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Architected materials such as lattices are capable of demonstrating extraordinary mechanical performance. Lattices are often used for their stretch-dominated behavior, which gives them a high degree of stiffness at low-volume fractions. At the other end of the stiffness spectrum, bending-dominated lattices tend to be more compliant and are of interest for their energy absorption performance. Here, we report a class of ultra-compliant interwoven lattices that demonstrate up to an order of magnitude improvement in compliance over their traditional counterparts at similar volume fractions. This is achieved by selectively decoupling nodes and interweaving struts in bending-dominated lattices, inspired by observations of this structural principle in the lattice-like arrangement of the Venus flower basket sea sponge. By decoupling nodes in this manner, we demonstrate a simple and near-universal design strategy for modulating stiffness in lattice structures and achieve among the most compliant lattices reported in the literature.

Ant cuticle texture presumably provides some type of function, and therefore is useful to research for ecological applications and bioinspired designs. In this study, we employ statistical image texture analysis and deep machine learning methods to classify similar ant species based on morphological features. We establish a public database of ant cuticle images for research. We provide a comparative study of the performance of image texture classification and deep machine learning methods on this ant cuticle dataset. Our results show that the deep learning methods give higher accuracy than statistical methods in recognizing ant cuticle textures. Our experiments also reveal that the deep learning networks designed for image texture performs better than the general deep learning networks.

Biodiversity provides a massive library of ideas for bio-inspired design, but the sheer number of species to consider can be daunting. Current approaches for sifting through biodiversity to identify relevant biological models include searching for champion adapters that are particularly adept at solving a specific design challenge. While the champion adapter approach has benefits, it tends to focus on a narrow set of popular models while neglecting the majority of species. An alternative approach to bio-inspired design is the comparative method, which leverages biodiversity by drawing inspiration across a broad range of species. This approach uses methods in phylogenetics to map traits across evolutionary trees and compare trait variation to infer structure–function relationships. Although comparative methods have not been widely used in bio-inspired design, they have led to breakthroughs in studies on gecko-inspired adhesives and multifunctionality of butterfly wing scales. Here we outline how comparative methods can be used to complement existing approaches to bio-inspired design, and we provide an example focused on bio-inspired lattices, including honeycomb, and glass sponges. We demonstrate how comparative methods can lead to breakthroughs in bio-inspired applications as well as answer major questions in biology, which can strengthen collaborations with biologists and produce deeper insights into biological function.

(2019). Hidden Treasures. Discovering the Design Potential of Natural History Collections. The Design Journal: Vol. 22, Running with Scissors: 13th International Conference of the European Academy of Design, Dundee, 10-12th April, 2019, edited by Louise Valentine, Joanna Bletcher and Leon Cruickshank, pp. 2189-2195.

Phenotypic plasticity allows organisms to respond to changing environments throughout their lifetime, but these changes are rarely reversible. Exceptions occur in relatively long-lived vertebrate species that exhibit seasonal plasticity in brain size, although similar changes have not been identified in short-lived species, such as insects. Here, we investigate brain plasticity in reproductive workers of the antHarpegnathos saltator. Unlike most ant species, workers ofH. saltatorare capable of sexual reproduction, and they compete in a dominance tournament to establish a group of reproductive workers, termed ‘gamergates'. We demonstrated that, compared to foragers, gamergates exhibited a 19% reduction in brain volume in addition to significant differences in behaviour, ovarian status, venom production, cuticular hydrocarbon profile, and expression profiles of related genes. In experimentally manipulated gamergates, 6–8 weeks after being reverted back to non-reproductive status their phenotypes shifted to the forager phenotype across all traits we measured, including brain volume, a trait in which changes were previously shown to be irreversible in honeybees andDrosophila. Brain plasticity inH. saltatoris therefore more similar to that found in some long-lived vertebrates that display reversible changes in brain volume throughout their lifetimes.

Because ectotherm activity and metabolism are sensitive to temperature, terrestrial arthropods may be especially responsive to ongoing climatic warming. Here, we quantified responses of arthropod abundance to two years of warming in an outdoor temperature manipulation experiment at Duke Forest, North Carolina, USA. Nine open‐top chambers were individually heated year‐round from 1.5° to 5.5°C above ambient temperature. From two years of monthly pitfall trapping, we collected and identified 4,468 arthropods representing 24 orders. We initially predicted that arthropods would experience the greatest negative effects of experimental warming during the summer months, when temperatures reach their yearly maximum and arthropods may be close to their maximum thermal tolerance limits. Instead, we found that the strongest negative effects on arthropod abundance occurred during the winter and spring, when ambient temperatures are relatively cooler, whereas the effects of experimental warming on abundance were not significant during the summer or fall. During the spring of 2012, the warmest spring on record for the southeastern USA, total arthropod abundance declined 20% per °C of experimental warming. Abundance declines were driven largely by flies (Diptera), which were the most abundant insect order, representing approximately a third of all arthropods collected. The most abundant arthropod family, Mycetophilidae (fungus gnats), declined 64% per °C of warming during the spring of 2012. Although previous research on climatic warming has focused on the impact of maximum yearly temperatures on organismal performance, our results are more consistent with the cool‐season sensitivity hypothesis, which posits that arthropods adapted for cooler conditions are likely to face the strongest negative effects of warming during the cooler seasons.

Collaborations between artists and scientists have a long history. In recent years, artists have joined with ecologists to showcase biodiversity, links between biodiversity and ecosystem function, and the effects of human activities on the broader environment. In many cases, artists also have provided “broader impacts” for ecological research activities, communicating scientific findings in creative and novel ways to audiences much broader than the readership of our technical journals.

Despite the remarkable diversity of food resources exploited by ants, fungi are rarely known to be part of their diet. The notable exception is the fungal-feeding specialists in the tribe Attini, which cultivate fungi inside their nests in what is one of the earliest forms of agriculture. Previous accounts of fungivory in ants outside these fungal-feeding specialists have been questioned due to whether or not ants consume fungal tissue or prey on mycophagous insect larvae present in or on mushrooms. Here we show that ants in the widespread genus Aphaenogaster recruit to mushroom baits in the field regardless of whether or not mushrooms contained insects upon which ants might prey. When dye-stained mushrooms were provided to colonies in the lab, ants fed on mushroom tissue and dye was visible throughout their digestive tract. Evidence of mushroom feeding in Aphaenogaster suggests that facultative fungal feeding is more common in ants than previously reported, including within the myrmecine clade that contains the attines. Previous accounts of fungivory in ants have been limited to fungal-feeding specialists, but mushroom feeding by Aphaenogaster shows that fungi can also be part of a generalist ant diet. By feeding on mushrooms and transporting mushroom tissues back to the nest, Aphaenogaster workers may also serve as dispersers of fungal spores.

Numerous studies have identified cuticular compounds that distinguish adult queens from workers in social insect colonies, but how future queens are identified at the larval stage is poorly understood. Nevertheless, the ability of workers to discriminate queen and worker larvae is necessary for them to regulate caste determination and queen production. In the ant Harpegnathos saltator, workers bite larvae to inhibit queen development, and we used biting as an assay to test how workers identify queens at the larval stage. The transfer of cuticular compounds from queen to worker larvae through direct physical contact (rubbing) or using a hexane extract both elicited biting. Gas chromatography revealed significant differences in cuticular hydrocarbon profiles of queen and worker larvae that could be induced by treatment with a juvenile hormone (JH) analogue. Finally, treatment of male larvae with a JH analogue also elicited worker biting, which suggests a direct connection between JH levels and the production of a larval queen signal. These results demonstrate that workers identify larval caste using a chemical signal present on the cuticle, a ‘princess pheromone’, that reflects endocrine changes associated with queen development. Based on the connection between JH levels and the production of a larval queen signal, we developed a model for caste determination in H. saltator that incorporates endocrine, pheromonal and behavioural control of caste development.

Few studies have quantified the relative importance of direct effects of climate change on communities versus indirect effects that are mediated thorough species interactions, and the limited evidence is conflicting. Trait-based approaches have been popular in studies of climate change, but can they be used to estimate direct versus indirect effects? At the species level, thermal tolerance is a trait that is often used to predict winners and losers under scenarios of climate change. But thermal tolerance might also inform when species interactions are likely to be important because only subsets of species will be able to exploit the available warmer climatic niche space, and competition may intensify in the remaining, compressed cooler climatic niche space. Here, we explore the relative roles of the direct effects of temperature change and indirect effects of species interactions on forest ant communities that were heated as part of a large-scale climate manipulation at high- and low-latitude sites in eastern North America. Overall, we found mixed support for the importance of negative species interactions (competition), but found that the magnitude of these interaction effects was predictable based on the heat tolerance of the focal species. Forager abundance and nest site occupancy of heat-intolerant species were more often influenced by negative interactions with other species than by direct effects of temperature. Our findings suggest that measures of species-specific heat tolerance may roughly predict when species interactions will influence responses to global climate change.

Summary How species respond to temperature change depends in large part on their physiology. Physiological traits, such as critical thermal limits (CTmax and CTmin), provide estimates of thermal performance but may not capture the full impacts of temperature on fitness. Rather, thermal performance likely depends on a combination of traits—including thermal limits—that vary among species. Here we examine how thermal limits correlate with the main components that influence fitness in ants. First, we compare how temperature affected colony survival and growth in two ant species that differ in their responses to warming in the field—Aphaenogaster rudis (heat-intolerant) and Temnothorax curvispinosus (heat-tolerant). We then extended our study to compare CTmax, thermal requirements of brood, and yearly activity season among a broader set of ant species. While thermal limits were higher for workers of T. curvispinosus than A. rudis, T. curvispinosus colonies also required higher temperatures for survival and colony growth. This pattern generalized across 17 ant species, such that species whose foragers had a high CTmax also required higher temperatures for brood development. Finally, species whose foragers had a high CTmax had relatively short activity seasons compared with less heat-tolerant species. The relationships between CTmax, thermal requirements of brood, and seasonal activity suggest two main strategies for growth and development in changing thermal environments: one where ants forage at higher temperatures over a short activity season, and another where ants forage at lower temperatures for an extended activity season. Where species fall on this spectrum may influence a broad range of life-history characteristics and aid in explaining the current distributions of ants as well as their responses to future climate change. This article is protected by copyright. All rights reserved.

Nests provide a buffer against environmental variation, but conditions may also vary at different locations within a nest. Conditions can vary based on abiotic factors, such as moisture and temperature, as well as biotic factors, such as the presence of microbes and potential pathogens. Therefore, characterizing how animals adjust their position inside their nests to track microclimate preferences while at the same time preventing pathogen exposure is necessary to understand the benefits nests provide. Here we studied how colonies of the acorn-nesting ant Temnothorax curvispinosus responded to experimental manipulation of moisture, temperature, and microbial growth inside their nests. Colonies showed no response to differences in moisture and moved to the bottom of the acorn regardless of moisture treatment. When nests were heated from the top to simulate warming by the sun, workers preferentially moved brood to the warm, upper half of the acorn, which would stimulate brood development. Finally, the strongest factor that influenced colony position was the presence of microbes inside the nest—colonies avoided the bottom of the nest when it was inoculated with microbes, and colonies in new acorns shifted to the top of the acorn over time as mold and other microbes had time to grow. The relatively strong response of T. curvispinosus to microbial growth inside their nests suggests that pathogen pressures—in addition to microclimate—have a significant impact on how colonies use nest spaces. Social insects are known to invest heavily in antimicrobial compounds that kill or slow the growth of microbes, but avoidance may represent an additional line of defense to prevent pathogen exposure.

Establishment and maintenance of the reproductive division of labor within social insect colonies relies on clear communication between nestmates. Fertile members convey their status to prevent others from becoming reproductively active. Recent findings in some basal termites indicate that cuticular hydrocarbon profiles may indicate reproductive state, but there is little evidence to show a direct link between reproductive status and hydrocarbon production—a prerequisite for an “honest” fertility signal. Here, we report that the putative signaling mechanism is influenced by juvenile hormone (JH), a primary regulator of gonadal development and activity in insects. Topical application of a JH-analog (pyriproxyfen) to reproductively inactive alates of the basal dampwood termite Zootermopsis nevadensis induced both females and males to express significantly more of a reproductive-specific hydrocarbon (6,9,17-tritriacontatriene). However, the JH-analog did not significantly enhance gonadal development or activity in treated termites beyond what is usually observed in maturing alates released from the inhibitory stimuli of their natal nest. These results suggest that a rise in JH following disinhibition drives the expression of reproductive-specific hydrocarbons, but that an individual’s hydrocarbon profile is not directly linked to its gonadal state. Rather than directly driving the expression of reproductive-specific hydrocarbons, the gonads may act indirectly through their influence on circulating JH.

Social dominance hierarchies are widespread, but little is known about the mechanisms that produce nonlinear structures. In addition to despotic hierarchies, where a single individual dominates, shared hierarchies exist, where multiple individuals occupy a single rank. In vertebrates, these complex dominance relationships are thought to develop from interactions that require higher cognition, but similar cases of shared dominance have been found in social insects. Combining empirical observations with a modeling approach, we show that all three hierarchy structures-linear, despotic, and shared-can emerge from different combinations of simple interactions present in social insects. Our model shows that a linear hierarchy emerges when a typical winner-loser interaction (dominance biting) is present. A despotic hierarchy emerges when a policing interaction is added that results in the complete loss of dominance status for an attacked individual (physical policing). Finally, a shared hierarchy emerges with the addition of a winner-winner interaction that results in a positive outcome for both interactors (antennal dueling). Antennal dueling is an enigmatic ant behavior that has previously lacked a functional explanation. These results show how complex social traits can emerge from simple behaviors without requiring advanced cognition.

The honeybee&rsquo, s comb has inspired the design of engineering honeycomb core that primarily abstract the hexagonal cell shape and exploit its mass minimizing properties to construct lightweight panels. This work explored three additional design features that are part of natural honeybee comb but have not been as well studied as design features of interest in honeycomb design: the radius at the corner of each cell, the coping at the top of the cell walls, and the interface between cell arrays. These features were first characterized in natural honeycomb using optical and X-ray techniques and then incorporated into honeycomb core design and fabricated using an additive manufacturing process. The honeycomb cores were then tested in out-of-plane compression and bending, and since all three design features added mass to the overall structure, all metrics of interest were examined per unit mass to assess performance gains despite these additions. The study concluded that the presence of an interface increases specific flexural modulus in bending, with no significant benefit in out-of-plane compression, coping radius positively impacts specific flexural strength, however, the corner radius has no significant effect in bending and actually is slightly detrimental for out-of-plane compression testing.

The chemosensory system is key to establishing and maintaining social structure in eusocial insects. Ants exhibit cooperative colonial behaviors reflective of an advanced form of sociality with an extensive dependency on communication. Cuticular hydrocarbons (CHCs) serve as pheromones and cues that regulate multiple aspects of social interactions and behaviors in ants. The perception of CHCs entails odorant receptor neurons (ORNs) that express specific odorant receptors (ORs) encoded by a dramatically expanded Or gene family in ants. Until recently, studies of the biological functions of ORs in eusocial insects were stymied by the lack of genetic tools. In most eusocial insect species, only one or a few queens in a colony can transmit the genetic information to their progeny. In contrast, any worker in the ant Harpegnathos saltator can be converted into a gamergate (pseudo-queen), and used as a foundress to engender an entire new colony and be crossed for genetic experiments. This feature facilitated CRISPR-Cas9 gene targeting to generate a germline mutation in the orco gene that encodes the obligate co-receptor whose mutation should significantly impact ant olfaction. Our results show that Orco exhibits a conserved role in the perception of general odorants but also a role in reproductive physiology and social behavior plasticity in ants. Surprisingly, and in contrast to other insect systems, the loss of OR functionality also dramatically reduces the development of the ant antennal lobe where ORNs project. Taken together, these findings open the possibility of studying the genetics of eusociality and provide inroads towards understanding the function of the expanded ORs family in eusocial insects in regulating caste determination, social communication and neuronal plasticity.

The microbiome of built structures has considerable influence over an inhabitant's well-being, yet the vast majority of research has focused on human-built structures. Ants are well-known architects, capable of constructing elaborate dwellings, the microbiome of which is underexplored. Here, we explore the bacterial and fungal microbiomes in functionally distinct chambers within and outside the nests of Azteca alfari ants in Cecropia peltata trees. We predicted that A. alfari colonies (1) maintain distinct microbiomes within their nests compared to the surrounding environment, (2) maintain distinct microbiomes among nest chambers used for different functions, and (3) limit both ant and plant pathogens inside their nests. In support of these predictions, we found that internal and external nest sampling locations had distinct microbial communities, and A. alfari maintained lower bacterial richness in their ‘nurseries’. While putative animal pathogens were suppressed in chambers that ants actively inhabited, putative plant pathogens were not, which does not support our hypothesis that A. alfari defends its host trees against microbial antagonists. Our results show that ants influence microbial communities inside their nests similar to studies of human homes. Unlike humans, ants limit the bacteria in their nurseries and potentially prevent the build-up of insect-infecting pathogens. These results highlight the importance of documenting how indoor microbiomes differ among species, which might improve our understanding of how to promote indoor health in human dwellings.

More than 13,000 ant species have been formally described, yet the general public recognizes only two main categories: red ants and black ants. This system has some limitations, so there has been increased effort to provide additional sensory information for ant identification, including the sense of smell. Nowhere has this been more strongly applied than to the “odorous house ant” ( Tapinoma sessile Say), a common household pest in North America that releases a curious odor when crushed (Smith 1928). At first, using scent to identify ants seems obvious and practical, because ants themselves communicate through smell. However, the sense of smell in humans is far less developed, and there has been recent controversy over what, exactly, the odorous house ant smells like. This species belongs to a …

The differences between adult male and female ants are often striking and obvious, yet both sexes appear virtually identical at the larval stage. Current methods for determining larval sex rely on genetic analyses or histology, both of which require killing all larvae examined. Here, we describe a method for identifying larval sex in vivo based on visible differences in genital imaginal discs. Using a light microscope, clear differences in genital disc morphology were observed between male and female larvae of the ponerine ant Harpegnathos saltator. Next, we investigated whether this technique could be broadly applied within ants and found similar differences in genital discs between male and female larvae of Aphaenogaster cockerelli and Camponotus floridanus. Taken together, our results show that genital discs can be used as a reliable indicator of larval sex in species from at least three major ant subfamilies. This technique should facilitate research into topics where information about larval sex is required.

The frequency of warm winter days is increasing under global climate change, but how organisms respond to warmer winters is not well understood. Most studies focus on growing season responses to warming. Locomotor performance is often highly sensitive to temperature, and can determine fitness outcomes through a variety of mechanisms including resource acquisition and predator escape. As a consequence, locomotor performance, and its impacts on fitness, may be strongly affected by winter warming in winter-active species. Here we use the acorn ant, Temnothorax curvispinosus, to explore how thermal performance (temperature-driven plasticity) in running speed is influenced by experimental winter warming of 3-5 degrees C above ambient in a field setting. We used running speed as a measure of performance as it is a common locomotor trait that influences acquisition of nest sites and food in acorn ants. Experimental winter warming significantly altered thermal performance for running speed at high (26 and 36 degrees C) but not low test temperatures (6 and 16 degrees C). Although we saw little differentiation in thermal performance at cooler test temperatures, we saw a marked increase in running speed at the hotter test temperatures for ants that experienced warmer winters compared with those that experienced cooler winters. Our results provide evidence that overwintering temperatures can substantially influence organismal performance, and suggest that we cannot ignore overwintering effects when forecasting organismal responses to environmental changes in temperature.

Ecological diversification into thermally divergent habitats can push species toward their physiological limits, requiring them to accommodate temperature extremes through plastic or evolutionary changes that increase persistence under the local thermal regime. One way to withstand thermal stress is to increase production of heat shock proteins, either by maintaining higher baseline abundance within cells or by increasing the magnitude of induction in response to heat stress. We evaluated whether environmental variation was associated with expression of three heat shock protein genes in two closely-related species of woodland ant, Aphaenogaster picea and A. rudis. We compared adult workers from colonies collected from 25 sites across their geographic ranges. Colonies were maintained at two different laboratory temperatures, and tested for the independent effects of environment, phylogeny, and acclimation temperature on baseline and heat-induced gene expression. The annual maximum temperature at each collection site (Tmax) was not a significant predictor of either baseline expression or magnitude of induction of any of the heat shock protein genes tested. A phylogenetic effect was detected only for basal expression of Hsp40, which was lower in the most southern populations of A. rudis and higher in a mid-range population of possible hybrid ancestry. In contrast, a higher acclimation temperature significantly increased baseline expression of Hsc70-4, and increased induction of Hsp40 and Hsp83. Thus, physiological acclimation to temperature variation appears to involve modulation of the heat shock response, whereas other mechanisms are likely to be responsible for evolutionary shifts in thermal performance associated with large-scale climate gradients.

In lower termites, functionally sterile larval helpers are totipotent—capable of becoming reproductively active with the loss of their colony’s king or queen. Full reproductive development may take several weeks, but initiation of this developmental response most likely occurs shortly after colony members detect when a reproductive-specific signal is missing. We investigated the early response of termite helpers to the removal of their king and queen in the basal termite species Zootermopsis nevadensis. Within 6–12 h after reproductives were removed, helpers displayed an increase in head-butting—a behavior associated with dominance in other termite species as well as in closely related roaches. The loss of just one reproductive, either the king or queen, was also sufficient to cause an increase in head-butting. We did not find evidence, however, that this response was sex-specific: males and females were equally likely to increase head-butting independent of the sex of the reproductive that was removed. Finally, we discovered that reproductive-specific compounds present on the cuticle of king and queen termites were also present in their feces, but the presence of the feces did not seem sufficient to inhibit the increased head-butting after the reproductives were removed. Collectively, these results indicate that termite workers readily detect the loss of reproductives in their colony and that they at least initially respond in a non sex-specific manner.

Extreme developmental plasticity within the eusocial insects defines the separation between the queen and worker castes. The switch between these 2 developmental pathways is thought to be under tight social control. Although a large emphasis has been placed on the effect of larval nutrition on caste determination, workers of many species have no direct control over larval feeding. This may be particularly relevant to the early evolution of ants, when behaviors that allow fine control over larval nutrition, such as mouth-to-mouth food exchange between larvae and workers, were probably not yet present. We investigated larval-directed aggression as an alternative means to regulate queen development in the ant Harpegnathos saltator, a species that retains ancestral characteristics. We tested worker response toward natural queen-destined larvae and larvae induced to develop as queens using a juvenile hormone analog (JHA). Workers from colonies that were not rearing queens bit queen-destined larvae, whereas worker-destined larvae were not attacked. When colonies were naturally rearing queens, workers did not bite JHA-treated larvae, and a larger proportion of these larvae developed into queens compared with larvae that received biting. This supports the hypothesis that workers of H. saltator use biting to inhibit queen development during periods when the conditions for queen rearing are not met. We propose that mechanical stress, in addition to nutrition, could serve as a mechanism to regulate queen development in species that lack fine control over larval food consumption, and this may have played a role during the early evolution of eusociality in ants. Key words: aggression, developmental plasticity, juvenile hormone, queen determination, social regulation, stress. [Behav Ecol]

The regulation of reproduction within insect societies is a key component of the evolution of eusociality. Differential patterns of hormone levels often underlie the reproductive division of labor observed among colony members, and further task partitioning among workers is also often correlated with differences in juvenile hormone (JH) and ecdysteroid content. We measured JH and ecdysteroid content of workers and queens of the ant Harpegnathos saltator. In this species, new colonies are founded by a single queen, but after she dies workers compete in an elaborate dominance tournament to decide a new group of reproductives termed "gamergates." Our comparisons revealed that queens, gamergates, and inside workers (non-reproductive) did not differ in levels of JH or ecdysteroids. However, increased JH and decreased ecdysteroid content was observed in outside workers exhibiting foraging behavior. Application of a JH analog to virgin queens of H. saltator, although effective at inducing dealation, failed to promote egg production. Together, these results support the hypothesis that JH has lost its reproductive function in H. saltator to regulate foraging among the worker caste.

Nest structure in ants is often designed to optimize the colony’s ability to thermoregulate, and this specialization is most highly developed in mound-building ant species. Solenopsis invicta invest a large amount of energy in building mounds and transporting their brood up and down in their nests as a means of thermoregulation. Because few ant species build true mounds, we wanted to determine the effectiveness of these mounds in harvesting solar heat as well as to distinguish what factors (temperature vs. circadian rhythm) govern where fire ants place their brood in the mound and when they place it. We measured temperature patterns in the mound over several days at different depths and under different conditions (under direct sunlight or shade), and then conducted a series of field experiments to manipulate the orientation and time of heating. On cool mornings in spring or fall, surface temperatures of the mound rise at the fastest rate on the side receiving the most direct sunlight (usually the south side). This heating causes a temperature gradient through different depths in the mound, and shows little difference from outside ground temperature at a depth greater than ~40 cm inside the nest. In the morning, fire ants move their brood up into the mound on the side most directly heated, and when temperatures exceed optimal (~32°C) they move their brood down the temperature gradient to lower depths in the nest. In addition to this, mound temperature does not only increase due to direct sunlight, but temperature also increases higher than ground temperatures when the mound is in the shade due to its low specific heat. Experiments in which sunlight was mirrored to the normally shaded side of the mound, or when mounds were heated at night, revealed that S. invicta primarily track temperature patterns and do not rely on behavioral habits or circadian rhythms for the thermoregulatory transport of their brood. When mounds were shaded, S. invicta brood was evenly distributed directly under the surface of the mound rather than aggregating towards a specific side. The fire ant mound is important for thermoregulation because, compared to moundless subterranean nests, it absorbs heat more rapidly both in direct sunlight and shady conditions. Temperature tracking within the nest is key to understanding thermoregulatory placement of fire ant brood, as well as insight into the production of sexual brood and reproduction.

The amount of energy consumed within an average city block is an order of magnitude higher than that consumed in any other ecosystem over a similar area. This is driven by human food inputs, but the consequence of these resources for urban animal populations is poorly understood. We investigated the role of human foods in ant diets across an urbanization gradient in Manhattan using carbon and nitrogen stable isotopes. We found that some—but not all—ant species living in Manhattan's most urbanized habitats had δ 13 C signatures associated with processed human foods. In particular, pavement ants ( Tetramorium sp. E) had increased levels of δ 13 C similar to δ 13 C levels in human fast foods. The magnitude of this effect was positively correlated with urbanization. By contrast, we detected no differences in δ 15 N, suggesting Tetramorium feeds at the same trophic level despite shifting to human foods. This pattern persisted across the broader ant community; species in traffic islands used human resources more than park species. Our results demonstrate that the degree urban ants exploit human resources changes across the city and among species, and this variation could play a key role in community structure and ecosystem processes where human and animal food webs intersect.

Social insects live in dense groups with a high probability of disease transmission and have therefore faced strong pressures to develop defences against pathogens. For this reason, social insects have been hypothesized to invest in antimicrobial secretions as a mechanism of external immunity to prevent the spread of disease. However, empirical studies linking the evolution of sociality with increased investment in antimicrobials have been relatively few. Here we quantify the strength of antimicrobial secretions among 20 ant species that cover a broad spectrum of ant diversity and colony sizes. We extracted external compounds from ant workers to test whether they inhibited the growth of the bacterium Staphylococcus epidermidis . Because all ant species are highly social, we predicted that all species would exhibit some antimicrobial activity and that species that form the largest colonies would exhibit the strongest antimicrobial response. Our comparative approach revealed that strong surface antimicrobials are common to particular ant clades, but 40% of species exhibited no antimicrobial activity at all. We also found no correlation between antimicrobial activity and colony size. Rather than relying on antimicrobial secretions as external immunity to control pathogen spread, many ant species have probably developed alternative strategies to defend against disease pressure.

Ants exhibit cooperative behaviors and advanced forms of sociality that depend on pheromone-mediated communication. Odorant receptor neurons (ORNs) express specific odorant receptors (ORs) encoded by a dramatically expanded gene family in ants. In most eusocial insects, only the queen can transmit genetic information, restricting genetic studies. In contrast, workers in Harpegnathos saltator ants can be converted into gamergates (pseudoqueens) that can found entire colonies. This feature facilitated CRISPR-Cas9 generation of germline mutations in orco, the gene that encodes the obligate co-receptor of all ORs. orco mutations should significantly impact olfaction. We demonstrate striking functions of Orco in odorant perception, reproductive physiology, and social behavior plasticity. Surprisingly, unlike in other insects, loss of OR functionality also dramatically impairs development of the antennal lobe to which ORNs project. Therefore, the development of genetics in Harpegnathos establishes this ant species as a model organism to study the complexity of eusociality.

Summary Dominance rank in animal societies is correlated with changes in both reproductive physiology and behavior. In some social insects, dominance status is used to determine a reproductive division of labor, where a few colony members reproduce while most remain functionally sterile. Changes in reproduction and behavior in this context must be coordinated through crosstalk between the brain and the reproductive system. We investigated a role for biogenic amines in forming this connection in the ant Harpegnathos saltator. In this species, workers engage in an elaborate dominance tournament to establish a group of reproductive workers termed gamergates. We analyzed biogenic amine content in the brains of gamergates, inside workers, and foragers under stable colony conditions and found that gamergates had the highest levels of dopamine. Dopamine levels were also positively correlated with increased ovarian activity among gamergates. Next, we experimentally induced workers to compete in a reproductive tournament to determine how dopamine may be involved in the establishment of a new hierarchy. Dopamine levels rose in aggressive workers at the start of a tournament, while workers that were policed by their nestmates (a behavior that inhibits ovarian activity) showed a rapid decline in dopamine. In addition to dopamine, levels of serotonin and tyramine differed among castes, and these changes could contribute to differences in caste-specific behavioral patterns observed among non-reproductive workers. Overall, these results provide support that biogenic amines link changes in behavior and dominance with reproductive activity in H. saltator as well as drive differences in worker task performance.

Physiological intolerance of high temperatures places limits on organismal responses to the temperature increases associated with global climatic change. Because ants are geographically widespread, ecologically diverse, and thermophilic, they are an ideal system for exploring the extent to which physiological tolerance can predict responses to environmental change. Here, we expand on simple models that use thermal tolerance to predict the responses of ants to climatic warming. We investigated the degree to which changes in the abundance of ants under warming reflect reductions in the thermal niche space for their foraging. In an eastern deciduous forest system in the United States with approximately 40 ant species, we found that for some species, the loss of thermal niche space for foraging was related to decreases in abundance with increasing experimental climatic warming. However, many ant species exhibited no loss of thermal niche space. For one well-studied species, Temnothorax curvispinosus, we examined both survival of workers and growth of colonies (a correlate of reproductive output) as functions of temperature in the laboratory, and found that the range of thermal tolerances for colony growth was much narrower than for survival of workers. We evaluated these functions in the context of experimental climatic warming and found that the difference in the responses of these two attributes to temperature generates differences in the means and especially the variances of expected fitness under warming. The expected mean growth of colonies was optimized at intermediate levels of warming (2-4°C above ambient); yet, the expected variance monotonically increased with warming. In contrast, the expected mean and variance of the survival of workers decreased when warming exceeded 4°C above ambient. Together, these results for T. curvispinosus emphasize the importance of measuring reproduction (colony growth) in the context of climatic change: indeed, our examination of the loss of thermal niche space with the larger species pool could be missing much of the warming impact due to these analyses being based on survival rather than reproduction. We suggest that while physiological tolerance of temperature can be a useful predictive tool for modeling responses to climatic change, future efforts should be devoted to understanding the causes and consequences of variability in models of tolerance calibrated with different metrics of performance and fitness.

A link between hormones and developmental plasticity has long been established, but understanding how evolution has shaped the physiological systems underlying plasticity remains a major question. Within the eusocial insects, developmental plasticity helps define a reproductive division of labor through the production of distinct queen and worker castes. Caste determination may be triggered via changes in juvenile hormone (JH) levels during specific JH-sensitive periods in development. The timing of these periods, however, can vary and may relate to phenotypic differences observed among species. In order to gain insight into the evolution of caste determining systems in eusocial insects, we investigated the presence of a JH-sensitive period for queen determination in the ant Harpegnathos saltator. This species displays a number of ancestral characteristics, including low queen-worker dimorphism, and should allow insight into the early evolution of caste determining systems in ants. We identified four larval instars in H. saltator, and we found that the application of a JH analog (JHA) to third and fourth instar larvae induced queen development while treatment of early instars did not. This indicated the presence of a JH-sensitive period for queen determination at the end of the larval stage. These results contrast with what has been found in other ant species, where queen determination occurs much earlier in development. Therefore, our results suggest that caste determination originally occurred late in the larval stage in the ancestral condition but has shifted earlier in development in species that began to acquire advanced characteristics. This shift may have facilitated the development of greater queen-worker dimorphism as well as multiple worker castes.

Concern for honey bee health has implicated both urbanization and nutritional stress as factors contributing to honey bee declines. The expansion of urban areas has resulted in reduced foraging habitat for bees, while at the same time introducing new food sources, such as foods unintentionally provided by humans as litter or in waste containers. While human foods play an important role in the diets of other urban animals, the extent to which honey bees feed on these resources has not been well characterized. Here, we compared the amount of human foods in honey bee diets across a rural-urban gradient in North Carolina, USA, using stable isotopes of carbon (δ13C). Human-produced sugars—primarily sugarcane and corn syrup—have a characteristic isotope signature that can be used to quantify the relative amount of human foods in animal diets. We predicted that urban bees would have an increase in δ13C if they were feeding on human-produced sugars, but we found no change in δ13C between urban and rural colonies. Instead, we found an increase in δ13C in managed bees in both habitats, which is indicative of supplemental sugar feeding by beekeepers. Rather than switching to human food sources, urban bees seem to be feeding on urban flowers or insect-produced honeydew. This suggests an important role for urban flowers and green spaces in maintaining healthy pollinator populations in cities.

The spatial organization within a social insect colony is a key component of colony life. It influences individual interaction rates, resource distribution, and division of labor within the nest. Yet studies of social insect behavior are most often carried out in artificial constructions, which may change worker behavior and colony organization. We observed how workers of the ant Pheidole rhea organized brood in nests with deep chambers and textured walls that were designed to mimic their natural constructions more closely. Instead of clumping larvae into piles on the chamber floor, workers suspended fourth-instar larvae from the vertical walls and ceiling of each chamber while young larvae and pupae were clumped at the base. Fourth-instar larvae possess five rows of anchor-tipped hairs on their dorsal side, and we predicted that these hairs functioned to attach larvae to the nest walls. We gave larvae "haircuts," where only the anchor-tipped hairs were removed, and then tested their ability to adhere to a textured surface raised to an angle of 90° and then 120° with respect to the horizontal plane. Larvae whose hairs had been clipped came unattached in almost all trials, while larvae whose hairs remained intact stayed attached. This confirmed that anchor-tipped hairs functioned to attach larvae to the walls of the nest. The presence of anchor-tipped hairs is widespread and has been documented in at least 22 genera from the ant subfamily Myrmicinae, including species that occur in a variety of environments and represent a broad range of nesting habits. Based on our results, it is likely that many species exhibit this larval hanging behavior, and this could impact colony characteristics such as spatial organization and the care of developing larvae by nurse workers.