This article has been retracted: please see Elsevier Policy on Article Withdrawal (https://www.elsevier.com/about/policies/article-withdrawal). This article has been retracted at the request of the authors C.N.K. and C.M.W. The data presented in this article, the data presented in Keiser et al. (Behavioural Processes, 2015), and the data presented in Keiser and Pruitt (Proceedings of the Royal Society B: Biological Sciences, 2014) were all collected in the same field experiment in 2014. A portion of the data presented in this article was published previously in Keiser and Pruitt (Proceedings of the Royal Society B: Biological Sciences, 2014) and this overlap was not disclosed at the time of publication. The aforementioned field experiment sought to answer two questions: how group phenotypic composition and group size influence collective behaviour (group-level analyses) and how individual traits influence participation in those collective behaviours (individual-level analyses). The experiment was designed and initiated by C.N.K. and J.N.P.; halfway through the experiment J.N.P. was replaced by C.M.W. until the conclusion of data collection. Keiser and Pruitt (Proceedings of the Royal Society B: Biological Sciences, 2014) published only the group-level analyses. C.M.W. took the lead on preparing the task participation data (individual-level analyses) for publication. C.N.K. and C.M.W. planned to publish only the individual-level data in Wright et al. (Animal Behaviour, 2015), because the group-level analyses were already published elsewhere. J.N.P. argued that the story would be incomplete without both levels of analysis and advised C.M.W. to include the group-level data in Wright et al. (Animal Behaviour, 2015). J.N.P. took responsibility for handling reviewer comments and reanalysing data and submitted the revised manuscript to Animal Behaviour. Recent inspection has identified minor discrepancies in the data between Wright et al. (Animal Behaviour, 2015) and Keiser and Pruitt (Proceedings of the Royal Society B: Biological Sciences, 2014), which should be identical, the source of which remains unknown. The data in Wright et al. (Animal Behaviour, 2015) are presented in four sections: 'task participation', 'state-dependent personality', 'collective foraging behaviour' and 'collective escape behaviour'. The first two sections are unique to this paper and the data have not been included in any other publications. The data in the second two sections had been previously published in Keiser and Pruitt (Proceedings of the Royal Society B: Biological Sciences, 2014). In the Animal Behaviour Guide for Authors (https://www.elsevier.com/journals/animal-behaviour/0003-3472/guide-for-authors), it states: 'Animal Behaviour will not consider submissions that have been published elsewhere, nor will it republish data found in other publications, unless the data are reevaluated to provide new information not found in the original'. As such, C.N.K. and C.M.W. request that this article is retracted. J.N.P. has not responded to the journal's request for comment. References Keiser, C. N., & Pruitt, J. N. (2014). Personality composition is more important than group size in determining collective foraging behaviour in the wild. Proceedings of the Royal Society B: Biological Sciences, 281, 20141424. https://doi.org/10.1098/rspb.2014.1424 Keiser, C. N., Wright, C. M., & Pruitt, J. N. (2015). Warring arthropod societies: social spider colonies can delay annihilation by predatory ants via reduced apparency and increased group size. Behavioural Processes, 119, 14-21. https://doi.org/10.1016/j.beproc.2015.07.005 Wright, C. M., Keiser, C. N., & Pruitt, J.N. (2015). Personality and morphology shape task participation, collective foraging and escape behaviour in the social spider Stegodyphus dumicola. Animal Behaviour, 105, 47-54. https://doi.org/10.1016/j.anbehav.2015.04.001

We write as medical and biological professionals who are increasingly concerned with how commercial and corporate interests of publishers are being allowed to unduly influence intellectual discourse, especially in relation to biological sex.We represent a variety of backgrounds, with interests ranging from malelethal genetic disorders in humans to sex behaviours in invertebrates.Human sex is an observable, immutable, and important biological classification; it is a fundamental characteristic of our species, foundational to many biology disciplines, and a major differentiator in medical/health outcomes.Public discourse around sex increasingly seeks to deny basic facts of human biology.One recent example has been the treatment of Suzanne Moore at The Guardian following her attempts to discuss sex-related issues (https://unherd.com/2020/11/why-i-had-to-leave-the-guardian/).This denialism is no longer confined to humanities departments and social media hashtags but has made inroads into mainstream culture, in part due to a highly sympathetic media environment.Of particular concern to us is the sight of respected scientific publications, such as Nature, now beginning to echo these popular trends.In a recent article discussing a research study of differential disease burden in male and female patients with cystic fibrosis, the following

Eusocial insect societies possess complex multilevel disease defences, including individual level protection conferred by physical (e.g. cuticle) and immunological obstacles and colony level protection mediated by collective behaviours (social immunity). It remains unclear whether and how these two levels of disease protection are related to one another in jointly driving colonies' susceptibility to disease. Here, we examine whether a relationship exists between individual worker survival after exposure to a fungal pathogen (a proxy for immunity) and corpse removal (a colony level social immunity metric) in the acorn ant Temnothorax curvispinosus. Since behavioural avoidance is the first line of defence against infection, we also tested whether individual ants exhibited parasite avoidance behaviour during exploration and whether colonies exhibit avoidance behaviour during foraging. We found that individual level and colony level immunity were negatively correlated: colonies that removed corpses more rapidly contained workers with weaker individual defences. We did not detect parasite avoidance behaviour by individual workers or whole colonies, nor were these two factors related. These data suggest that individual immunity and social immunity may trade off, regulating overall parasite protection. Alternatively, optimized social immunity at the colony level may compensate for disease vulnerability to infection at the individual level, and thus provide a protective benefit in overall colony defence in the absence of pathogen avoidance. (c) 2021 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Many animal societies are susceptible to mass mortality events and collapse. Elucidating how environmental pressures determine patterns of collapse is important for understanding how such societies function and evolve. Using the social spider Stegodyphus dumicola, we investigated the environmental drivers of colony extinction along two precipitation gradients across southern Africa, using the Namib and Kalahari deserts versus wetter savanna habitats to the north and east. We deployed experimental colonies (n = 242) along two ~ 800-km transects and returned to assess colony success in the field after 2 months. Specifically, we noted colony extinction events after the 2-month duration and collected environmental data on the correlates of those extinction events (e.g., evidence of ant attacks, no. of prey captured). We found that colony extinction events at desert sites were more frequently associated with attacks by predatory ants as compared with savanna sites, while colony extinctions in wetter savannas sites were more tightly associated with fungal outbreaks. Our findings support the hypothesis that environments vary in the selection pressures that they impose on social organisms, which may explain why different social phenotypes are often favored in each habitat.

The founders of a social group or colony have the potential to greatly influence the success or failure of the societies that they initiate. Whether through their genetic contribution, or through behavioral maternal effects related to temperament, resource acquisition, or brood care, the traits of founders deserve special consideration. The queens of many social insects are ideally suited to address questions relating to the importance of founder traits due to her solitary involvement in performing a multitude of tasks necessary to produce a functioning colony. While it has been suggested that a queen’s behavioral phenotype might contribute to differences in colony success, no study has yet demonstrated these links in situ. Here, we use the singly founding (haplometrotic) paper wasp Polistes metricus to examine whether queen personality, measured shortly after colony founding (pre-emergent phase), and morphology, can predict colony size (a proxy for queen fitness) and parasite load in the wild. We found that larger and bolder queens gave rise to larger colonies than smaller and shyer queens, and there was no relationship between queen personality and parasite load. Differences in queen traits therefore appear to be a major determinant of colony success under field conditions. In social species, the personality composition of groups has been shown to be a large determinant of group behavior, which in turn can determine group performance and survival. In this study, we explore the degree to which the behavioral tendencies of key group members—wasp queens—influence the performance of their descendant societies. We tested the behavioral tendencies and morphology of Polistes metricus paper wasp queens in the early spring, placed them back in the field in nest boxes, and recorded each nest’s cell count at the end of the season—a reliable proxy for queen fitness. We found that larger and bolder queens gave rise to larger descendant colonies on average than their smaller, shyer counterparts. Individual variation in queen traits therefore appears to be a determinant of colony success under field conditions.

Nest parasites attempt to shift the cost of rearing young from themselves to others. Despite strong selection to avoid this exploitation, there is considerable variation among-individuals in susceptibility to nest parasites. We evaluated the effects of individual variation in boldness, aggressiveness, and olfactory responsiveness on egg discrimination in wasps: Polistes metricus, which founds nests as singleton individuals, and P. dominula, which founds nests in small coalitions. Aggressiveness and boldness were evaluated using individuals’ response to mechanical disturbance, and olfactory responsiveness was evaluated using individuals’ tendency to respond to a novel rewarding stimulus. Egg discrimination was evaluated by presenting each queen with a variety of foreign eggs: 1) unaltered eggs laid by the resident (negative control group), 2) eggs produced by the resident female that were removed and replaced (procedural control), 3) eggs of foreign conspecifics (conspecific egg), and 4) eggs of a heterospecific congener (heterospecific egg). Females of both species never rejected untampered eggs and rejected procedural controls in only 35% of cases. Both species were twice as likely (70% rejection rate) to reject eggs of foreign conspecifics or heterospecifics. In P. dominula, bolder individuals and those with low olfactory responsiveness were more likely to reject foreign eggs. In P. metricus, boldness was not associated with egg rejection, but individuals with heightened olfactory responsiveness were more likely to reject foreign eggs. Thus, there are contrasting associations between behavioral phenotypes and egg rejection across species. These results are discussed in light of differences in the colony founding behavior of these species.

Collective personalities refer to temporally consistent behavioral differences between distinct social groups. This phenomenon is a ubiquitous and key feature of social groups in nature, as virtually every study conducted to date has documented repeatable between-group differences in collective behavior, and has revealed ongoing selection on these traits in both the laboratory and field environments. Five years ago, foundational reviews by Bengston and Jandt pioneered this topic and delimited the present knowledge on collective personality. Here, we update these reviews by summarizing the recent works conducted in the field's most prominent model systems: social spiders and eusocial insects. After presenting how these recent works helped scientists tobetter understand the determinants of collective personality, we used a trait-by-trait format to compare and contrast the results and thematic trends obtained in these taxaon 10 major aspects of collective personality: division of labor, foraging, exploration, boldness, defensive behavior, aggressiveness, decision-making, cognition, learning, and nest construction. We then discuss why similarities and dissimilarities in these results open the door to applying numerous theories developed in evolutionary behavioral ecology for individual traits (e.g., life history theory, game theory, optimal foraging theory) at the colony level, and close by providing examples of unexamined questions in this field that are ripe for new inquiries. We conclude that collective personality, as a framework, has the potential to improve our general understanding of how selection acts on intraspecific variation in collective phenotypes that are of key importance across arthropod societies and beyond.

Division of labor (DOL) is a pattern of work organization where individual group members specialize on different tasks. DOL is argued to have been instrumental for the success of eusocial insects, where it scales positively with group size both within and across species. Here we evaluate whether DOL scales positively with group size in a society of cooperative breeders (social spiders) and whether this pattern is impacted by the behavioral composition of the group. To do this we engineered experimental colonies of contrasting group sizes and behavioral compositions and tracked individuals participation in two colony maintenance tasks: prey capture and web construction. As with some eusocial insects, we found that larger groups exhibited DOL metrics up to 10-times greater than smaller groups, conveying that individuals specialize on particular tasks more in larger colonies. This scalar relationship did not differ by a groups behavioral composition, though groups composed of only bold spiders exhibited reduced DOL relative to all-shy or mixed groups. We also found that per capita participation in prey capture, but not web construction, decreased as a function of group size. This suggests that individuals in larger groups may save energy by reducing their involvement in some tasks. Together, our results convey that similar scalar relationships between DOL and group size can emerge both inside and outside the eusocial insects. Thus, theory developed for understanding DOL in eusocial societies may inform our understanding of group function in a larger swath of animal social diversity than is broadly appreciated.Significance Statement Division of labor (DOL) has been a major area of research in the eusocial insects for decades, and is argues to underlie their ecological success. Only recently have other social arthropods, such as social spiders, been considered for studies concerning DOL. Given their smaller colony sizes, and absence of morphological castes, DOL was not thought to be an important facet of spider societies. However, we found that spider societies do indeed exhibit high degrees of DOL that is positively correlated to colony size, as seen in many eusocial insects. These findings suggest that the scalar relationship between group size and social organization seen in social insects is likely generalizable to a larger diversity of social taxa, and that cooperative breeders can show levels of division of labor equaling or exceeding those of eusocial systems evaluated to date.

Stable between-group variation in collective behavior has been observed in a variety of taxa. We examine here whether climate of origin (arid/wet), colony personality composition (shy/bold/mixed), and group size determine the repeatability of collective foraging behavior in Stegodyphus dumicola Pocock, 1898 (Eresidae). Experimental colonies were created with contrasting ratios of bold/shy group members and run through 20 simulated prey capture events in a greenhouse. We found that (i) larger colonies and colonies composed of bold spiders were more repeatable in how many attackers they deployed to prey stimuli, (ii) colonies composed of shy spiders were more repeatable in their latency of attack, and (iii) climate of origin had no effect on the repeatability of colony behavior. Colony bold/shy composition had no effect on within-group variation in foraging behavior. Thus, differences in repeatability were the result of increases in between-group differences in foraging behavior, and not shifts in the behavioral flexibility of individual colonies. These results indicate that changes to colony composition and group size can alter the extent to which colonies exhibit characteristic behavioral differences.

The individual behavioral traits of predators and prey sometimes determine the outcome of their interactions. Here, we examine whether changes to habitat complexity alter the effects of predator and prey behavior on their survival rates. Specifically, we test whether behavioral traits (activity level, boldness, and perch height) measured in predators and prey or multivariate behavioral volumes best predict the survival rates of both trophic levels in staged mesocosms with contrasting structural complexity. Behavioral volumes and hypervolumes are a composite group-level behavioral diversity metric built from the individual-level behavioral traits we measured in predators and prey. We stocked mesocosms with a host plant and groups of cannibalistic predators (n = 5 mantises/mesocosm) and their prey (n = 15 katydids/mesocosm), and mesocosms varied in the presence/absence of additional non-living climbing structures. We found that mantis survival rates were unrelated to any behavioral metric considered here, but were higher in structurally complex mesocosms. Unexpectedly, katydids were more likely to survive when mantis groups occupied larger behavioral volumes, indicating that more behaviorally diverse predator groups are less lethal. Katydid mortality was also increased when both predators and prey exhibited higher average perch heights, but this effect was increased by the addition of supplemental structure. This is consistent with the expectation that structural complexity increases the effect of intraspecific behavioral variation on prey survival rates. Collectively, these results convey that the effects of predator and prey behavior on prey survival could depend highly on the environment in which they are evaluated.

A major benefit of living in a group is the ability to learn from others. We investigated how spider societies learn and respond to important information when that information is held by the majority or by single influential or generic individuals. We found that groups adopted a “better safe than sorry” strategy and exhibited caution when the group or any individual, regardless of their presumed social influence, had been previously exposed to danger.

Colonies of social insects exhibit a spectacular variety of life histories. Here we documented the degree of variation in colony life-history traits, mostly related to productivity, in two species of wild paper wasps. We then tested for associations between colony life-history traits to look for trade-offs or positively associated syndromes, and examined whether individual differences in the behavioral tendencies of foundresses ( Polistes metricus ) or the number of cofoundresses ( P. fuscatus ) influenced colony life-history. The majority of our measures of colony life-history were positively related, indicating no obvious resource allocation trade-offs. Instead, the positive association of traits into a productivity syndrome appears to be driven by differences in queen or microhabitat quality. Syndrome structure differed only marginally between species. Queen boldness and body size were not associated with colony life-history in P. metricus . Colonies initiated by multiple P. fuscatus foundresses were generally more productive, and this advantage was approximately proportional to the number of cofoundresses. These findings demonstrate that colony life-history traits can be associated together much like individual life-history traits, and the associations seen here convey that differences in overall productivity drive between-colony differences in life-history.

Social spiders are thought to predominantly receive information about their environment through vibrational cues. Thus, group living introduces the challenge of distinguishing useful vibrational information from the background noise of nestmates. Here we investigate whether spatial proximity between colony-mates may allow social spiders (Stegodyphus dumicola) to reduce background noise that might obstruct vibrational information from prey. To do so, we constructed experimental colonies and measured whether the number of spiders in proximity to one another whilst resting could predict the number of spiders that participated in prey capture. Additionally, we exposed spider colonies to five different simulated vibrational cues mimicking prey to determine which cue types spiders were most responsive to. We found that the number of spiders huddled together prior to foraging trials was positively correlated with the number of spiders participating in collective foraging. Furthermore, colonies responded more quickly to pulsed vibrational cues over other types of vibrational patterns. Together these data reveal that both social interactions and prey cues shape how social sit-and-wait predators experience and respond to their environment.

Particularly socially influential individuals are present in many groups [1-8], but it is unclear whether their emergence is determined by their social influence versus the social susceptibility of others [9]. The social spider Stegodyphus dumicola shows regional variation in apparent leader-follower dynamics. We use this variation to evaluate the relative contributions of leader social influence versus follower social susceptibility in driving this social order. Using chimeric colonies that combine potential leaders and followers, we discover that leader-follower dynamics emerge from the site-specific social susceptibility of followers. We further show that the presence of leaders increases colony survival in environments where leader-follower dynamics occur. Thus, leadership is driven by the "social susceptibility" of the population majority, rather than the social influence of key group members.

Cooperative breeding decreases the direct reproductive output of subordinate individuals, but cooperation can be evolutionarily favored when there are challenges or constraints to breeding independently. Environmental factors, including temperature, precipitation, latitude, high seasonality, and environmental harshness have been hypothesized to correlate with the presence of cooperative breeding. However, to test the relationship between cooperation and ecological constraints requires comparative data on the frequency and variation of cooperative breeding across differing environments, ideally replicated across multiple species. Paper wasps are primitively social species, forming colonies composed of reproductively active dominants and foraging subordinates. Adult female wasps, referred to as foundresses, initiate new colonies. Nests can be formed by a single solitary foundress (noncooperative) or by multiple foundress associations (cooperative). Cooperative behavior varies within and among species, making paper wasps species well suited to disentangling ecological correlates of variation in cooperative behavior. This data set reports the frequency and extent of cooperative nest founding for 87 paper wasp species. Data were assembled from more than 170 published sources, previously unpublished field observations, and photographs contributed by citizen scientists to online natural history repositories. The data set includes 25,872 nest observations and reports the cooperative behavioral decisions for 45,297 foundresses. Species names were updated to reflect modern taxonomic revisions. The type of substrate on which the nest was built is also included, when available. A smaller population-level version of this data set found that the presence or absence of cooperative nesting in paper wasps was correlated with temperature stability and environmental harshness, but these variables did not predict the extent of cooperation within species. This expanded data set contains details about individual nests and further increases the power to address the relationship between the environment and the presence and extent of cooperative breeding. Beyond the ecological drivers of cooperation, these high-resolution data will be useful for future studies examining the evolutionary consequences of variation in social behavior. This data set may be used for research or educational purposes provided that this data paper is cited.

The field of animal behavior has experienced a surge of studies focusing on functional differences among individuals in their behavioral tendencies (‘animal personalities’) and the relationships between different axes of behavioral variation (‘behavioral syndromes’). Many important developments in this field have arisen through research using insects and other terrestrial arthropods, in part, because they present the opportunity to test hypotheses not accessible in other taxa. This chapter reviews how studies on insects and spiders have advanced the study of animal personalities by describing the mechanisms underlying the emergence of individual variation and their ecological consequences. Furthermore, studies accounting for animal personalities can expand our understanding of phenomena in insect science like metamorphosis, eusociality, and applied insect behavior. In addition, this chapter serves to highlight some of the most exciting issues at the forefront of our field and to inspire entomologists and behaviorists alike to seek the answers to these questions.

Consistent differences in behavior between individuals, otherwise known as animal personalities, have become a staple in behavioral ecology due to their ability to explain a wide range of phenomena. Social organisms are especially serviceable to animal personality techniques because they can be used to explore behavioral variation at both the individual and group level. Despite the success of personality research in social organisms generally, and social Hymenoptera in particular, social wasps (Vespidae) have received little to no attention in the personality literature. In the present study, we test Polistes metricus (Vespidae; Polistinae) paper wasp queens for the presence of repeatable variation in, and correlations (“behavioral syndromes”) between, several commonly used personality metrics: boldness, aggressiveness, exploration, and activity. Our results indicate that P. metricus queens exhibit personalities for all measured traits and correlations between different behavioral measures. Given that paper wasps have served as a model organism for a wide range of phenomena such as kin selection, dominance hierarchies, mate choice, facial recognition, social parasitism, and chemical recognition, we hope that our results will motivate researchers to explore whether, or to what degree, queen personality is important in their research programs.