Sarah E. Bengston

Behavioural syndromes and social insects: personality at multiple levels.

Animal personalities or behavioural syndromes are consistent and/or correlated behaviours across two or more situations within a population. Social insect biologists have measured consistent individual variation in behaviour within and across colonies for decades. The goal of this review is to illustrate the ways in which both the study of social insects and of behavioural syndromes has overlapped, and to highlight ways in which both fields can move forward through the synergy of knowledge from each. Here we, (i) review work to date on behavioural syndromes (though not always referred to as such) in social insects, and discuss mechanisms and fitness effects of maintaining individual behavioural variation within and between colonies; (ii) summarise approaches and principles from studies of behavioural syndromes, such as trade‐offs, feedback, and statistical methods developed specifically to study behavioural consistencies and correlations, and discuss how they might be applied specifically to the study of social insects; (iii) discuss how the study of social insects can enhance our understanding of behavioural syndromes—research in behavioural syndromes is beginning to explore the role of sociality in maintaining or developing behavioural types, and work on social insects can provide new insights in this area; and (iv) suggest future directions for study, with an emphasis on examining behavioural types at multiple levels of organisation (genes, individuals, colonies, or groups of individuals).

Group Size and Its Effects on Collective Organization

Many insects and arthropods live in colonies or aggregations of varying size. Group size may affect collective organization either because the same individual behavior has different consequences when displayed in a larger group or because larger groups are subject to different constraints and selection pressures than smaller groups. In eusocial colonies, group size may have similar effects on colony traits as body size has on organismal traits. Social insects may, therefore, be useful to test theories about general principles of scaling, as they constitute a distinct level of organization. However, there is a surprising lack of data on group sizes in social insects and other group-living arthropods, and multiple confounding factors have to be controlled to detect effects of group size. If such rigorous studies are performed, group size may become as important to understanding collective organization as is body size in explaining behavior and life history of individual organisms.

Be meek or be bold? A colony-level behavioural syndrome in ants

Consistent individual variation in animal behaviour is nearly ubiquitous and has important ecological and evolutionary implications. Additionally, suites of behavioural traits are often correlated, forming behavioural syndromes in both humans and other species. Such syndromes are often described by testing for variation in traits across commonly described dimensions (e.g. aggression and neophobia), independent of whether this variation is ecologically relevant to the focal species. Here, we use a variety of ecologically relevant behavioural traits to test for a colony-level behavioural syndrome in rock ants (Temnothorax rugatulus). Specifically, we combine field and laboratory assays to measure foraging effort, how colonies respond to different types of resources, activity level, response to threat and aggression level. We find evidence for a colony level syndrome that suggests colonies consistently differ in coping style—some are more risk-prone, whereas others are more risk-averse. Additionally, by collecting data across the North American range of this species, we show that environmental variation may affect how different populations maintain consistent variation in colony behaviour.

The development of collective personality: the ontogenetic drivers of behavioral variation across groups

For the past decade, the study of personality has become a topic on the frontier of behavioral ecology. However, most studies have focused on exploring inter-individual behavioral variation in solitary animals, and few account for the role that social interactions may have on the development of an individual’s personality. Moreover, a social group may exhibit collective personality: an emergent behavioral phenotype displayed at the group-level, which is not necessarily the sum or average of individual personalities within that group. The social environment, in many cases, can determine group success, which then influences the relative success of all the individuals in that group. In addition, group-level personality may itself evolve, subject to the same selection pressures as individual-level behavioral variation, when the group is a unit under selection. Therefore, we reason that understanding how collective personalities emerge and change over time will be imperative to understanding individual- and group-level behavioral evolution. Personality is considered to be fixed over an individual’s lifetime. However, behavior may shift throughout development, particularly during adolescence. Therefore, juvenile behavior should not be compared with adult behavior when assessing personality. Similarly, as conditions within a group and/or the local environment can shift, group behavior may similarly fluctuate as it matures. We discuss potential within-group factors, such as group initiation, group maturation, genetic make-up of the group, and the internal social environment, and external factors, such as well as how local environment may play a role in generating group-level personalities. There are a variety of studies that explore group development or quantify group personality, but few that integrate both processes. Therefore, we conclude by discussing potential ways to evaluate development of collective personality, and propose several focal areas for future research.

Differences in environmental enrichment generate contrasting behavioural syndromes in a basal spider lineage

Behavioural syndrome studies are commonly descriptive and often find a relationship between boldness, shyness and exploration. However, the mechanisms underlying behavioural syndromes are not well understood. In the present study, we examined the extent to which early experience acts as a modifier of behavioural tendencies in the basal tarantula, Brachypelma smithi. Juvenile individuals were housed for 2 years either in enriched controlled conditions, or in restricted (minimal) conditions. Behavioural assays were completed both in short-term and in long-term increments. We found both short-term and long-term differences in multiple contexts of behaviour between treatments. In addition, individuals in the enriched treatment developed correlations between several behavioural traits whereas individuals in the restricted treatment did not. This result suggests that early environment can induce behavioural syndromes in some populations, or conversely, that continual stress may break down normal behavioural development and thus prevent a behavioural syndrome from emerging. This study provides a cautionary tale for those studying behavioural syndromes in captivity, and because this is a basal spider species, it provides important insight into the evolution of spider behavioural syndromes.

Colony size does not predict foraging distance in the ant Temnothorax rugatulus: a puzzle for standard scaling models

Body size is often positively correlated with ecologically relevant traits such as fecundity, survival, resource requirements, and home range size. Ant colonies, in some respects, behave like organisms, and their colony size is thought to be a significant predictor of many behavioral and ecological traits similar to body size in unitary organisms. In this study, we test the relationship between colony size and field foraging distance in the ant species Temnothorax rugatulus. These ants forage in the leaf litter presumably for small arthropod prey. We found colonies did not differ significantly in their foraging distances, and colony size is not a significant predictor of foraging distance. This suggests that large colonies may not exhaust local resources or that foraging trips are not optimized for minimal distance, and thus that food may not be the limiting resource in this species. This study shows T. rugatulus are behaving in ways that differ from existing models of scaling.

Latitudinal variation in behaviors linked to risk tolerance is driven by nest-site competition and spatial distribution in the ant Temnothorax rugatulus

Geographic range has long been noted to be associated with many organismic and ecological traits such as body size and species richness. However, much less is known about whether and how ecological variation across latitudinal gradients reflects behavioral variation. Ant colonies may also show behavioral variation, and Temnothorax rugatulus show a colony-level behavioral syndrome that seems to reflect risk tolerance across their North American range. While it is presumed that this pattern is the result of adaptation to local environmental conditions, which ecological factors are driving this variation are unknown. Here, we test if colony risk tolerance is affected by competition, predation, resource availability, or environmental stress at each site. Our results show that increased competition, specifically for nest sites, as well as increased spatial clustering of colonies predicts higher risk tolerance. Additionally, the spatial clustering of colonies influences the structure of the risk-taking syndrome, i.e., which colony-level behaviors are correlated and how strongly. This emphasizes the need for understanding large-scale geographic variation in behavior, as it may explain how ecological factors drive the evolution and maintenance of intraspecific behavioral variation across populations.

Morphological differences between extranidal and intranidal workers in the ant Temnothorax rugatulus, but no effect of body size on foraging distance

Most ant genera are thought to have monomorphic workers, indicating perhaps a high degree of flexibility in task allocation, and the well-studied genus Temnothorax is an example of this. However, considerable size variation may exist between individuals. In addition, though workers can show flexible behavior, it has been shown that individuals may consistently differ in their task profiles. Here we test whether body size variation among workers affects foraging behavior. Two main hypotheses were tested: first, whether larger ants forage at greater distance from the nest, and second, whether larger individuals show a higher propensity to work outside of the nest. Our results showed that ant body size does not significantly affect foraging distance. However, larger ants were more likely to be found outside the nest. Though Temnothorax ants are a common model system, this is the first study demonstrating task allocation based on body size, which is fixed in adults. Our study suggests that particularly small species may have to be examined carefully for body size variation before concluding that body size is uniform and therefore irrelevant for task allocation.

Activity Trends and Movement Distances in the Arizona Bark Scorpion (Scorpiones: Buthidae)

ABSTRACT The bark scorpion, Centruroides sculpturatus Ewing, is a nocturnal, cryptic, nonburrowing, mobile species that is common in urban landscapes spanning the desert southwest. Bark scorpions are often found in dense localized populations in cities, but the question of whether this is because the species is metabolically movement limited or choose to aggregate has not been addressed. Field observations lead us to believe that the scorpions move very little. Their ability to move is tested here. A circular pacing ring was constructed to observe the distance individuals could move in 2 h under both dark and light conditions. Observations under light motivate the arthropods to move, and significantly greater distances were observed in light trials, the maximum travel distance being 104.37 m, while the maximum distance in dark trials was 14.63 m. To monitor movement in the field, telemetry tags were used to mark female and male scorpions over 21 d during which relocation distances were recorded daily. Additionally, 12-h and 6-h overnight observational periods took place during which, scorpion movements were recorded hourly. Overall, it was found that scorpions moved significantly more in the pacing ring than in the field, indicating that field individuals are not moving at their maximum potential. Movement limitation does not explain their distribution pattern. In both the pacing ring and field, gender and pregnancy status had significant influence on distances moved. We conclude that C. sculpturatus is capable of much greater movement than is typically observed in the field.

Exploration of Refuge Preference in the Arizona Bark Scorpion (Scorpiones: Buthidae)

ABSTRACT The ongoing difficulty in understanding how Centruroides sculpturatus (Ewing) uses the built environment has prompted this study in exploring refuge choices in a school environment and in laboratory behavioral assays. Radio telemetry tags were used at an urban site heavily populated with C. sculpturatus to track scorpions for a period of 21 d from 1 to 21 August 2013. Complimentary laboratory work tested scorpions in refuge choice arenas targeting variables of four crevice widths—14.0, 9.3, 7.0, and 4.6 mm—or of a vertical versus horizontal orientation preference. These crevice sizes were picked as a response to crevices being naturally used in the field. Telemetry and observations tallied significant preference for artificial structural harborage. Ninety-five percent of the structure refuge use occurred in hollow block walls. Vegetative harborage, debris, and underground burrows were not selected with any significance compared with each other or structures. Generalized additive models (GAMs) indicated the strongest predictive power from individual preference. The behavioral choice assays yielded a significant preference for the largest of crevice widths offered, 14.0 mm and to a lesser extent 9.3 mm, both horizontally and vertically. GAMs for these assays indicated size as the strongest predictive factor in choices. The orientation tests and GAMs showed individual preference driving choice favoring vertical planes. Observations about negative geotaxis in assay and refuge use details from the field are also reported.