Competition and habitat availability interact to structure arboreal ant communities across scales of ecological organization.

Understanding how resource limitation and biotic interactions interact across spatial scales is fundamental to explaining the structure of ecological communities. However, empirical studies addressing this issue are often hindered by logistical constraints, especially at local scales. Here, we use a highly tractable arboreal ant study system to explore the interactive effects of resource availability and competition on community structure across three local scales: an individual tree, the nest network created by each colony and the individual ant nest. On individual trees, the ant assemblages are primarily shaped by availability of dead wood, a critical nesting resource. The nest networks within a tree are constrained by the availability of nesting resources but also influenced by the co-occurring species. Within individual nests, the distribution of adult ants is only affected by distance to interspecific competitors. These findings demonstrate that resource limitation exerts the strongest effects on diversity at higher levels of local ecological organization, transitioning to a stronger effect of species interactions at finer scales. Collectively, these results highlight that the process exerting the strongest influence on community structure is highly dependent on the scale at which we examine the community, with shifts occurring even across fine-grained local scales.

Identifying the Role of Elevation, Geography, and Species Identity in Structuring Turtle Ant (Cephalotes Latreille, 1802) Bacterial Communities.

Bacterial communities in animals are often necessary for hosts to survive, particularly for hosts with nutrient-limited diets. The composition, abundance, and richness of these bacterial communities may be shaped by host identity and external ecological factors. The turtle ants (genus Cephalotes) are predominantly herbivorous and known to rely on bacterial communities to enrich their diet. Cephalotes have a broad Neotropical distribution, with high diversity in the South American Cerrado, a geologically and biologically diverse savanna. Using 16S rRNA amplicon sequencing, we examined the bacterial communities of forty-one Cephalotes samples of sixteen different species collected from multiple locations across two sites in the Cerrado (MG, Brazil) and compared the bacterial communities according to elevation, locality, species, and species group, defined by host phylogeny. Beta diversity of bacterial communities differed with respect to all categories but particularly strongly when compared by geographic location, species, and species group. Differences seen in species and species groups can be partially explained by the high abundance of Mesorhizobium in Cephalotes pusillus and Cephalotes depressus species groups, when compared to other clades via the Analysis of Composition of Microbiome (ANCOM). Though the Cephalotes bacterial community is highly conserved, results from this study indicate that multiple external factors can affect and change bacterial community composition and abundance.

Trait evolution is reversible, repeatable, and decoupled in the soldier caste of turtle ants.

The scope of adaptive phenotypic change within a lineage is shaped by how functional traits evolve. Castes are defining functional traits of adaptive phenotypic change in complex insect societies, and caste evolution is expected to be phylogenetically conserved and developmentally constrained at broad phylogenetic scales. Yet how castes evolve at the species level has remained largely unaddressed. Turtle ant soldiers (genus Cephalotes), an iconic example of caste specialization, defend nest entrances by using their elaborately armored heads as living barricades. Across species, soldier morphotype determines entrance specialization and defensive strategy, while head size sets the specific size of defended entrances. Our species-level comparative analyses of morphotype and head size evolution reveal that these key ecomorphological traits are extensively reversible, repeatable, and decoupled within soldiers and between soldier and queen castes. Repeated evolutionary gains and losses of the four morphotypes were reconstructed consistently across multiple analyses. In addition, morphotype did not predict mean head size across the three most common morphotypes, and head size distributions overlapped broadly across all morphotypes. Concordantly, multiple model-fitting approaches suggested that soldier head size evolution is best explained by a process of divergent pulses of change. Finally, while soldier and queen head size were broadly coupled across species, the level of head size disparity between castes was decoupled from both queen head size and soldier morphotype. These findings demonstrate that caste evolution can be highly dynamic at the species level, reshaping our understanding of adaptive morphological change in complex social lineages.

Distributing Defenses: How Resource Defendability Shapes the Optimal Response to Risk.

AbstractMany organisms divide limited defenses among heterogeneous assets. Plants allocate defensive chemicals among tissues differing in value, cost of defense, and risk of herbivory. Some ant colonies allocate specialized defenders among multiple nests differing in volume, entrance size, and risk of attack. We develop a general mathematical model to determine the optimal strategy for dividing defenses among assets depending on their value, defendability, and risk of attack. We build on plant defense theory by focusing on defendability, which we define as the functional relationship between defensive investment and successful defense. We show that if hard-to-defend assets cost more to defend, as assumed in resource defense theory, the optimal strategy allocates more defenses to those assets, regardless of risk. Inspired by cavity-nesting ants, we also consider the possibility that hard-to-defend assets have a lower chance to be successfully defended, even when defensive investment is high. Under this assumption, the optimal response to elevated risk is to reduce defensive allocation to hard-to-defend assets, a conservative strategy previously observed in turtle ants (Cephalotes). This new perspective on defendability suggests that in systems where assets differ in the chance of successful defense, defensive strategies may evolve to be sensitive to risk in surprising ways.

Warm and arid regions of the world are hotspots of superorganism complexity.

Biologists have long been fascinated by the processes that give rise to phenotypic complexity of organisms, yet whether there exist geographical hotspots of phenotypic complexity remains poorly explored. Phenotypic complexity can be readily observed in ant colonies, which are superorganisms with morphologically differentiated queen and worker castes analogous to the germline and soma of multicellular organisms. Several ant species have evolved 'worker polymorphism', where workers in a single colony show quantifiable differences in size and head-to-body scaling. Here, we use 256 754 occurrence points from 8990 ant species to investigate the geography of worker polymorphism. We show that arid regions of the world are the hotspots of superorganism complexity. Tropical savannahs and deserts, which are typically species-poor relative to tropical or even temperate forests, harbour the highest densities of polymorphic ants. We discuss the possible adaptive advantages that worker polymorphism provides in arid environments. Our work may provide a window into the environmental conditions that promote the emergence of highly complex phenotypes.

Informing conservation decisions to target private lands of highest ecological value and risk of loss.

Natural habitats on private lands are potentially important components of national biodiversity conservation strategies, yet they are being rapidly lost to development. Conservation easements and other means of protecting these habitats have expanded in use and will be most effective if they target private lands of highest biodiversity value and risk of loss. We developed a Biodiversity Conservation Priority Index (BCPI) based on ecological value and risk of habitat loss for remaining areas of natural vegetation cover (NVC) in the northwestern United States and addressed two questions: (1) Which remaining NVC on private lands is the highest priority for biodiversity conservation based on ecological value and risk of development? And (2) are conservation easements in NVC placed preferentially in locations of high biodiversity conservation priority? Drawing on the concept of ecological integrity, we integrated five metrics of ecological structure, function, and composition to quantify ecological value of NVC. These included net primary productivity, species richness, ecosystem type representation, imperiled species range rarity, and connectivity among "Greater Wildland Ecosystems." Risk of habitat loss was derived from analysis of biophysical and sociodemographic predictors of NVC loss. Ecological value and risk of loss were combined into the BCPI. We then analyzed spatial patterns of BCPI to identify the NVC highest in biodiversity conservation priority and examined the relationship between BCPI and conservation easement status. We found that BCPI varied spatially across the study area and was highest in western and southern portions of the study area. High BCPI was associated with suburban and rural development, roads, urban proximity, valley bottom landforms, and low intensity of current development. Existing conservation easements were distributed more towards lower BCPI values than unprotected NVC at both the study area and region scales. The BCPI can be used to better inform land use decision making at local, regional, and potentially national scales in order to better achieve biodiversity goals.

From foraging trails to transport networks: how the quality-distance trade-off shapes network structure.

Biological systems are typically dependent on transportation networks for the efficient distribution of resources and information. Revealing the decentralized mechanisms underlying the generative process of these networks is key in our global understanding of their functions and is of interest to design, manage and improve human transport systems. Ants are a particularly interesting taxon to address these issues because some species build multi-sink multi-source transport networks analogous to human ones. Here, by combining empirical field data and modelling at several scales of description, we show that pre-existing mechanisms of recruitment with positive feedback involved in foraging can account for the structure of complex ant transport networks. Specifically, we find that emergent group-level properties of these empirical networks, such as robustness, efficiency and cost, can arise from models built on simple individual-level behaviour addressing a quality-distance trade-off by the means of pheromone trails. Our work represents a first step in developing a theory for the generation of effective multi-source multi-sink transport networks based on combining exploration and positive reinforcement of best sources.

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

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

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

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

Patterns of transaminase elevation in rhabdomyolysis versus acetaminophen toxicity.

BACKGROUND: Transaminase elevations can occur from liver injury or in the setting of rhabdomyolysis. The goal of this study is to evaluate indices that could differentiate acetaminophen toxicity from muscle injury in the setting of transaminase elevations. METHODS: A retrospective chart review of consecutive cases reported to our regional poison center. Patients with transaminase (AST and ALT) elevation were grouped as those with acetaminophen exposure (AT) and those with elevated creatine phosphokinase (CPK) without evidence of acetaminophen exposure (RHB). RESULTS: Of the 345 patients included in the study, elevated AST/ALT levels were attributed to rhabdomyolysis in 168 patients and attributed to acetaminophen toxicity in 177 patients. The median AST: ALT values also differed between groups, with patients in the RHB group had higher median ratios (p < 0.001). Using an AST: ALT value of 2.02 as a diagnostic cutoff produced a specificity of 0.52 (95% CI: 0.37, 0.64) and sensitivity of 0.84 (95% CI: 0.73, 0.94) for acetaminophen detection in the test dataset (N = 104). CONCLUSIONS: Elevated transaminases due to liver injury from acetaminophen ingestion had a different pattern than elevated transaminases due to rhabdomyolysis. Lower AST:ALT ratios were found in acetaminophen cases, however, the specificity using a ratio threshold of ≤1 would be 83%.

Risk Assessment for the Establishment of Vespa mandarinia (Hymenoptera: Vespidae) in the Pacific Northwest, United States.

The recent introduction of the Asian giant hornet, Vespa mandarinia Smith, in the United States in late 2019 has raised concerns about its establishment in the Pacific Northwest and its potential deleterious effects on honey bees, Apis spp., and their pollination services in the region. Therefore, we conducted a risk assessment of the establishment of V. mandarinia in Washington, Oregon, Montana, and Idaho on a county-by-county basis. Our highly conservative tier-1 qualitative and semiquantitative risk assessment relied on the biological requirements and ecological relationships of V. mandarinia in the environments of the Pacific Northwest. Our risk characterization was based on climate and habitat suitability estimates for V. mandarinia queens to overwinter and colonize nests, density and distribution of apiaries, and locations of major human-mediated introduction pathways that may increase establishment of the hornet in the counties. Our results suggest that 32 counties in the region could be at low risk, 120 at medium risk, and 23 at high risk of establishment. Many of the western counties in the region were estimated to be at the highest risk of establishment mainly because of their suitable climate for queens to overwinter, dense forest biomass for nest colonization, and proximity to major port and freight hubs in the region. By design, our tier-1 risk assessment most likely overestimates the risk of establishment, but considering its negative effects, these counties should be prioritized in ongoing monitoring and eradication efforts of V. mandarinia.

Detection of prey odours underpins dietary specialization in a Neotropical top-predator: How army ants find their ant prey.

Deciphering the mechanisms that underpin dietary specialization and niche partitioning is crucial to understanding the maintenance of biodiversity. New world army ants live in species-rich assemblages throughout the Neotropics and are voracious predators of other arthropods. They are therefore an important and potentially informative group for addressing how diverse predator assemblages partition available prey resources. New World army ants are largely specialist predators of other ants, with each species specializing on different ant genera. However, the mechanisms of prey choice are unknown. In this study, we addressed whether the army ant Eciton hamatum: (a) can detect potential prey odours, (b) can distinguish between odours of prey and non-prey and (c) can differentiate between different types of odours associated with its prey. Using field experiments, we tested the response of army ants to the following four odour treatments: alarm odours, dead ants, live ants and nest material. Each treatment had a unique combination of odour sources and included some movement in two of the treatments (alarm and live ants). Odour treatments were tested for both prey and non-prey ants. These data were used to determine the degree to which E. hamatum are using specific prey stimuli to detect potential prey and direct their foraging. Army ants responded strongly to odours derived from prey ants, which triggered both increased localized recruitment and slowed advancement of the raid as they targeted the odour source. Odours from non-prey ants were largely ignored. Additionally, the army ants had the strongest response to the nest material of their preferred prey, with progressively weaker responses across the live ant, dead ant and alarm odours treatments respectively. This study reveals that the detection of prey odours, and especially the most persistent odours related to the prey's nest, provides a mechanism for dietary specialization in army ants. If ubiquitous across the Neotropical army ants, then this olfaction-based ecological specialization may facilitate patterns of resource partitioning and coexistence in these diverse predator communities.

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

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

Correlates and Consequences of Worker Polymorphism in Ants.

Body size is a key life-history trait influencing all aspects of an organism's biology. Ants provide an interesting model for examining body-size variation because of the high degree of worker polymorphism seen in many taxa. We review worker-size variation in ants from the perspective of factors internal and external to the colony that may influence body-size distributions. We also discuss proximate and ultimate causes of size variation and how variation in worker size can promote worker efficiency and colony fitness. Our review focuses on two questions: What is our current understanding of factors influencing worker-size variation? And how does variation in body size benefit the colony? We conclude with recommendations for future work aimed at addressing current limitations and ask, How can we better understand the contribution of worker body-size variation to colony success? And, what research is needed to address gaps in our knowledge?

Army ants dynamically adjust living bridges in response to a cost-benefit trade-off.

The ability of individual animals to create functional structures by joining together is rare and confined to the social insects. Army ants (Eciton) form collective assemblages out of their own bodies to perform a variety of functions that benefit the entire colony. Here we examine ?bridges" of linked individuals that are constructed to span gaps in the colony's foraging trail. How these living structures adjust themselves to varied and changing conditions remains poorly understood. Our field experiments show that the ants continuously modify their bridges, such that these structures lengthen, widen, and change position in response to traffic levels and environmental geometry. Ants initiate bridges where their path deviates from their incoming direction and move the bridges over time to create shortcuts over large gaps. The final position of the structure depended on the intensity of the traffic and the extent of path deviation and was influenced by a cost-benefit trade-off at the colony level, where the benefit of increased foraging trail efficiency was balanced by the cost of removing workers from the foraging pool to form the structure. To examine this trade-off, we quantified the geometric relationship between costs and benefits revealed by our experiments. We then constructed a model to determine the bridge location that maximized foraging rate, which qualitatively matched the observed movement of bridges. Our results highlight how animal self-assemblages can be dynamically modified in response to a group-level cost-benefit trade-off, without any individual unit's having information on global benefits or costs.

Accuracy of palpation-directed intra-articular glenohumeral injection confirmed by magnetic resonance arthrography.

PURPOSE: The aim of this study was to determine the accuracy of anatomic palpation-directed injections in the office setting. METHODS: Two hundred twenty-six shoulders in 208 patients were studied using a 0.2-Tesla extremity scanner after the injection of gadolinium-diethylene triamine pentaacetic acid-saline. All patients were injected in a sterile fashion by a single board-certified shoulder surgeon using an anterior approach by palpating the rotator interval anterior to the acromioclavicular joint and angling the needle 45° lateral and 45° caudad. All injections, successful or otherwise, were single injections. Magnetic resonance (MR) arthrograms were retrospectively read by 2 musculoskeletal fellowship-trained, board certified radiologists to determine whether the injection was in the glenohumeral joint. RESULTS: Two hundred one of the 226 injections were successful (88.9%). Of the 25 unsuccessful injections, the contrast material extravasated out of the capsule in 5 cases and into the subscapularis tendon in 10 cases. The contrast material was injected into the subacromial space in 9 cases, into the rotator interval fat in 9 cases, and into extracapsular tissue in 6 cases. There was insufficient volume of contrast material in 10 cases. The accuracy rate was 88.9%. There were no complications. CONCLUSIONS: The palpation-directed rotator interval anterior approach technique for intra-articular glenohumeral MR arthrogram injections performed by a single surgeon was 88.9% accurate. LEVEL OF EVIDENCE: Level IV, therapeutic case series.

Effect of thematic map misclassification on landscape multi-metric assessment.

Advancements in remote sensing and computational tools have increased our awareness of large-scale environmental problems, thereby creating a need for monitoring, assessment, and management at these scales. Over the last decade, several watershed and regional multi-metric indices have been developed to assist decision-makers with planning actions of these scales. However, these tools use remote-sensing products that are subject to land-cover misclassification, and these errors are rarely incorporated in the assessment results. Here, we examined the sensitivity of a landscape-scale multi-metric index (MMI) to error from thematic land-cover misclassification and the implications of this uncertainty for resource management decisions. Through a case study, we used a simplified floodplain MMI assessment tool, whose metrics were derived from Landsat thematic maps, to initially provide results that were naive to thematic misclassification error. Using a Monte Carlo simulation model, we then incorporated map misclassification error into our MMI, resulting in four important conclusions: (1) each metric had a different sensitivity to error; (2) within each metric, the bias between the error-naive metric scores and simulated scores that incorporate potential error varied in magnitude and direction depending on the underlying land cover at each assessment site; (3) collectively, when the metrics were combined into a multi-metric index, the effects were attenuated; and (4) the index bias indicated that our naive assessment model may overestimate floodplain condition of sites with limited human impacts and, to a lesser extent, either over- or underestimated floodplain condition of sites with mixed land use.

Mimicry and eavesdropping enable a new form of social parasitism in ants.

Social parasitism is defined by the exploitation of the social mechanisms of one society by another whole society. Here, we use quantitative ecological data and experiments to identify the components of a new form of social parasitism by the recently discovered "mirror turtle ant," Cephalotes specularis. We show that C. specularis workers visually mimic and actively avoid contact with foragers of the hyperaggressive host ant Crematogaster ampla, allowing them to move freely in the extensive and otherwise defended foraging networks of host colonies. Workers from parasite colonies have immediate access to these networks by nesting exclusively within host territories, and 89% of all potential host territories were parasitized. Inside the network, parasite workers eavesdrop on the host's trail pheromones to locate and exploit food resources that are defended by the host to the exclusion of all other ants. Experiments demonstrated the unprecedented capacity of the parasite for superior foraging performance on its host's pheromone trails than on trails of its own. Considered together, the apparent Batesian-Wallacian mimicry, pheromone-based interceptive eavesdropping, kleptoparasitism, and xenobiotic nesting ecology displayed by C. specularis within the territory and foraging network of a host ant represents a novel adaptive syndrome for social exploitation.

Stability and phylogenetic correlation in gut microbiota: lessons from ants and apes.

Correlation between gut microbiota and host phylogeny could reflect codiversification over shared evolutionary history or a selective environment that is more similar in related hosts. These alternatives imply substantial differences in the relationship between host and symbiont, but can they be distinguished based on patterns in the community data themselves? We explored patterns of phylogenetic correlation in the distribution of gut bacteria among species of turtle ants (genus Cephalotes), which host a dense gut microbial community. We used 16S rRNA pyrosequencing from 25 Cephalotes species to show that their gut community is remarkably stable, from the colony to the genus level. Despite this overall similarity, the existing differences among species' microbiota significantly correlated with host phylogeny. We introduced a novel analytical technique to test whether these phylogenetic correlations are derived from recent bacterial evolution, as would be expected in the case of codiversification, or from broader shifts more likely to reflect environmental filters imposed by factors such as diet or habitat. We also tested this technique on a published data set of ape microbiota, confirming earlier results while revealing previously undescribed patterns of phylogenetic correlation. Our results indicated a high degree of partner fidelity in the Cephalotes microbiota, suggesting that vertical transmission of the entire community could play an important role in the evolution and maintenance of the association. As additional comparative microbiota data become available, the techniques presented here can be used to explore trends in the evolution of host-associated microbial communities.

Description of Cephalotes specularis n. sp. (Formicidae: Myrmicinae)-the mirror turtle ant.

We describe here Cephalotes specularis n. sp. (Hymenoptera: Formicidae: Myrmicinae: Cephalotini) based on minor and major workers, gynes and larvae from Uberlândia, Minas Gerais state, Brazil. On morphological and molecular grounds, this new species belongs to the C. fiebrigi + C. bruchi species complex, of which there are 11 previously described species (one in C. bruchi group and 10 in the C. fiebrigi group). All members of these groups are found in, or are limited to the South American "arid diagonal", comprised of the Argentinian Chaco, the Cerrados of central South America, and the Brazilian northeastern caatingas. Workers of C. specularis n. sp. have an extremely shiny gaster which is mirror-like, notwithstanding its sparse covering by minute hairs. This species engages in a form of resource-based social parasitism of the host ant Crematogaster ampla (Myrmicinae: Crematogastrini). Cephalotes specularis foragers move freely in the dense traffic of Crematogaster ampla foraging trails. They exhibit highly atypical body posturing for turtle ants, which makes them hard to distinguish from the Crematogaster foragers.