How within-group behavioural variation and task efficiency enhance fitness in a social group

How task specialization, individual task performance and within-group behavioural variation affects fitness is a longstanding and unresolved problem in our understanding of animal societies. In the temperate social spider, Anelosimus studiosus, colony members exhibit a behavioural polymorphism; females either exhibit an aggressive 'asocial' or docile 'social' phenotype. We assessed individual prey-capture success for both phenotypes, and the role of phenotypic composition on group-level prey-capture success for three prey size classes. We then estimated the effect of group phenotypic composition on fitness in a common garden, as inferred from individual egg-case masses. On average, asocial females were more successful than social females at capturing large prey, and colony-level prey-capture success was positively associated with the frequency of the asocial phenotype. Asocial colony members were also more likely to engage in prey-capture behaviour in group-foraging situations. Interestingly, our fitness estimates indicate females of both phenotypes experience increased fitness when occupying colonies containing unlike individuals. These results imply a reciprocal fitness benefit of within-colony behavioural variation, and perhaps division of labour in a spider society.     

Sources of individual variation in plasma testosterone levels

The steroid hormone testosterone (T) plays a central role in the regulation of breeding in males, because many physiological, morphological and behavioural traits related to reproduction are T dependent. Moreover, in many seasonally breeding vertebrates, male plasma T levels typically show a pronounced peak during the breeding season. While such population-level patterns are fairly well worked out, the sources and the implications of the large variability in individual T levels within the seasonal cycle remain surprisingly little understood. Understanding the potential sources of individual variation in T levels is important for behavioural and evolutionary ecologists, for at least two reasons. First, in 'honest signalling' theory, T is hypothesized to play a critical role as the assumed factor that enforces honesty of the expression of sexually selected quality indicators. Second, T is often considered a key mediator of central life-history trade-offs, such as investment in survival versus reproduction or in mating versus parental care. Here, we discuss the patterns of within- and between-individual variation in male plasma T levels in free-living populations of birds. We argue that it is unclear whether this variability mainly reflects differences in underlying individual quality (intrinsic factors such as genetic or maternal effects) or in the environment (extrinsic factors including time of day, individual territorial status and past experience). Research in avian behavioural endocrinology has mainly focused on the effects of extrinsic factors, while other sources of variance are often ignored. We suggest that studies that use an integrative approach and investigate the relative importance of all potential sources of variation are essential for the interpretation of data on individual plasma T levels.     

Parasitism and the evolutionary ecology of animal personality

The ecological factors responsible for the evolution of individual differences in animal personality (consistent individual differences in the same behaviour across time and contexts) are currently the subject of intense debate. A limited number of ecological factors have been investigated to date, with most attention focusing on the roles of resource competition and predation. We suggest here that parasitism may play a potentially important, but largely overlooked, role in the evolution of animal personalities. We identify two major routes by which parasites might influence the evolution of animal personality. First, because the risk of acquiring parasites can be influenced by an individual's behavioural type, local parasite regimes may impose selection on personality traits and behavioural syndromes (correlations between personality traits). Second, because parasite infections have consequences for aspects of host 'state', parasites might induce the evolution of individual differences in certain types of host behaviour in populations with endemic infections. Also, because infection often leads to specific changes in axes of personality, parasite infections have the potential to decouple behavioural syndromes. Host-parasite systems therefore provide researchers with valuable tools to study personality variation and behavioural syndromes from a proximate and ultimate perspective.     

Predation mediated population divergence in complex behaviour of nine‐spined stickleback (Pungitius pungitius)

The proximate and ultimate explanations for behavioural syndromes (correlated behaviours – a population trait) are poorly understood, and the evolution of behavioural types (configuration of behaviours – an individual trait) has been rarely studied. We investigated population divergence in behavioural syndromes and types using individually reared, completely predator‐ or conspecific‐naïve adult nine‐spined sticklebacks (Pungitius pungitius) from two marine and two predatory fish free, isolated pond populations. We found little evidence for the existence of behavioural syndromes, but population divergence in behavioural types was profound: individuals from ponds were quicker in feeding, bolder and more aggressive than individuals from marine environments. Our data reject the hypothesis that behavioural syndromes exist as a result of genetic correlations between behavioural traits, and support the contention that different behavioural types can be predominant in populations differing in predation pressure, most probably as a result of repeated independent evolution of separate behavioural traits.     

Consistent individual differences in cooperative behaviour in meerkats (Suricata suricatta)

Although recent models for the evolution of personality, using game theory and life‐history theory, predict that individuals should differ consistently in their cooperative behaviour, consistent individual differences in cooperative behaviour have rarely been documented. In this study, we used a long‐term data set on wild meerkats to quantify the repeatability of two types of cooperative care (babysitting and provisioning) within individuals and examined how repeatability varied across age, sex and status categories. Contributions to babysitting and provisioning were significantly repeatable and positively correlated within individuals, with provisioning more repeatable than babysitting. While repeatability of provisioning was relatively invariant across categories of individuals, repeatability of babysitting increased with age and was higher for subordinates than dominants. These results provide support for theoretical predictions that life‐history trade‐offs favour the evolution of consistent individual differences in cooperative behaviour and raise questions about why some individuals consistently help more than others across a suite of cooperative behaviours.     

Individual variation in behavioural plasticity: direct and indirect effects of boldness,exploration and sociability on habituation to predators in lizards

Little is known about the factors causing variation in behavioural plasticity and the interplay between personality and plasticity. Habituation to predators is a special case of behavioural plasticity. We investigated the direct and indirect effects of boldness, exploration and sociability traits on the habituation ability of Iberian wall lizards, considering exposure and sex effects. Individual boldness was consistent across several non-habituation contexts, but it did not significantly affect habituation. Exploration had a strong direct effect on habituation, with more exploratory individuals being able to habituate faster than less exploratory ones, probably because of their ability to assess risk better. Individual variation in habituation was also affected by sociability, but this was an indirect effect mediated by exposure to the predator. Less social individuals avoided refuges with conspecific cues, increasing exposure to the predator and eventually habituation. Finally, the direct effects of sex (females habituated faster than males) were opposite to its indirect effects through exposure. We conclude that risk assessment, instead of the proactivity–reactivity gradient usually considered in the literature, can affect behavioural plasticity through complex interactions between direct and indirect effects, including exploratory behaviour, degree of exposure to the predator and sex, which represent novel mechanisms generating inter-individual variation in plasticity.     

First- and second-order sociality determine survival and reproduction in cooperative cichlids

Cooperative breeders serve as a model to study the evolution of cooperation, where costs and benefits of helping are typically scrutinized at the level of group membership. However, cooperation is often observed in multi-level social organizations involving interactions among individuals at various levels. Here, we argue that a full understanding of the adaptive value of cooperation and the evolution of complex social organization requires identifying the effect of different levels of social organization on direct and indirect fitness components. Our long-term field data show that in the cooperatively breeding, colonial cichlid fish Neolamprologus pulcher, both large group size and high colony density significantly raised group persistence. Neither group size nor density affected survival at the individual level, but they had interactive effects on reproductive output; large group size raised productivity when local population density was low, whereas in contrast, small groups were more productive at high densities. Fitness estimates of individually marked fish revealed indirect fitness benefits associated with staying in large groups. Inclusive fitness, however, was not significantly affected by group size, because the direct fitness component was not increased in larger groups. Together, our findings highlight that the reproductive output of groups may be affected in opposite directions by different levels of sociality, and that complex forms of sociality and costly cooperation may evolve in the absence of large indirect fitness benefits and the influence of kin selection.     

Fitness consequences of variation in developmental temperature in a butterfly

We explored the adaptive significance of developmental plasticity in the tropical butterfly Bicyclus anynana using two experiments including temperature changes during ontogeny. In contrast to previous findings on adult acclimation, we could not find any evidence in support of adaptive developmental plasticity, as survival until adulthood was not enhanced when larval rearing temperatures matched the temperatures experienced during prepupal or pupal development. Extreme temperatures substantially reduced survival, supporting the ‘optimal developmental temperature’ hypothesis. Metamorphosis was more efficient at the higher rearing temperature of 27 °C, where egg hatching success was also higher, indicating that the lower temperature of 20 °C is already slightly stressful for this tropical butterfly.     

Mutants affecting plasticity of phototactic behaviour in Drosophila melanogaster

Eight mutants of Drosophila m. have been isolated that fail to display longlasting modification of “choice tube phototaxis”. Two different classes of stocks defective in plasticity of phototaxis (dipp) can be discriminated: (1) Mutants which are affected in the stability of the Prolonged Depolarising Afterpotential (PDA) in the peripheral retinula cells R1–R6. (2) Mutants with normal PDA but abnormalities in fast escape phototaxis especially under conditions where only receptor cells R7–R8 serve as input for this behavioural trait.     

Behavioural phenotype affects social interactions in an animal network

Animal social networks can be extremely complex and are characterized by highly non-random interactions between group members. However, very little is known about the underlying factors affecting interaction preferences, and hence network structure. One possibility is that behavioural differences between individuals, such as how bold or shy they are, can affect the frequency and distribution of their interactions within a network. We tested this using individually marked three-spined sticklebacks (Gasterosteus aculeatus), and found that bold individuals had fewer overall interactions than shy fish, but tended to distribute their interactions more evenly across all group members. Shy fish, on the other hand, tended to associate preferentially with a small number of other group members, leading to a highly skewed distribution of interactions. This was mediated by the reduced tendency of shy fish to move to a new location within the tank when they were interacting with another individual; bold fish showed no such tendency and were equally likely to move irrespective of whether they were interacting or not. The results show that animal social network structure can be affected by the behavioural composition of group members and have important implications for understanding the spread of information and disease in social groups.     

Host plant effects on the genetic variation and correlations in the individual performance of the Gypsy Moth

1. Quantitative genetic analysis of variation in host-use ability was performed in an extremely polyphagous species – the Gypsy Moth. Various life-history traits were investigated by applying a split-family two-environment experimental design, where 30 full-sibling families were reared on oak and Locust Tree leaves.
2. Feeding on Locust Tree leaves decreased preadult viability, prolonged development time, decreased pupal mass both in males and females, and decreased reproductive effort in females.
3. The majority of broad-sense heritabilities did not change across host plants.
4. Significant expression of genetic variation in diet breadth was observed for development time and pupal mass both in males and females, but not for female reproductive effort traits. The heritabilities of plasticities were, on average, lower than heritabilities of the traits themselves.
5. The majority of genetic correlations between the host plants were significantly positive. The only trade-off was found between reciprocal value of reproductive index and the average mass of a fertilized egg within oak. This means that selection for the decrease in relative reproductive investment (under starvation during gradation) will be followed by laying larger eggs. Given that the Gypsy Moth has a cyclic population dynamics, this negative genetic correlation could have a role in maintaining genetic variability in this species.     

Nectar production rates of Asclepias quadrifolia: causes and consequences of individual variation

The causes and reproductive consequences of individual variation in nectar production rates within a population of Asclepias quadrifolia were investigated. Two parameters were correlated with nectar production rate per flower: the root weight of the plant and the number of flowers in the inflorescence (umbel). Nectar production increased with increasing root weight but levelled off after a root size of about 3 g was reached. Nectar production decreased with increasing umbel size, but only for umbels that were greater than average size. A total of 57% of the variance in individual nectar production could be explained by these two variables with root weight accounting for 67% of the explained variance. Root weight is a good indicator of a plant's energetic status, indicating the importance of available energy in determining quantity of nectar produced. About 30% of the energy devoted to flowering is utilized in nectar production. Nectar production was significantly correlated with the male component of reproductive fitness, pollinaria removal, but not with the female component, pollinia insertion. Since pod production is limited by resource availability rather than the number of pollinia insertions, nectar production in A. quadrifolia is most closely associated with the maximization of the male function.