Individual-level personality influences social foraging and collective behaviour in wild birds

There is increasing evidence that animal groups can maintain coordinated behaviour and make collective decisions based on simple interaction rules. Effective collective action may be further facilitated by individual variation within groups, particularly through leader–follower polymorphisms. Recent studies have suggested that individual-level personality traits influence the degree to which individuals use social information, are attracted to conspecifics, or act as leaders/followers. However, evidence is equivocal and largely limited to laboratory studies. We use an automated data-collection system to conduct an experiment testing the relationship between personality and collective decision-making in the wild. First, we report that foraging flocks of great tits (Parus major) show strikingly synchronous behaviour. A predictive model of collective decision-making replicates patterns well, suggesting simple interaction rules are sufficient to explain the observed social behaviour. Second, within groups, individuals with more reactive personalities behave more collectively, moving to within-flock areas of higher density. By contrast, proactive individuals tend to move to and feed at spatial periphery of flocks. Finally, comparing alternative simulations of flocking with empirical data, we demonstrate that variation in personality promotes within-patch movement while maintaining group cohesion. Our results illustrate the importance of incorporating individual variability in models of social behaviour.     

Social foragers adopt a riskier foraging mode in the centre of their groups

Foraging in groups provides many benefits that are not necessarily experienced the same way by all individuals. I explore the possibility that foraging mode, the way individuals exploit resources, varies as a function of spatial position in the group, reflecting commonly occurring spatial differences in predation risk. I show that semipalmated sandpipers (Calidris pusilla), a social foraging avian species, tended to adopt a riskier foraging mode in the central, more protected areas of their groups. Central birds effectively used the more peripheral group members as sentinels, allowing them to exploit a wider range of resources within the same group at the same time. This finding provides a novel benefit of living in groups, which may have a broad relevance given that social foraging species often exploit a large array of resources.     

Previous foraging success influences web building in the spider Stegodyphus lineatus (Eresidae)

Stegodyphus lineatus (Eresidae) is a desert spider that buildsan aerial capture web on bushes in the Negev desert in southernIsrael. Web building for spiders is costly in energy, time,and risk of predation. Spiders should trade-off these costswith the benefits in terms of prey capture. We tested the hypothesisthat the previous foraging success of the spider influencesthe effort invested in foraging. Specifically, we asked whetheran increase in food intake causes spiders to reduce web renewalactivity and web size. Alternatively, time constraints on foragingand development, resulting from a short growing season, couldinduce spiders to continue foraging even when supplemented withprey. The cost of web building was measured as time and massloss. To build an average size web (about 150 cm²), we calculatedthat a spider requires 6 h and that spiders lose 3%-7% of their weight.In field experiments, spiders responded differently to food supplementationin 2 different years. In 1994, they improved their condition comparedto individuals whose webs were removed to reduce foraging opportunitiesand compared to control spiders. In 1995, spiders tested earlier inthe season than the previous year did not improve their conditionin response to prey supplementation. Nonetheless, in both years, food-supplementedspiders built significantly smaller webs than food-deprived andcontrol spiders. This result was confirmed in a laboratory experiment whereprey intake was controlled. We conclude that for S. lineatus immediateforaging risks outweigh the potential time constraints on foraging.     

Less is more: density influences the development of behavioural life skills in trout

Theory suggests that habitat structure and population density profoundly influence the phenotypic development of animals. Here, we predicted that reduced rearing density and increased structural complexity promote food search ability, anti-predator response and the ability to forage on novel prey, all behavioural skills important for surviving in the wild. Brown trout were reared at three densities (conventional hatchery density, a fourth of conventional hatchery density and natural density) in tanks with or without structure. Treatment effects on behaviour were studied on trout fry and parr, whereupon 20 trout from each of the six treatment groups were released in an enclosed natural stream and recaptured after 36 days. Fry reared at natural density were faster to find prey in a maze. Moreover, parr reared at natural density were faster to eat novel prey, and showed more efficient anti-predator behaviour than fish reared at higher densities. Furthermore, parr reared at reduced densities were twice as likely to survive in the stream as trout reared at high density. In contrast, we found no clear treatment effects of structure. These novel results suggest that reduced rearing densities can facilitate the development of behavioural life skills in captive animals, thereby increasing their contribution to natural production.     

The role of conformity in foraging when personal and social information conflict

There is currently considerable interest in the interplay betweenpersonal and social information in decision-making processes.Two experiments are presented exploring the relative use ofprior personal information and subsequent social informationin foraging decisions of guppies. Experiment 1 tested the assumptionthat when the use of information acquired through personal experienceis not costly, conflicting social information will be ignored.The assumption was confirmed because, when given a choice betweenfeeding at two food patches, at one of which they had previouslyseen conspecifics feed, individual fish with prior experienceof feeding at the alternative site chose the alternative, whereasfish with no prior experience chose the site at which theirconspecifics had fed. Experiment 2 tested theoretical predictionsthat when the use of information acquired through personal experienceis potentially costly, conflicting social information will beweighed more heavily than will personal information. The predictionwas confirmed because, when given a choice between feeding attwo food patches, one at which they had previously seen conspecificsfeed and one behind a visual barrier, individual fish with priorexperience of feeding behind the barrier chose the site at whichtheir conspecifics had fed. These findings suggest that conformitycan promote social learning in naïve individuals, but priorexperience can insulate individuals from conformity providedthe costs of relying on that experience are small. In addition,the experiments highlight the fact that personal and socialinformation are not always weighed equally.     

Flights of fear: a mechanical wing whistle sounds the alarm in a flocking bird

Animals often form groups to increase collective vigilance and allow early detection of predators, but this benefit of sociality relies on rapid transfer of information. Among birds, alarm calls are not present in all species, while other proposed mechanisms of information transfer are inefficient. We tested whether wing sounds can encode reliable information on danger. Individuals taking off in alarm fly more quickly or ascend more steeply, so may produce different sounds in alarmed than in routine flight, which then act as reliable cues of alarm, or honest ‘index’ signals in which a signal''s meaning is associated with its method of production. We show that crested pigeons, Ocyphaps lophotes, which have modified flight feathers, produce distinct wing ‘whistles’ in alarmed flight, and that individuals take off in alarm only after playback of alarmed whistles. Furthermore, amplitude-manipulated playbacks showed that response depends on whistle structure, such as tempo, not simply amplitude. We believe this is the first demonstration that flight noise can send information about alarm, and suggest that take-off noise could provide a cue of alarm in many flocking species, with feather modification evolving specifically to signal alarm in some. Similar reliable cues or index signals could occur in other animals.     

The fitness consequences of kin-biased dispersal in a cooperatively breeding bird

Cooperative alliances among kin may not only lead to indirect fitness benefits for group-living species, but can also provide direct benefits through access to mates or higher social rank. However, the immigrant sex in most species loses any potential benefits of living with kin unless immigrants disperse together or recruit relatives into the group in subsequent years. To look for evidence of small subgroups of related immigrants within social groups (kin substructure), we used microsatellites to assess relatedness between immigrant females of the cooperatively breeding superb starling, Lamprotornis superbus. We determined how timing of immigration led to kin subgroup formation and if being part of one influenced female fitness. Although mean relatedness in groups was higher for males than females, 26% of immigrant females were part of a kin subgroup with a sister. These immigrant sibships formed through kin recruitment across years more often than through coalitions immigrating together in the same year. Furthermore, females were more likely to breed when part of a kin subgroup than when alone, suggesting that female siblings form alliances that may positively influence their fitness. Ultimately, kin substructure should be considered when determining the role of relatedness in the evolution of animal societies.     

The effects of foraging demand on social interactions in a laboratory group of Bonnet Macaques

The relationship between foraging demand and social behavior was experimentally studied in a laboratory group of bonnet macaques. Fourteen adult animals were housed in a large outdoor enclosure containing three shallow gravelfilled circular containers that served as the foraging sites. During the experimental foraging sessions raisins were placed in the containers and the social and foraging behaviors of the group were observed for 50 min following the distribution of raisins. Three types of foraging conditions were inter-spersed with one another on different test days: (1) surface load— raisins placed on top of the gravel; (2) buried load— raisins hidden underneath the gravel; and (3) sham load— no raisins placed at the foraging sites. Three basic foraging patterns, defined along a temporal dimension, were seen. One group of animals completed 50% of their total foraging by the end of the first 15 min. A second group foraged more steadily through the session. A third group foraged late, completing 50% of their foraging during the last half of the session. The foraging patterns were similar in the buried and surface condition, although the patterns were more compressed during the surface condition. More aggression and more avoidance of other animals occurred in the buried condition than in the surface condition. Very little foraging occurred during the sham condition. There was no clear relationship between the patterns of interaction during foraging and nonforaging observation sessions. The results suggest the value of manipulative laboratory studies in examining the relationship between ecological variables and social behavior in nonhuman primates.     

The effect of social facilitation on foraging success in vultures: a modelling study

The status of many Gyps vulture populations are of acute conservation concern as several show marked and rapid decline. Vultures rely heavily on cues from conspecifics to locate carcasses via local enhancement. A simulation model is developed to explore the roles vulture and carcass densities play in this system, where information transfer plays a key role in locating food. We find a sigmoid relationship describing the probability of vultures finding food as a function of vulture density in the habitat. This relationship suggests a threshold density below which the foraging efficiency of the vulture population will drop rapidly towards zero. Management strategies should closely study this foraging system in order to maintain effective foraging densities.     

The early bird gets the worm: foraging strategies of wild songbirds lead to the early discovery of food sources

Animals need to manage the combined risks of predation and starvation in order to survive. Theoretical and empirical studies have shown that individuals can reduce predation risk by delaying feeding (and hence fat storage) until late afternoon. However, little is known about how individuals manage the opposing pressures of resource uncertainty and predation risks. We suggest that individuals should follow a two-part strategy: prioritizing the discovery of food early in the day and exploiting the best patch late in the day. Using automated data loggers, we tested whether a temporal component exists in the discovery of novel foraging locations by individuals in a mixed-species foraging guild. We found that food deployed in the morning was discovered significantly more often than food deployed in the afternoon. Based on the diurnal activity patterns in this population, overall rates of new arrivals were also significantly higher than expected in the morning and significantly lower than expected in the afternoon. These results align with our predictions of a shift from patch discovery to exploitation over the course of the day.     

Factors affecting offspring survival and development in a cooperative bird: social, maternal and environmental effects

1. In many noncooperative vertebrates, maternal effects commonly influence offspring survival and development. In cooperative vertebrates, where multiple adults help to raise young from a single brood, social effects may reduce or replace maternal effects on offspring. 2. Factors affecting offspring survival and development at different stages (fledging, nutritional independence and adulthood) were tested in the cooperatively breeding Arabian babbler to determine the relative importance of social, maternal and environmental factors at each stage. An influence of maternal effects was found during the nestling stage only. 3. Social factors affected the survival and development of young at all stages. The amount of food received from helpers influenced post-fledging weight gain, development of foraging skills, and survival to reproductive age. Environmental effects were also important, with groups occupying high-quality territories more likely to produce young that survived to maturity. 4. The strong influence of helper contributions on the survival and development of young at all stages from hatching to maturity suggests social factors may have important long-term effects on offspring fitness in cooperative societies. Traditional measures of offspring survival in cooperative birds, which commonly measure survival to fledging age only, may underestimate the significant benefit of helper contributions on the survival and development of young.     

Variable postfledging care in a cooperative bird: causes and consequences

Prolonged postfledging care is a commonly observed behaviorin many cooperatively breeding species and has been shown toprovide young with both survival and developmental benefits.However, the causes of intraspecific variation in postfledgingcare and the consequences of this variation on the developmentof young remain unclear. Here we investigate factors affectingthe duration of postfledging care in the cooperatively breedingpied babbler (Turdoides bicolor). We show that the durationof care is variable (40–97 days) and is determined primarilyby the cost of care. Adults in groups with a low adult:fledglingratio were unable to maintain body mass during the period ofchick provisioning and subsequently ceased care of young earlier.This had a strong influence on offspring development: fledglingsthat received longer periods of care attained higher foragingefficiency and body mass than their counterparts at 6 monthsof age. The duration of postfledging care also had long-termeffects, with individuals that received longer periods of postfledgingcare more likely to successfully disperse from their natal group.This had important fitness implications as successful dispersersbecame reproductively active at an earlier age than their "failed-disperser"counterparts. These findings highlight the importance of consideringlong-term influences when assessing the benefits of prolongedpostfledging care on offspring fitness and development in cooperativesocieties.     

Stress and the costs of extra-territorial movement in a social carnivore

Costs associated with extra-territorial movement are believed to have favoured the evolution of delayed dispersal and sociality across a range of social vertebrates, but remain surprisingly poorly understood. Here we reveal a novel mechanism that may contribute substantially to the costs of extra-territorial movement: physiological stress. We show that subordinate male meerkats, Suricata suricatta, exhibit markedly elevated faecal glucocorticoid metabolite levels (a non-invasive measure of hypothalamic–pituitary–adrenal axis activity) while conducting extra-territorial prospecting forays. While brief increases in glucocorticoid levels are unlikely to be costly, chronic elevations, arising from prolonged and/or frequent forays, are expected to compromise fitness through their diverse negative effects on health. Our findings strongly suggest that prolonged extra-territorial movements do result in chronic stress, as the high glucocorticoid levels of prospectors do not diminish on longer forays and are no lower among males with greater prospecting experience. A generalized ‘stress’ of extra-territorial movement may therefore have strengthened selection for delayed dispersal and sociality in this and other species, and favoured the conduct of brief forays over extended periods of floating. Our findings have implications too for understanding the rank-related distribution of physiological stress in animal societies, as extra-territorial movements are often conducted solely by subordinates.     

The consequences of adult foraging success on the components of lifetime fitness in a semelparous,sit and wait predator

Summary We evaluated the role of adult foraging success in the lifetime fitness of female crab spidersMisumena vatia. Misumena are semelparous, sit and wait predators that hunt for insect prey on flowers, in this study primarily on inflorescences (umbels) of milkweedAsclepias syriaca. We used path analysis to integrate previously performed experimental and observational studies, thereby establishing the magnitude, correlations and causal relationships of key foraging and life history variables and their roles in lifetime fitness. A path proceeding from maternal hunting patch choice through maternal mass, clutch mass and number of dispersal-age young was the dominant element and explained a large part of the variation. Other paths that incorporated parasitism of the egg mass and predation of young leaving the nests made only small impacts on variation. No trade-offs were found, primarily because a single factor, maternal mass (a maternal effect) resulting from foraging success, provided major benefits for successive life history stages. Since differences in the numbers of eggs, egg loss and mortality at dispersal resulted almost entirely from differences in maternal mass, they are controlled by the maternal generation and, thus, are appropriately attributed to the lifetime fitnesses of the mothers, rather than to those of their offspring.     

The adaptive significance of stealing in a marine bird and its relationship to parental quality

Kleptoparasitism (food theft) is a tactic used opportunisticallyby many foraging birds, but little is known about its fitnessbenefits. Here we show that habitual kleptoparasitism by individualparent roseate terns (Sterna dougallii) is associated with consistentlysuperior reproductive performance relative to nonkleptoparasitic("honest") parents, as measured by growth and survival to fledgingamong their offspring. In broods of two, both chicks of kleptoparasiticparents exhibited superior growth performance during the middleand later stages of the rearing period, relative to chicks ofhonest parents. This difference was especially pronounced insecond-hatched chicks, whose survival is highly variable amongyears and dependent on food availability. Over a 10-year period,average productivity (number of chicks fledged per pair) wassignificantly higher among kleptoparasites than among honestparents, with a larger relative difference during years of foodshortage. Our study indicates that kleptoparasitism in roseateterns is an important component of parental quality and providesthe first evidence that food stealing is associated with enhancedfitness in a facultatively kleptoparasitic seabird.