Exploring the effects of individual traits and within-colony variation on task differentiation and collective behavior in a desert social spider

Social animals are extraordinarily diverse and ecologically abundant. In understanding the success of complex animal societies, task differentiation has been identified as a central mechanism underlying the emergence and performance of adaptive collective behaviors. In this study, we explore how individual differences in behavior and body size determine task allocation in the social spider Stegodyphus dumicola. We found that individuals with high body condition indices were less likely to participate in prey capture, and individuals’ tendency to engage in prey capture was not associated with either their behavioral traits or body size. No traits were associated with individuals’ propensity to participation in web repair, but small individuals were more likely to engage in standard web-building. We also discovered consistent, differences among colonies in their collective behavior (i.e., colony-level personality). At the colony level, within-colony variation in behavior (aggressiveness) and body size were positively associated with aggressive foraging behavior. Together, our findings reveal a subtly complex relationship between individual variation and collective behavior in this species. We close by comparing the relationship between individual variation and social organization in nine species of social spider. We conclude that intraspecific variation is a major force behind the social organization of multiple independently derived lineages of social spider.     

Evidence for differential selection and potential adaptive evolution in the worker caste of an inquiline social parasite

Social parasites exploit the socially managed resources of social insect colonies in order to maximise their own fitness. The inquilines are among the most specialised social parasites, because they are dependent on being fully integrated into their host's colony throughout their lives. They are usually relatives of their host and so share ancestral characteristics (Emery's rule). Closely related inquiline-host combinations offer a rare opportunity to study trade-offs in natural selection. This is because ancestral adaptations to a free-living state (e.g. the production of a worker caste) become redundant and may be replaced by novel, parasitic traits as the inquiline becomes more specialised. The dynamics of such processes are, however, unknown as virtually all extant inquiline social parasites have completely lost their worker caste. An exception is Acromyrmex insinuator, an incipient permanent social parasite of the leaf-cutting ant Acromyrmex echinatior. In the present study, we document the size distribution of parasite and host workers and infer how selection has acted on A. insinuator to reduce, but not eliminate, its investment in a worker caste. We show that the antibiotic producing metapleural glands of these parasite workers are significantly smaller than in their host counterparts and we deduce that the metapleural gland size in the host represents the ancestral state. We further show experimentally that social parasite workers are more vulnerable to the general insect pathogenic fungus Metarhizium than are host workers. Our findings suggest that costly disease resistance mechanisms are likely to have been lost early in inquiline evolution, possibly because active selection for maintaining these traits became less when parasite workers had evolved the ability to exploit the collective immune system of their host societies.     

Spatial proximity loggers for recording animal social networks: consequences of inter-logger variation in performance

Social network analysis has become an increasingly popular method to link individual behaviour to population level patterns (and vice versa). Technological advances of recent years, such as the development of spatial proximity loggers, have enhanced our abilities to record contact patterns between animals. However, loggers are often deployed without calibration which may lead to sampling biases and spurious results. In particular, loggers may differ in their performance (i.e., some loggers may over-sample and other loggers may under-sample social associations). However, the consequences of inter-logger variation in logging performance has not been thoroughly considered or quantified. In this study, proximity loggers made by Sirtrack Ltd. were fitted to 20 dairy cows over a 3-week period. Contact records resulting from field deployment demonstrated variability in logger performance when recording contact duration, which was highly consistent for each logger over time. Testing loggers under standardised conditions suggested that inter-logger variation observed in the field was due to a combination of intrinsic variation in devices, and environmental/behavioural effects. We demonstrate the potential consequences that inter-logger variation in logging performance can have for social network analysis; particularly how measures of connectivity can be biased by logging performance. Finally, we suggest some approaches to correct data generated by proximity loggers with imperfect performance, that should be used to improve the robustness of future analyses.     

Habitat structure helps guide the emergence of colony-level personality in social spiders

Studying the environmental factors that guide the emergence of collective behaviors is instrumental to understanding the ecology and evolution of animal societies. Although recent work has provided insights into the demographic factors that influence inter-colony variation in collective behavior (i.e., colony-level personality or collective personality), relatively few studies have investigated how the physical environment (e.g., habitat structure) affects colony-level personality. Here, we study the emergence of collective personality in prey capture behavior in the social spider, Stegodyphus dumicola. We measured collective prey capture behavior four times over 36 days in a classic repeated measures design. We used four different artificial habitat (web support) structures in three different treatments: habitat structure was either (1) fixed and undisturbed, (2) disturbed with a complete removal of webbing between each measurement, or (3) disturbed with changes of habitat structure between each measurement. Our results revealed that repeatability in colony-level personality was retained as long as habitat structure was not altered. However, the repeatability of colony-level personality declined precipitously when groups were forced to build their webs on novel habitat structures. Furthermore, habitat structure affected collective capture behavior, that is, latency to attack and the number of attackers differed among colonies on different habitat structures. Collectively, our data demonstrate that habitat structure is instrumental in shaping both the mean and repeatability of the collective behavior of colonies and may influence overall foraging success.     

Association patterns of spider monkeys: the influence of ecology and sex on social organization

Summary In this paper I consider how the costs and benefits of group living in spider monkeys (Ateles geoffroyi) vary between troop members. The results suggest that ecological factors set an upper limit to the number of spider monkeys that can associate and still efficiently exploit the available resources. In addition, the needs of the individual appears to influence the type and size of the subgroup it chooses. Adult males band together, travel over a large area, and are frequently sighted near the community's boundary. In contrast, adult females spend more time solitary than males and have association patterns that are strongly influenced by the presence of a dependent infant. Females with dependent infants tend to travel in small subgroups or alone, avoid the boundaries of the community's home range, and exhibit a restricted pattern of use of their range. The results suggest that males may be attempting to locate females with which they can breed, while mothers attempt to protect their infants by avoiding conspecifics and potentially dangerous situations near territorial boundaries.     

Sympatric species of threespine stickleback differ in their performance in a spatial learning task

Increasing evidence suggests that cognitive function is under selection in diverse taxa and that this results in different cognitive abilities in taxa experiencing different selective environments. For example, species inhabiting spatially complex environments might be expected to have good spatial learning ability. We investigated whether local habitat conditions influence learning by comparing the performance of two recently diverged species of threespine stickleback (Gasterosteus aculeatus complex) in a spatial learning task. The two species reside sympatrically in several lakes. Benthics occupy the spatially structured vegetated littoral zone, whereas limnetics occupy the spatially homogenous open-water pelagic zone. We trained fish to locate a hidden reward in a T-maze and asked whether they did so by learning a body-centred pattern of movement (turn left or right) or by using plant landmarks. Both benthics and limnetics used turn and landmark cues, but benthics learnt the task almost twice as quickly as limnetics. This difference is consistent with the hypothesis that benthic and limnetic sticklebacks are equipped with spatial learning abilities well suited to the spatial complexity of their littoral and pelagic habitats. Our findings add to the understanding of the evolution of learning.     

Molecular genetic and behavioral analysis of social organization in the Asian elephant (Elephas maximus)

We report on the genetic evaluation and behavioral study of social organization in the Asian elephant (Elephas maximus). Although Asian elephants and African elephants (Loxodonta africana) were previously thought to have similar social organizations, our results demonstrate a substantial difference in the complexity and structure of Asian elephant social groupings from that described for African savanna elephants. Photographic cataloging of individuals, radio telemetry, and behavioral observations in Ruhuna National Park, Sri Lanka, enabled us to assign associated females and young to four groups with overlapping ranges. Genetic sampling of individuals from the four groups in Ruhuna National Park and three other groups in surrounding areas, conducted through PCR amplification and sequencing of mitochondrial DNA from dung, supported the matriarchal nature of female groups and the lack of inter-group transfer of females. Behaviorally and genetically, the identified social groups were best described as ”family groups”. We did not find any evidence for the existence of social groups of higher complexity than family groups. Received: 25 March 2000 / Received in revised form: 28 March 2000 / Accepted: 1 April 2000     

Morphological variation in a population of grey-crowned babblers: Correlations with variables affecting social behavior

Summary Variation in weight, wing length and bill length in a population of grey-crowned babblers is influenced primarily by sex and age, but correlations with size of social unit, with reproductive success, and with vegetation are also detectable (Table 1). The latter correlations vary with sex, age, and status as helper or breeder. Differential wear according to behavioral role, competition for status, incubation, and inheritance are discussed as possible causal mechanisms. Helpers were not detectably smaller in any dimension than breeders of the same age and sex. Male and female non-breeding helpers differ in patterns of morphological correlation, suggesting that they have different behavioral roles. Breeding males have a unique pattern of morphological correlation, suggesting that their foraging behavior differs from breeding females and non-breeders.     

Spatial,temporal, and behavioral patterns in emergence of zooplankton in the lagoon of Heron Reef,Great Barrier Reef,Australia

A total of 34 zooplanktonic taxa were common in emergence trap, reentry trap, and net-tow samples taken in the lagoon of Heron Reef, Great Barrier Reef, between 27 February and 22 March and between 11 June and 4 July 1985. Twenty-nine of these taxa were classified as demersal (17 taxa), meroplanktonic and larval (10 taxa), or incidental (2 taxa). The remaining five multispecific groups yielded variable results. Differences were observed between two locations separated by 200 m, with emergence 2 to 62 times greater for 22 taxa at a deeper site with larger coral formations. Most zooplankters were more common (2 to 122 times) in samples from 1 m² areas around 0.25 to 0.5 m² patches of branching coral. However, two species of copepods, Pseudodiaptomus colefaxi and Metis holothuriae, were taken in greater numbers (3 to 12 times) from open sand. Seasonal increases (2 to 322 times) were observed for 13 adult taxa and 8 classes of larvae or juveniles in the summer and for 10 adult taxa and one larval group in the winter. Fourteen of the 17 demersal taxa and 4 groups of large larvae or juveniles emerged in numbers 2 to 323 times greater during lunar quarters or new moons. In contrast, 6 larval taxa and 4 groups of small or transparent adults displayed significant emergence during full moons. Five diurnal emergence patterns were shown by 27 taxa, with patterns varying primarily among lunar periods. The variety of taxon-specific patterns observed in this study highlight a need for caution when generalizing about demersal zooplankton.     

Food exploitation by a winter flock of greylag geese: behavioral dynamics,competition and social status

Summary To investigate the dynamics of the winter flock patch exploitation, feeding experiments were performed with 140 semi-tame, free-roaming greylag geese (Anser anser). Three different initial densities of barley were offered on a 50-m² patch: low (1600 grains/m²), intermediate (3300/m²) and high (I 1500/m²). Goose numbers on the patch, peck rates, and frequencies of agonistic encounters and of alert postures were observed by scanning the flock and in focal individuals. At low and intermediate initial food densities, peck rates decreased with food density, whereas at high food density, peck rates decreased only slightly over the feeding bout. Agonistic interactions increased as food decreased. A switch from exploitation to interference competition occurred at a threshold of approximately 900 remaining grains per square meter. With high initial food density, agonistic encounters began to increase gradually after 20–30 min. The same general patterns were observed for frequency of alertness. Family members occupied the patch for the longest time periods. The family female and the offspring fed most intensely. The family gander however, fed significantly less but was more aggressive and vigilant than all other social categories. Solitary geese spent their time on the patch doing little else but feeding and were the first to leave. It seems that high-ranking families and low-ranking singles adopt different competitive strategies, the former being superior interference competitors whereas the latter are forced scramble competitors.This study is dedicated to the memory of Konrad Lorenz Correspondence to: K. Kotrschal     

Physiological correlates of genetic variation for rate of behavioral development in the honeybee, Apis mellifera

Two factors that influence age at onset of foraging in honeybees are juvenile hormone (JH) and colony age demography (older bees inhibit behavioral development of younger bees). We tested the hypothesis that genetic variation among bees for these factors influences genetic variation in behavioral development. Pairs of colonies showing genetic differences in rates of behavioral development were identified in a screening experiment and bees from these colonies were used for physiological and behavioral assays. Six pairs were assayed, three with European bees only and three with both European and Africanized bees. There was genetic variation for the following four components: (1) production of JH in four pairs (experiment 1); (2) sensitivity to JH in three pairs (experiment 2); (3) sensitivity to social inhibition in three pairs (experiment 3), and (4) potency of social inhibition in four pairs (experiment 4). Cross-fostering assays (experiment 5), which allowed all four components to be evaluated simultaneously, revealed genetic variation for production of JH, sensitivity to JH, or sensitivity to social inhibition in five of six pairs, and potency of social inhibition in five of six pairs. There was often evidence for genotypic differences in more than one component, and no consistent pattern of association among any of the components. Africanized bees had faster rates of behavioral development than European bees, but there were no racial differences in patterns of variation among the four components. These results indicate that there are at least several, apparently distinct, physiological processes associated with JH and colony age demography upon which natural selection can act to alter the rate of behavioral development in honeybees. Received: 8 December 1998 / Received in revised form: 29 July 1999 / Accepted: 8 August 1999     

Recognition of caste and mating status maintains monogyny in the ant Aphaenogaster senilis

In ants dispersing through colony fission, queens mate near their natal nest and found a new society with the help of workers. This allows potential future queens to challenge the mother queen’s reproductive monopoly. Conflicts might be resolved if the mated queen signals her presence and the workers control the developmental fate of the diploid larvae (whether they develop to worker or queen). In this study we sought to determine whether, in the fission-performing ant Aphaenogaster senilis, conflicts between queens for control of the colony are resolved by the resident queen signalling her mating status. Virgin queens were less effective than newly mated queens in inhibiting queen rearing. Moreover, potential challenger queens were recognized and heavily aggressed independent of mating status. Chemical analyses showed that mating status was associated with changes in cuticular hydrocarbon and poison gland composition, but not in Dufour’s gland composition. Cuticular dimethylalkanes were identified as potential constituents that signal both caste (present in queens only) and mating status (mated queens have higher amounts). We hypothesised that pheromone emission by virgin queens did not reach the threshold needed to fully inhibit larval development into queens but was sufficiently high to stimulate overt aggression by mated queens. These findings provide evidence for the complexity of chemical communication in social insects, in which a small number of signals may have a variety of effects, depending on the context.     

Habitat-specific variation and performance trade-offs in shell armature of New Zealand mudsnails

Studies documenting phenotypic variation among populations show that ecological performance in one activity is sometimes traded off against another. Identifying environment-specific costs and benefits associated with performance trade-offs is fundamental to knowing how conflicting selection pressures shape phenotype-environment matching in populations. We studied phenotypic variation in shell armature (spininess) of the New Zealand mudsnail, Potamopyrgus antipodarum (Gray), and explored how this variability relates to performance trade-offs between flow resistance and predator deterrence. Smooth- and spiny-shell morphotypes exist in populations in New Zealand streams and lakes, but the patterns and correlates of spatial variation of these phenotypes, and the possible hydrodynamical constraints and antipredatory benefits associated with spiny shell armature, are unknown. Samples from 11 rivers and nine lakes on the South Island showed that, on average, nearly 70% of snails in streams were smooth-shelled, whereas >80% of snails in lakes were spiny, suggesting dissimilar selective pressures between habitats. A laboratory flume experiment revealed that spines collected seston (i.e., suspended algae) at current speeds <40 cm/s, making spiny morphs more prone to flow-induced dislodgment than smooth morphs. However, a fish feeding experiment showed that one benefit of spines on shells was a decrease in predation risk from the common bully (Gobiomorphus cotidianus), a widespread predator of mudsnails in both streams and lakes. All snails egested by bullies were dead, further suggesting that these fishes may exert strong lethal effects on mudsnail populations in nature. Spine expression in lakes also appeared to be temperature related. We conclude that functional trade-offs between risk of flow-induced dislodgment and risk of fish predation affect shell armature frequencies of Potamopyrgus in freshwater habitats.     

Ecological costs and benefits of communal behavior in a presocial spider

Summary Philoponella oweni (Uloboridae) is a facultatively communal spider found in the southwestern United States. Solitary females and communal groups of females coexist in the same habitats. Communal females are known to produce more eggs than solitary females. Communal females might acquire the energy for the production of additional eggs by one of three means: they might have a longer life span, and thus more time in which to acquire energy; web maintenance might be more efficient in a communal group; or communal females might spend more time feeding. Field data from a marked population in Arizona showed the third alternative was true.Two possible causes for the greater time spent feeding by communal females were investigated; either the communal web or group is a more efficient trap for insects, or the communal groups are located at sites where insects are more abundant. Analysis of feeding rates and sticky trap data confirmed the second alternative.Potential web attachment sites are limited in some habitats, and mortality during dispersal is very high. When potential web attachment sites are scarce more of the population is found in communal groups; when they are abundant more of the population is solitary. The pattern of dispersal and overwintering is this species suggests that females renewing a colony in the spring could be related.     

Material, energy and environmental performance of technological and social systems under a Life Cycle Assessment perspective

Selected energy and material resource conversion systems are compared in this paper under an extended LCA point of view. A multi-method multi-scale assessment procedure is applied in order to generate consistent performance indicators based on the same set of input data, to ascertain the existence of constraints or crucial steps characterized by low conversion efficiency and to provide the basis for improvement patterns. Optimizing the performance of a given process requires that many different aspects are taken into account. Some of them, mostly of technical nature, relate to the local scale at which the process occurs. Other technological, economic and environmental aspects are likely to affect the dynamics of the larger space and time scales in which the process is embedded. These spatial and time scale effects require that a careful evaluation of the relation between the process and its surroundings is performed, so that hidden consequences and possible sources of inefficiency and impact are clearly identified. In this paper we analyse and compare selected electricity conversion systems, alternative fuels and biofuels, waste management strategies and finally the time evolution of an urban system, in order to show the importance of a multiple perspective point of view for the proper evaluation of a system's environmental and resource use performance.     

Dispersal behaviour in a subsocial spider: group conflict and the effect of food availability

This study investigates proximate factors influencing dispersal behaviour in the subsocial spider Amaurobius ferox. Dispersal of spiderlings from the natal web occurred as a progressive sigmoidal function (mean duration of dispersal period=31 days), with considerable individual variability in developmental instar and body mass within the clutch at the time of dispersal. The spiderlings showed a significant decrease in group cohesion on the 6th day after their second moult, which corresponded to the beginning of the dispersal period. Mutual aggressiveness appeared when the spiderlings began to show predatory behaviour (4th day post-second moult) and increased over the course of the dispersal period. While lack of prey in the maternal nest accelerated dispersal behaviour, the addition of prey items lengthened the dispersal period in previously non-fed clutches. Individuals that dispersed were smaller than the remaining individuals when measured on the day 50% of the clutch had dispersed. Timing of the appearance of the developmental characters (second moult, predation activity, agonistic behaviour against siblings, reduction of group cohesion, dispersal) suggests that the dispersal trait might have evolved in consequence of these different functional behaviours. Received: 10 August 1999 / Received in revised form: 15 March 2000 / Accepted: 1 April 2000