Jennifer M. Jandt

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).

Spatial organization and division of labour in the bumblebee Bombus impatiens

Individuals in many types of animal groups show both reproductive and task-related division of labour. In some social insect species, such division of labour may be related to the spatial organization of workers inside the nest. We examined colonies of bumblebees and found that (1) 11–13% of workers maintained small spatial fidelity zones inside the nest, and all workers tended to remain at a specific distance from the colony centre independent of their age; (2) smaller individuals maintained smaller spatial zones and tended to be closer to the centre; and (3) individuals that were more likely to perform the in-nest task of larval feeding tended to remain in the centre of the nest, whereas foragers were more often found on the periphery of the nest when not foraging. Individuals that performed other tasks did not maintain a predictable distance to the centre, and there was no evidence that spatial preferences changed over time. Instead, spatial patterns may result from inherent differences between individuals in terms of activity level, and may be a self-organized sorting mechanism that influences division of labour among workers.

Polistes paper wasps: a model genus for the study of social dominance hierarchies

Polistes are an ideal system to study ultimate and proximate questions of dominance, and to test theoretical predictions about social evolution. The behaviors typically associated with dominance in Polistes are similar to those observed in many vertebrate societies. Here, we review recent ethological, mechanistic, and evolutionary studies on how social dominance hierarchies are established and maintained in Polistes spp. From the ultimate perspective, we address individual and group benefits of hierarchy formation, as well as issues such as reproductive skew, queen-worker conflict, and costs of challenging the dominant. From the proximate perspective, we review social, physical, and physiological factors influencing hierarchy formation, including co-foundress interactions, age structure, body size, endocrine system, and chemical and visual signals. We also discuss the extensive inter- and intra-specific variation of Polistes in the formation and maintenance of hierarchies, as well as levels of within-colony aggression. We conclude the review by highlighting the utility of this variation for comparative studies and the immense potential of the genus Polistes to address fundamental and unanswered questions about the evolution and maintenance of dominance behavior in animals.

Weak specialization of workers inside a bumble bee ( Bombus impatiens ) nest

Division of labor is common across social groups. In social insects, many studies focus on the differentiation of in-nest and foraging workers and/or the division of foraging tasks. Few studies have specifically examined how workers divide in-nest tasks. In the bumble bee, Bombus impatiens, we have shown previously that smaller workers are more likely to feed larvae and incubate brood, whereas larger workers are more likely to fan or guard the nest. Here, we show that in spite of this, B. impatiens workers generally perform multiple tasks throughout their life. The size of this task repertoire size does not depend on body size, nor does it change with age. Further, individuals were more likely to perform the task they had been performing on the previous day than any other task, a pattern most pronounced among individuals who guarded the nest. On the other hand, there was no predictable sequence of task switching. Because workers tend to remain in the same region of the nest over time, in-nest workers may concentrate on a particular task, or subset of tasks, inside that region. This division of space, then, may be an important mechanism that leads to this weak specialization among in-nest bumble bee workers.

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.

Ontogeny of worker body size distribution in bumble bee (Bombus impatiens) colonies

1. Bumble bees exhibit worker size polymorphisms; highly related workers within a colony may vary up to 10‐fold in body mass. As size variation is an important life history feature in bumble bees, the distribution of body sizes within the colony and how it fluctuates over the colony cycle were analysed.

Bumblebee response thresholds and body size: does worker diversity increase colony performance?

Although models of colony organization in social insects often rely on the assumption that within-group variation increases group performance, empirical support for this is mostly confined to studies of genetic variation. However, workers in ant or bee colonies often vary in behaviour and morphology even when genetic variation is low. Bumblebees provide a unique opportunity to explore the consequences of such variation: colonies have a wide range of worker body sizes compared to other social bee species, and workers also vary in response thresholds (i.e. stimulus levels at which workers respond by performing a task), in spite of queens being singly mated (and thus, low genetic variation). Here we test how body size and response threshold diversity affect colony performance in two unrelated in-nest tasks (thermoregulation and undertaking). We manipulated worker diversity using worker removals to restrict threshold or body size variation within the colony. We also quantified the degree of intracolony variation across colonies and related this to colony performance. In general, colonies took longer to cool the nest after bees were removed, but there was no significant effect of treatment on fanning or undertaking success. Furthermore, when intracolony variation was analysed as a continuous variable, we found no effect on colony-level thermoregulation or undertaking performance. Instead, average threshold was a more useful predictor of thermoregulation success, and colonies with a narrower range of size variation had more success at undertaking. These results emphasize the importance of understanding how different types of variation (e.g. behavioural, morphological, etc.) contribute to colony performance.

Competition and cooperation: bumblebee spatial organization and division of labor may affect worker reproduction late in life

Within-group conflict may influence the degree to which individuals within a group cooperate. For example, the most dominant individuals within a group often gain access to the best resources and may be less inclined to perform risky tasks. We monitored space use and division of labor among all workers in three colonies of bumblebees, Bombus impatiens, during the ergonomic and queenless phases of their colony cycle. We then measured the two largest oocytes in each worker to estimate each individuals reproductive potential at the end of the colony cycle. We show that workers that remained farther from the queen while inside the nest and avoided risky or more energy-expensive tasks during the ergonomic phase developed larger oocytes by the end of the colony cycle. These individuals also tended to be the largest, oldest workers. After the queen died, these workers were more likely than their nestmates to increase brood incubation. Our results suggest that inactive bumblebees may be storing fat reserves to later develop reproductive organs and that the spatial organization of workers inside the nest, particularly the distance workers maintain from the queen, may predict which individuals will later have the greatest reproductive potential in the colony.

Different axes of environmental variation explain the presence vs. extent of cooperative nest founding associations in Polistes paper wasps

Ecological constraints on independent breeding are recognised as major drivers of cooperative breeding across diverse lineages. How the prevalence and degree of cooperative breeding relates to ecological variation remains unresolved. Using a large data set of cooperative nesting in Polistes wasps we demonstrate that different aspects of cooperative breeding are likely to be driven by different aspects of climate. Whether or not a species forms cooperative groups is associated with greater short-term temperature fluctuations. In contrast, the number of cooperative foundresses increases in more benign environments with warmer, wetter conditions. The same data set reveals that intraspecific responses to climate variation do not mirror genus-wide trends and instead are highly heterogeneous among species. Collectively these data suggest that the ecological drivers that lead to the origin or loss of cooperation are different from those that influence the extent of its expression within populations.

Individual bumblebees vary in response to disturbance: a test of the defensive reserve hypothesis

Bees may leave their nest in the event of an attack, but this is not their only response. Here, we examine the behavior of those individuals that remain inside the nest during a disturbance. Specifically, we test the hypothesis that bee workers usually exhibiting high levels of inactivity (i.e., ‘lazy’ bees) may function as defensive reserves that are more likely to respond when the colony is disturbed. We explore this hypothesis by simulating vertebrate attacks by vibrating or blowing carbon dioxide into two colonies on alternating days and measuring the movements and tasks performed by bees inside the nest. Our results show that regardless of the disturbance type, workers increase guarding behavior after a disturbance stops. Although previously inactive bees increased their movement speed inside the nest when the disturbance was vibration, they were not more likely to leave the nest (presumably to attack the simulated attacker) or switch to guarding behavior for any disturbance type. We therefore reject the hypothesis that inactive Bombus impatiens bumblebees act as defensive reserves, and propose alternative hypotheses regarding why many workers remain inactive inside the nest.