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Emergence is a fundamental concept in biology and other disciplines, but whether emergent phenotypes evolve similarly to nonemergent phenotypes is unclear. The hypothesized process of emergent evolution posits that evolutionary change in at least some collective behaviors will differ from evolutionary change in the corresponding intrinsic behaviors of isolated individuals. As a result, collective behavior might evolve more rapidly and diversify more between populations compared to individual behavior. To test whether collective behavior evolves emergently, we conducted a large comparative study using 22 ant species and gathered over 1,500 behavioral rhythm time series from hundreds of colonies and isolated individuals, totaling over 1.5 y of behavioral data. We show that analogous traits measured at individual and collective levels exhibit distinct evolutionary patterns. The estimated rates of phenotypic evolution for the rhythmicity of activity in ant colonies were faster than the evolutionary rates of the same behavior measured in isolated individual ants, and total variation across species in collective behavior was higher than variation in individual behavior. We hypothesize that more rapid evolution and higher variation is a widespread feature of emergent phenotypes relative to lower-level phenotypes across complex biological systems., Competing Interests: Competing interests statement:The authors declare no competing interest.

Synchronization is a conspicuous form of collective behaviour that is of crucial importance in numerous biological systems. Ant colonies from the genera Leptothorax and Temnothorax form small colonies, typically made up of only a few hundred workers, and exhibit a form of synchronized behaviour where workers inside colonies' nests become active together in rhythmic cycles that have a period of approximately 20-200 min. However, it is not currently known if these synchronized rhythms of locomotion confer any functional benefit to colonies. By using a combination of multiple image analysis techniques, we show that inactive Leptothorax ants can act as immobile obstacles to moving ants, and that synchronized activity has the potential to reduce the likelihood that individual ants will encounter regions of immobile obstacles that impede access to portions of the nest. We demonstrate qualitatively similar findings using a computational model of confined active particles with oscillating activity.

Evolutionary theory predicts that animals make decisions that maximize fitness. If so, they are expected to adhere to principles of rational choice, which a decision-maker must follow to reliably maximize net benefit. For example, evaluation of an option should not be influenced by the quality of other unchosen options. However, humans and other animals are known to evaluate a mediocre option more favorably after encountering poor options than after encountering no options, a phenomenon known as the 'anchoring effect'. Rationality is also expected in the consensus decisions of animal societies, but the anchoring effect has not previously been tested in that context. Here we show that colonies of the rock ant, Temnothorax rugatulus, demonstrate the anchoring effect during nest site selection - colonies moved more readily from a mediocre nest to a good nest when exposed to poor nests than when exposed to mediocre nests. This effect depended on both current conditions and past experience; movement probability was affected only when colonies were exposed to surrounding nests before and during the emigration. The effect was small, reaching statistical significance in only one of two experimental replicates. We discuss possible mechanisms and ultimate explanations for why colonies show this seemingly suboptimal behavior., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Biology is suffused with rhythmic behaviour, and interacting biological oscillators often synchronize their rhythms with one another. Colonies of some ant species are able to synchronize their activity to fall into coherent bursts, but models of this phenomenon have neglected the potential effects of intrinsic noise and interspecific differences in individual-level behaviour. We investigated the individual and collective activity patterns of two Leptothorax ant species. We show that in one species ( Leptothorax sp. W), ants converge onto rhythmic cycles of synchronized collective activity with a period of about 20 min. A second species ( Leptothorax crassipilis ) exhibits more complex collective dynamics, where dominant collective cycle periods range from 16 min to 2.8 h. Recordings that last 35 h reveal that, in both species, the same colony can exhibit multiple oscillation frequencies. We observe that workers of both species can be stimulated by nest-mates to become active after a refractory resting period, but the durations of refractory periods differ between the species and can be highly variable. We model the emergence of synchronized rhythms using an agent-based model informed by our empirical data. This simple model successfully generates synchronized group oscillations despite the addition of noise to ants' refractory periods. We also find that adding noise reduces the likelihood that the model will spontaneously switch between distinct collective cycle frequencies., Competing Interests: We declare we have no competing interests., (© 2022 The Authors.)

Group cohesion and collective decision-making are important for many social animals, like social insects, whose societies depend on the coordinated action of individuals to complete collective tasks. A useful model for understanding collective, consensus-driven decision-making is the fluid nest selection dynamics of ant colonies. Certain ant species oscillate between occupying multiple nests simultaneously (polydomy) and reuniting at a single location (monodomy), but little is known about how colonies achieve a consensus around these dynamics. To investigate the factors underpinning the splitting-reunification dynamics of ants, we manipulated the availability and quality of nest sites for the ant Temnothorax rugatulus and measured the likelihood and speed of reunification from contrasting starting conditions. We found that pursuing reunification was more likely for smaller colonies, that rates of initial splitting were lower when colonies could coordinate their activity from a central hub, and that diluting colonies among additional sites did not impair reaching consensus on a single nest. We further found mixed support for a specific threshold of social density that prevents reunification (i.e., prolonged polydomy) and no evidence that nest quality influences reunification behavior. Together our data reveal that consensus driven decisions can be influenced by both external and intrinsic group-level factors and are in no way simple stereotyped processes., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Populations of independently oscillating agents can sometimes synchronize. In the context of animal societies, conspicuous synchronization of activity is known in some social insects. However, the causes of variation in synchrony within and between species have received little attention. We repeatedly assessed the short-term activity cycle of ant colonies ( Temnothorax rugatulus ) and monitored the movements of individual workers and queens within nests. We detected persistent differences between colonies in the waveform properties of their collective activity oscillations, with some colonies consistently oscillating much more erratically than others. We further demonstrate that colony crowding reduces the rhythmicity (i.e., the consistent timing) of oscillations. Workers in both erratic and rhythmic colonies spend less time active than completely isolated workers, but workers in erratic colonies oscillate out of phase with one another. We further show that the queen's absence can impair the ability of colonies to synchronize worker activity and that behavioral differences between queens are linked with the waveform properties of their societies., (© The Author(s) 2019. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

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.

Maintenance of a social group requires the ability to reach consensus when faced with divisive choices. Thus, when migrating colonies of the ant Temnothorax rugatulus split among multiple sites, they can later reunify on the basis of queen location or differences in site quality. In this study, we found that colonies can reunify even without obvious cues to break the symmetry between sites. To learn how they do so, we observed both symmetric reunifications (between identical nests) and asymmetric reunifications (between nests of unequal quality) by colonies of individually marked ants. Both reunification types were accomplished by a tiny minority that carried nestmates from the 'losing' to the 'winning' site. Reunification effort was highly skewed in asymmetric splits, where the majority of the work was done by the first ant to transport, which nearly always came from the winning site. This contrasted with symmetric splits, where the initiator did not play an outsize role and was just as likely to come from the losing site. Symmetric reunifications were also characterized by high transporter attrition, which may help to prevent deadlocks. Tandem runs were abundant in both types and were typically led by transporters as they returned to the losing site to fetch another nestmate. Few tandem followers joined the transport effort, suggesting that tandem runs do not serve to recruit transporters but may have another, as yet unidentified role. Our results underscore the potentially large contribution of highly active individuals to group behaviour, even in decentralized societies such as ant colonies., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

The physical environment occupied by group-living animals can profoundly affect their cooperative social interactions and therefore their collective behavior and success. These effects can be especially apparent in human-modified habitats, which often harbor substantial variation in the physical environments available within them. For nest-building animal societies, this influence of the physical environment on collective behavior can be mediated by the construction of nests-nests could either buffer animal behavior from changes in the physical environment or facilitate shifts in behavior through changes in nest structure. We test these alternative hypotheses by examining the differences in collective prey-attacking behavior and colony persistence between fence-dwelling and tree-dwelling colonies of Stegodyphus dumicola social spiders. Fences and trees represent substantially different physical environments: fences are 2-dimensional and relatively homogenous environments, whereas tree branches are 3-dimensional and relatively heterogeneous. We found that fence-dwelling colonies attack prey more quickly and with more attackers than tree-dwelling colonies in both field and controlled settings. Moreover, in the field, fence-dwelling colonies captured more prey, were more likely to persist, and had a greater number of individuals remaining at the end of the experiment than tree-dwelling colonies. Intriguingly, we also observed a greater propensity for colony fragmentation in tree-dwelling colonies than fence-dwelling colonies. Our results demonstrate that the physical environment is an important influence on the collective behavior and persistence of colonies of social spiders, and suggest multiple possible proximate and ultimate mechanisms-including variation in web complexity, dispersal behavior, and bet-hedging-by which this influence may be realized.

Animal social groups are complex systems that are likely to exhibit tipping points-which are defined as drastic shifts in the dynamics of systems that arise from small changes in environmental conditions-yet this concept has not been carefully applied to these systems. Here, we summarize the concepts behind tipping points and describe instances in which they are likely to occur in animal societies. We also offer ways in which the study of social tipping points can open up new lines of inquiry in behavioural ecology and generate novel questions, methods, and approaches in animal behaviour and other fields, including community and ecosystem ecology. While some behaviours of living systems are hard to predict, we argue that probing tipping points across animal societies and across tiers of biological organization-populations, communities, ecosystems-may help to reveal principles that transcend traditional disciplinary boundaries., (© 2018 The Authors.)

Living systems sometimes experience abrupt tipping points in response to stress. Here we investigate the factors contributing to the appearance of such abrupt state transitions in animal societies. We first construct a mathematical account of how the personality compositions of societies could alter their propensity to shift from calm to violent states in response to thermal stress. To evaluate our model, we subjected experimental societies of the spider Anelosimus studiosus to heat stress. We demonstrate that both colony size and personality composition influence the timing of and recoverability from sudden transitions in social state. Groups composed of aggressive personalities transitioned into violent within-group dynamics sooner during heating, and also resisted recovery to baseline non-aggressive behaviour during cooling. We further observed hysteresis in groups composed of aggressive individuals, where group behaviour depended strongly on whether the colony had previously been in a calm or agitated state. These results demonstrate that a society's susceptibility to sudden state shifts and their recoverability from them can be driven by the personalities of their constituents.

This study showed that there is a relationship between acid phosphatase levels and life cycle stages in the nematode, Panagrellus silusiae. Ten different isozymes of acid phosphatase were separated electrophoretically. Relative activity peaked at different stages in the life cycle for the different isozymes. Later in the life cycle, there is a general decrease in the relative activity of acid phosphatase itself, while there is a concurrent increase in the number of isozymes present. At least 97% of the acid phosphatase in P. silusiac is soluble (unbound). Acid phosphatase appears to be present in large quantities in the entire gastrointestinal tract, the excretory canals, and the reproductive system of mature Panagrellus silusiae.