of leaves is not among the dominant factors influencing abundance and mean body mass of the community of chewing arthropod herbivores. If other factors, such as the microclimate, predation or further (a-)biotic interactions are more important has to be analyzed in refined studies., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2023 Schön et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

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., (© 2023. Springer Nature Limited.)

This work updates the invertebrate type specimen catalog published by Donoso et al. (2009). The catalog is increased by 2281 type specimens (from 454 species or subspecies) to a total of 4180 type specimens (from 770 species or subspecies) hosted at the Pontificia Universidad Católica del Ecuador and Escuela Politécnica Nacional natural history collections. The new material adds 307 holotypes, 1910 paratypes, and 64 allotypes. It provides original information from four phyla (Arthropoda, Mollusca, Nemata, and Platyhelminthes), eight classes, 21 orders, 73 families, and 156 genera. This updated catalog includes a map showing the type localities in the country, a list of the 71 new type specimens (from 23 species or subspecies) from other countries hosted at both museums, corrections to the previous catalog published by Donoso et al. (2009), and label information from each new specimen., Competing Interests: No conflict of interest was declared., (Fernanda Salazar-Buenaño, Diego Guevara, Alvaro Barragán, Vladimir Carvajal, David A. Donoso.)

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., (© 2023 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and species-poor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. Considering interactions between grazing and local abiotic and biotic factors is key for understanding the fate of dryland ecosystems under climate change and increasing human pressure.

Invertebrates constitute the majority of animal species and are critical for ecosystem functioning and services. Nonetheless, global invertebrate biodiversity patterns and their congruences with vertebrates remain largely unknown. We resolve the first high-resolution (~20-km) global diversity map for a major invertebrate clade, ants, using biodiversity informatics, range modeling, and machine learning to synthesize existing knowledge and predict the distribution of undiscovered diversity. We find that ants and different vertebrate groups have distinct features in their patterns of richness and rarity, underscoring the need to consider a diversity of taxa in conservation. However, despite their phylogenetic and physiological divergence, ant distributions are not highly anomalous relative to variation among vertebrate clades. Furthermore, our models predict that rarity centers largely overlap (78%), suggesting that general forces shape endemism patterns across taxa. This raises confidence that conservation of areas important for small-ranged vertebrates will benefit invertebrates while providing a "treasure map" to guide future discovery.

The soil fauna of the tropics remains one of the least known components of the biosphere. Long-term monitoring of this fauna is hampered by the lack of taxonomic expertise and funding. These obstacles may potentially be lifted with DNA metabarcoding. To validate this approach, we studied the ants, springtails and termites of 100 paired soil samples from Barro Colorado Island, Panama. The fauna was extracted with Berlese-Tullgren funnels and then either sorted with traditional taxonomy and known, individual DNA barcodes ("traditional samples") or processed with metabarcoding ("metabarcoding samples"). We detected 49 ant, 37 springtail and 34 termite species with 3.46 million reads of the COI gene, at a mean sequence length of 233 bp. Traditional identification yielded 80, 111 and 15 species of ants, springtails and termites, respectively; 98%, 37% and 100% of these species had a Barcode Index Number (BIN) allowing for direct comparison with metabarcoding. Ants were best surveyed through traditional methods, termites were better detected by metabarcoding, and springtails were equally well detected by both techniques. Species richness was underestimated, and faunal composition was different in metabarcoding samples, mostly because 37% of ant species were not detected. The prevalence of species in metabarcoding samples increased with their abundance in traditional samples, and seasonal shifts in species prevalence and faunal composition were similar between traditional and metabarcoding samples. Probable false positive and negative species records were reasonably low (13-18% of common species). We conclude that metabarcoding of samples extracted with Berlese-Tullgren funnels appear suitable for the long-term monitoring of termites and springtails in tropical rainforests. For ants, metabarcoding schemes should be complemented by additional samples of alates from Malaise or light traps., (© 2022. The Author(s).)

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., (© 2022 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Tropical forests sustain many ant species whose mating events often involve conspicuous flying swarms of winged gynes and males. The success of these reproductive flights depends on environmental variables and determines the maintenance of local ant diversity. However, we lack a strong understanding of the role of environmental variables in shaping the phenology of these flights. Using a combination of community-level analyses and a time-series model on male abundance, we studied male ant phenology in a seasonally wet lowland rainforest in the Panama Canal. The male flights of 161 ant species, sampled with 10 Malaise traps during 58 consecutive weeks (from August 2014 to September 2015), varied widely in number (mean = 9.8 weeks, median = 4, range = 1 to 58). Those species abundant enough for analysis (n = 97) flew mainly towards the end of the dry season and at the start of the rainy season. While litterfall, rain, temperature, and air humidity explained community composition, the time-series model estimators elucidated more complex patterns of reproductive investment across the entire year. For example, male abundance increased in weeks when maximum daily temperature increased and in wet weeks during the dry season. On the contrary, male abundance decreased in periods when rain receded (e.g., at the start of the dry season), in periods when rain fell daily (e.g., right after the beginning of the wet season), or when there was an increase in the short-term rate of litterfall (e.g., at the end of the dry season). Together, these results suggest that the BCI ant community is adapted to the dry/wet transition as the best timing of reproductive investment. We hypothesize that current climate change scenarios for tropical regions with higher average temperature, but lower rainfall, may generate phenological mismatches between reproductive flights and the adequate conditions needed for a successful start of the colony., Competing Interests: The authors have declared that no competing interests exist.

Biodiversity and ecosystem functions are highly threatened by global change. It has been proposed that geodiversity can be used as an easy-to-measure surrogate of biodiversity to guide conservation management. However, so far, there is mixed evidence to what extent geodiversity can predict biodiversity and ecosystem functions at the regional scale relevant for conservation planning. Here, we analyse how geodiversity computed as a compound index is suited to predict the diversity of four taxa and associated ecosystem functions in a tropical mountain hotspot of biodiversity and compare the results with the predictive power of environmental conditions and resources (climate, habitat, soil). We show that combinations of these environmental variables better explain species diversity and ecosystem functions than a geodiversity index and identified climate variables as more important predictors than habitat and soil variables, although the best predictors differ between taxa and functions. We conclude that a compound geodiversity index cannot be used as a single surrogate predictor for species diversity and ecosystem functions in tropical mountain rain forest ecosystems and is thus little suited to facilitate conservation management at the regional scale. Instead, both the selection and the combination of environmental variables are essential to guide conservation efforts to safeguard biodiversity and ecosystem functions., (© 2021. The Author(s).)

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

Water contamination is a major environmental problem in many cities of the world. Most water contamination results from industry and human activities that generate toxic substances (e.g., metals). Rheophilic and aquatic mosses are found in lotic ecosystems, and their morphological and physiological traits are responsive to ecological and pollution gradients. Here we hypothesized that the native rheophilic moss Platyhypnidium aquaticum (A. Jaeger) M. Fleisch exposed to polluted waters can bioaccumulate greater amounts of metals, and a metalloid, than P. aquaticum exposed to pollution-free water. To this aim, we tested the bioindicator capacity of the aquatic P. aquaticum for 15 metals (Cd, Pb, Zn, Fe, K, Ca, Na, Mn, V, Co, Ba, Cr, Al, Sr, and Mg) and one metalloid (As), in twelve river samples coming from three urban and one control zone along the Zamora river in the city of Loja. When compared to the control, our results showed that P. aquaticum in the Southern, Central, and Northern zones of the city bioaccumulated higher concentrations of Ba, Cd, Co, Fe, Mg, Mn, Na, Sr, Zn, and the metalloid As. On the other hand, concentrations of Al, Ca, Cr, Pb, and V in P. aquaticum tended to be lower in the control zone, but these differences were not significant. We suggest that the presence of these contaminants may be related to water pollution (e.g., residual discharges and a lack of treatment systems) along urban zones of the river. We report for the first time the utility of P. aquaticum as a model species for development of long-term biomonitoring programs of water contamination in South America. Passive biomonitoring with P. aquaticum can be a simple and low-cost method to obtain reliable data of the current state of water contamination with metals and metalloids in tropical regions.

One of the largest species in its genus, Odontomachus davidsoni Hoenle, Lattke & Donoso, sp. nov. is described from workers and queens collected at lowland forests in the Chocó-Darién bioregion in coastal Ecuador. The workers are characterized by their uniform red coloration, their large size (16-18 mm body length), and their frontal head striation that reaches the occipital margin. DNA barcodes (COI) and high resolution 2D images of the type material are provided, as well as an updated key for the Neotropical species of Odontomachus . In addition, a three-dimensional digital model of the worker holotype and a paratype queen scanned with DISC3D based on photogrammetry is presented, for the first time in a species description. Findings of large and conspicuous new species are uncommon around the world and suggest that these Ecuadorian rainforests may conceal many more natural treasures that deserve conservation., (Philipp Hoenle, John Edwin Lattke, David Donoso, Christoph von Beeren, Michael Heethoff, Sebastian Schmelzle, Adriana Argoti, Luis Camacho, Bernhard Ströbel, Nico Blüthgen.)

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., (© 2019 John Wiley & Sons Ltd.)

Poison frogs acquire chemical defenses from the environment for protection against potential predators. These defensive chemicals are lipophilic alkaloids that are sequestered by poison frogs from dietary arthropods and stored in skin glands. Despite decades of research focusing on identifying poison frog alkaloids, we know relatively little about how environmental variation and subsequent arthropod availability impacts alkaloid loads in poison frogs. We investigated how seasonal environmental variation influences poison frog chemical profiles through changes in the diet of the Climbing Mantella (Mantella laevigata). We collected M. laevigata females on the Nosy Mangabe island reserve in Madagascar during the wet and dry seasons and tested the hypothesis that seasonal differences in rainfall is associated with changes in diet composition and skin alkaloid profiles of M. laevigata. The arthropod diet of each frog was characterized into five groups (i.e. ants, termites, mites, insect larvae, or 'other') using visual identification and cytochrome oxidase 1 DNA barcoding. We found that frog diet differed between the wet and dry seasons, where frogs had a more diverse diet in the wet season and consumed a higher percentage of ants in the dry season. To determine if seasonality was associated with variation in frog defensive chemical composition, we used gas chromatography / mass spectrometry to quantify alkaloids from individual skin samples. Although the assortment of identified alkaloids was similar across seasons, we detected significant differences in the abundance of certain alkaloids, which we hypothesize reflects seasonal variation in the diet of M. laevigata. We suggest that these variations could originate from seasonal changes in either arthropod leaf litter composition or changes in frog behavioral patterns. Although additional studies are needed to understand the consequences of long-term environmental shifts, this work suggests that alkaloid profiles are relatively robust against short-term environmental perturbations., Competing Interests: The authors have declared that no competing interests exist.

Leaf-cutting ants are often considered agricultural pests, but they can also benefit local people and serve important roles in ecosystems. Throughout their distribution, winged reproductive queens of leaf-cutting ants in the genus Atta Fabricius, 1804 are consumed as a protein-rich food source and sometimes used for medical purposes. Little is known, however, about the species identity of collected ants and the accuracy of identification when ants are sold, ambiguities that may impact the conservation status of Atta species as well as the nutritional value that they provide to consumers. Here, 21 samples of fried ants bought in San Gil, Colombia, were identified to species level using Cytochrome Oxidase I (COI) barcoding sequences. DNA was extracted from these fried samples using standard Chelex extraction methods, followed by phylogenetic analyses with an additional 52 new sequences from wild ant colonies collected in Panama and 251 publicly available sequences. Most analysed samples corresponded to Atta laevigata (Smith, 1858), even though one sample was identified as Atta colombica Guérin-Méneville, 1844 and another one formed a distinct branch on its own, more closely related to Atta texana (Buckley, 1860) and Atta mexicana (Smith, 1858). Analyses further confirm paraphyly within Atta sexdens (Linnaeus, 1758) and A. laevigata clades. Further research is needed to assess the nutritional value of the different species.

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., (© 2018 John Wiley & Sons Ltd.)

Coevolutionary models suggest that herbivores drive diversification and community composition in plants. For herbivores, many questions remain regarding how plant defenses shape host choice and community structure. We addressed these questions using the tree genus Inga and its lepidopteran herbivores in the Amazon. We constructed phylogenies for both plants and insects and quantified host associations and plant defenses. We found that similarity in herbivore assemblages between Inga species was correlated with similarity in defenses. There was no correlation with phylogeny, a result consistent with our observations that the expression of defenses in Inga is independent of phylogeny. Furthermore, host defensive traits explained 40% of herbivore community similarity. Analyses at finer taxonomic scales showed that different lepidopteran clades select hosts based on different defenses, suggesting taxon-specific histories of herbivore-host plant interactions. Finally, we compared the phylogeny and defenses of Inga to phylogenies for the major lepidopteran clades. We found that closely related herbivores fed on Inga with similar defenses rather than on closely related plants. Together, these results suggest that plant defenses might be more evolutionarily labile than the herbivore traits related to host association. Hence, there is an apparent asymmetry in the evolutionary interactions between Inga and its herbivores. Although plants may evolve under selection by herbivores, we hypothesize that herbivores may not show coevolutionary adaptations, but instead "chase" hosts based on the herbivore's own traits at the time that they encounter a new host, a pattern more consistent with resource tracking than with the arms race model of coevolution., Competing Interests: The authors declare no conflict of interest.

Biotic interactions underlie ecosystem structure and function, but predicting interaction outcomes is difficult. We tested the hypothesis that biotic interaction strength increases toward the equator, using a global experiment with model caterpillars to measure predation risk. Across an 11,660-kilometer latitudinal gradient spanning six continents, we found increasing predation toward the equator, with a parallel pattern of increasing predation toward lower elevations. Patterns across both latitude and elevation were driven by arthropod predators, with no systematic trend in attack rates by birds or mammals. These matching gradients at global and regional scales suggest consistent drivers of biotic interaction strength, a finding that needs to be integrated into general theories of herbivory, community organization, and life-history evolution., (Copyright © 2017, American Association for the Advancement of Science.)

What forces structure ecological assemblages? A key limitation to general insights about assemblage structure is the availability of data that are collected at a small spatial grain (local assemblages) and a large spatial extent (global coverage). Here, we present published and unpublished data from 51 ,388 ant abundance and occurrence records of more than 2,693 species and 7,953 morphospecies from local assemblages collected at 4,212 locations around the world. Ants were selected because they are diverse and abundant globally, comprise a large fraction of animal biomass in most terrestrial communities, and are key contributors to a range of ecosystem functions. Data were collected between 1949 and 2014, and include, for each geo-referenced sampling site, both the identity of the ants collected and details of sampling design, habitat type, and degree of disturbance. The aim of compiling this data set was to provide comprehensive species abundance data in order to test relationships between assemblage structure and environmental and biogeographic factors. Data were collected using a variety of standardized methods, such as pitfall and Winkler traps, and will be valuable for studies investigating large-scale forces structuring local assemblages. Understanding such relationships is particularly critical under current rates of global change. We encourage authors holding additional data on systematically collected ant assemblages, especially those in dry and cold, and remote areas, to contact us and contribute their data to this growing data set., (© 2016 by the Ecological Society of America.)

How ecological context shapes mutualistic relationships remains poorly understood. We combined long-term tree census data with ant censuses in a permanent 25-ha Amazonian forest dynamics plot to evaluate the effect of the mutualistic ant Myrmelachista schumanni (Formicinae) on the growth and survival of the common Amazonian tree Duroia hirsuta (Rubiaceae), considering its interactions with tree growth, population structure, and habitat. We found that the mutualist ant more than doubled tree relative growth rates and increased odds of survival. However, host tree size and density of conspecific neighbors modified the effect of the ant. Smaller trees hosting the mutualist ant consistently grew faster when surrounded by higher densities of conspecifics, suggesting that the benefit to the tree outweighs any negative effects of high conspecific densities. Moreover, our findings suggest that the benefit afforded by the ant diminishes with plant age and also depends on the density of conspecific neighbors. We provide the first long-term large-scale evidence of how mutualism affects the population biology of an Amazonian tree species.

Recently a new species of bombyliid fly, Marleyimyia xylocopae, was described by Marshall & Evenhuis (2015) based on two photographs taken during fieldwork in the Republic of South Africa. This species has no preserved holotype. The paper generated some buzz, especially among dipterists, because in most cases photographs taken in the field provide insufficient information for properly diagnosing and documenting species of Diptera.

Many studies have focused on the impacts of climate change on biological assemblages, yet little is known about how climate interacts with other major anthropogenic influences on biodiversity, such as habitat disturbance. Using a unique global database of 1128 local ant assemblages, we examined whether climate mediates the effects of habitat disturbance on assemblage structure at a global scale. Species richness and evenness were associated positively with temperature, and negatively with disturbance. However, the interaction among temperature, precipitation and disturbance shaped species richness and evenness. The effect was manifested through a failure of species richness to increase substantially with temperature in transformed habitats at low precipitation. At low precipitation levels, evenness increased with temperature in undisturbed sites, peaked at medium temperatures in disturbed sites and remained low in transformed sites. In warmer climates with lower rainfall, the effects of increasing disturbance on species richness and evenness were akin to decreases in temperature of up to 9°C. Anthropogenic disturbance and ongoing climate change may interact in complicated ways to shape the structure of assemblages, with hot, arid environments likely to be at greatest risk., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

Nutrient pulses can profoundly impact ecosystem processes and urine is a frequently deposited source of N and K, and Na. Na is unimportant to plants, but its addition can increase decomposition and change invertebrate community structure in Na-poor tropical forests. Here we used synthetic urine to separate the effects of Na from urine's other nutrients and contrasted their roles in promoting decomposition and detritivore recruitment in both a Na-poor inland Ecuadorian and Na-rich coastal Panamanian tropical forest. After 2 days, invertebrate communities did not vary among +Na, H2O, Urine+Na, and Urine-Na treatments. But after 2 weeks, Ecuador wood, but not cellulose, decomposition was twofold higher on Urine+Na and +Na plots compared to H2O and Urine-Na plots accompanied by >20-fold increases in termite abundance on these plots. Panama, in contrast, showed no effect of Na on decomposition. In both forests, plots fertilized with urine had nearly twofold decrease in detritivores after 2 weeks that was likely a shock effect from ammonification. Moreover, the non-Na nutrients in urine did not enhance decomposition at this time scale. On control plots, Panama had higher decomposition rates for both cellulose and wood than Ecuador, but the addition of Na in Ecuador alleviated these differences. These results support the hypothesis that in Na-poor tropical forests, urine can enhance wood decomposition and generate an important source of heterogeneity in the abundance and activity of brown food webs.

Added Na was used to determine whether litter decomposition and associated fungal biomass and termites are limited by Na availability in a lowland tropical rainforest at Yasuni, Ecuador. This is a partial test of the "sodium ecosystem respiration" (SER) hypothesis that posits Na is critical for consumers but not plants, that Na shortfall is more likely on highly weathered soils inland from oceanic aerosols, and that this shortfall results in decreased decomposer activity. We fertilized 4 x 4 m plots twice a month for a year with quantities of Na comparable to those falling on a coastal tropical rainforest. Decomposition rates of four substrates were consistently higher on +NaCl plots by up to 70% for cellulose, and 78%, 68%, and 29% for three woods of increasing percentage lignin. The density of termite workers averaged 17-fold higher on +NaCl plots; fungal biomass failed to differ. After controlling for temperature and precipitation, which co-limit gross primay productivity (GPP) and ecosystem respiration (ER), these results suggest that Na shortfall is an agent enhancing the storage of coarse woody debris in inland tropical forests.

Increased tree species diversity in the tropics is associated with even greater herbivore diversity, but few tests of tree effects on litter arthropod diversity exist. We studied whether tree species influence patchiness in diversity and abundance of three common soil arthropod taxa (ants, gamasid mites, and oribatid mites) in a Panama forest. The tree specialization hypothesis proposes that tree-driven habitat heterogeneity maintains litter arthropod diversity. We tested whether tree species differed in resource quality and quantity of their leaf litter and whether more heterogeneous litter supports more arthropod species. Alternatively, the abundance-extinction hypothesis states that arthropod diversity increases with arthropod abundance, which in turn tracks resource quantity (e.g., litter depth). We found little support for the hypothesis that tropical trees are templates for litter arthropod diversity. Ten tree species differed in litter depth, chemistry, and structural variability. However, the extent of specialization of invertebrates on particular tree taxa was low and the more heterogeneous litter between trees failed to support higher arthropod diversity. Furthermore, arthropod diversity did not track abundance or litter depth. The lack of association between tree species and litter arthropods suggests that factors other than tree species diversity may better explain the high arthropod diversity in tropical forests.