Foraging bumblebees do not rate social information above personal experience

Foraging animals can acquire new information about food sources either individually or socially, but they can also opt to rely on information that they have already acquired, termed “personal information”. Although social information can provide an adaptive shortcut to new resources, recent theory predicts that investing too much time in acquiring new information can be detrimental. Here, we investigate whether foraging bumblebees (Bombus terrestris) strategically prioritize personal information unless there is evidence of environmental change. All bees in our study had personal information that one species of artificial flower was rewarding, and bees in the scent group then experienced social information about an alternative-scented species inside the nest, while a control group did not. On their next foraging bout, bees in both groups overwhelmingly used personal information when deciding where to forage. When bees subsequently learnt that the rewards offered by their preferred species had dwindled, bees that had social information were no quicker to abandon their personal information than control bees, but once they had sampled the alternative flowers, they showed greater commitment to that species than control bees. Thus, we found no evidence that social information is particularly important when personal information fails to produce rewards (a “copy when established behaviour is unproductive” strategy). Instead, bees used social information specifically to complement personal information.     

Subhabitat selection by foraging threespine stickleback (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>): previous experience and social conformity

Any mechanism that allows animals to increase their foraging efficiency is likely to be selected for, including the ability to learn to recognise and subsequently discriminate between habitat types based on their profitability. In a series of laboratory studies, we manipulated prey densities across two different experimental subhabitats and demonstrated that threespine stickleback (Gasterosteus aculeatus) can develop foraging preferences for subhabitats that have previously yielded prey. Fish were not recalling the spatial location of prey patches; rather, they were discriminating between subhabitats based on foraging experience there and allocating foraging effort accordingly. Foraging preferences took around 14 days to develop, and once established, they persisted independently of experimental prey density, suggesting that fish were using experience rather than real-time sampling to select foraging grounds. When we presented focal fish with social information cues, we found that they preferentially used local enhancement and current public information cues when they conflicted with previous experience, but that they did not use prior public information. This suggests that in the presence of conspecifics, individuals prioritise social conformity over the use of private information. We discuss our results in the context of optimal foraging and the trade-offs associated with balancing conflicting private and social information.     

Vicarious sampling: the use of personal and public information by starlings foraging in a simple patchy environment

Group foragers may be able to assess patch quality more efficiently by paying attention to the sampling activities of conspecifics foraging in the same patch. In a previous field experiment, we showed that starlings foraging on patches of hidden food could use the successful foraging activities of others to help them assess patch quality. In order to determine whether a starling could also use another individual’s lack of foraging success to assess and depart from empty patches more quickly, we carried out two experimental studies which compared the behaviour of captive starlings sampling artificial patches both when alone and when in pairs. Solitary starlings were first trained to assess patch quality in our experimental two-patch system, and were then tested on an empty patch both alone and with two types of partner bird. One partner sampled very few holes and thus provided a low amount of public information; the other sampled numerous holes and thus provided a high amount of public information. In experiment 1, we found no evidence of vicarious sampling. Subjects sampled a similar number of empty holes when alone as when with the low and high information partners; thus they continued to rely on their own personal information to make their patch departure decisions. In experiment 2, we modified the experimental patches, increasing the ease with which a bird could watch another’s sampling activities, and increasing the difficulty of acquiring accurate personal sampling information. This time, subjects apparently did use public information, sampling fewer empty holes before departure when with the high-information partner than when with the low-information partner, and sampling fewer holes when with the low-information partner than when alone. We suggest that the degree to which personal and public information are used is likely to depend both on a forager’s ability to remember where it has already sampled and on the type of environment in which foraging takes place. Received: 31 January 1995/Accepted after revision: 11 September 1995     

Reciprocity and trades in wild West African chimpanzees

Why do animals help other individuals and provide benefits to the recipient, sometimes at personal cost? In this study, we aim to determine if some of the helpful behaviors observed in a group of wild chimpanzees (Taï chimpanzee group, Côte d'Ivoire, West Africa) are exchanged among individuals resulting in a net benefit for both participants. We adopted an inclusive view of exchanges by considering that all commodities (i.e., social behaviors as grooming, sex, support, as well as resources, such as meat) can be exchanged. This would result in “market” type social interactions in which debts of one commodity can be met by giving the same or other commodities. We investigated whether both in-kind exchanges and trades of commodities occurred. The Taï chimpanzees reciprocated the amount of grooming they received and were more likely to give support in agonistic conflicts to those who also supported them. In addition, they traded support for meat and meat for sex. Both male and female chimpanzees exchanged many different commodities, which seemed to result in long-term balanced relationships. Our results suggest that wild chimpanzees rely on other group members to obtain many of the important commodities they need.     

The anxious vole: the impact of group and gender on collective behavior under life-threat

Social animals behave collectively in order to maintain a cohesive group. This collective behavior is often led by a few individuals of specific gender, social rank, or spatial physical location in the group (i.e., perimeter or front). We examined how individual social voles (Microtus socialis) in same-gender compared with mixed-gender groups respond to an owl attack. We found that anxiety level, as measured by the time that each individual spent in less-sheltered sectors (open arms of elevated plus-maze and center of open arena), was affected by both the social context and the gender of the tested individuals. While both female and male voles generally reduced their activity in the open following owl attack, males in mixed-gender groups were exceptional in dichotomizing into those that spent a short period and those that spent a long period in the open arms of the plus-maze. Based on the similar responses of the same-gender groups, we suggest that anxiety is contagious, and based on the lower anxiety level of the mixed-gender groups, we suggest that natural groups that comprise both males and females are better able to cope with life-threat compared with same-gender groups. Finally, we suggest that the differential responses of males in the mixed-gender groups were due to a few males that displayed a low level of anxiety. These males were probably individuals of high social rank, and their response reflects their natural protective role, as previously described in social voles.     

Socially acquired predator recognition in complex ecosystems

Social animals acquire information on predator identities through social learning, where individuals with no prior experience learn from experienced members of the group. However, a large amount of uncertainty is often associated with socially acquired information especially in cases of cross-species learning. Theory predicts that socially acquired information from heterospecifics should take more repetitions to develop in complex ecosystems where the number of participants is greater. Our work focuses on coral reef fish as their social and communal lifestyles, along with their complex life histories, make them an ideal model to test for socially acquired predator recognition. Specifically, we tested if Pomacentrus wardi were capable of transmitting the recognition of an unknown predator, Pseudochromis fuscus, to closely related Pomacentrus moluccensis and phylogenetically distant Apogon trimaculatus. Individuals of both species were able to learn the predator's identity from experienced P. wardi based on a single conditioning event. It is somewhat surprising how fast social learning occurred particularly for the distantly related cardinalfish. This study demonstrates the widespread nature of social learning as a method of predator recognition in biologically complex ecosystems, and highlights that the benefits of responding to uncertain information may override the costs associated with lost foraging opportunities.     

Humans use social information to adjust their quorum thresholds adaptively in a simulated predator detection experiment

Quorum sensing is used in many biological systems to increase decision accuracy. In quorum sensing, the probability that an individual adopts a behavior is a nonlinear function of the number of other individuals adopting this behavior. From an optimal decision-making perspective, individuals should adjust their quorum threshold to the particulars of the decision problem. Recent work predicts that a key factor here is the quality of social information. In particular, it is predicted that individuals should adjust their quorum thresholds such that it lies in between the average true and false positive rate of the other group members. We here test this prediction with a predator detection experiment. First, human groups observed a group of animals (projected on a white screen) in which a predator was present or absent, and each individual made an independent decision to escape or not. Second, individuals received social information on the decisions of their group members, after which individuals decided again. This social information, however, did not represent their own decisions but consisted of responses that either came from a high-performing group (i.e., many individuals detecting the predator) or from a low-performing group (i.e., few individuals detecting the predator). We found that individuals adaptively adjust their quorum threshold to the quality of the social information: when receiving social information from high-performing groups, individuals employed higher quorum thresholds than when receiving information from low-performing groups. Our study demonstrates that humans can quickly evaluate the quality of publicly available information and adaptively adjust their decision rules.     

The need for formal reflexivity in conservation science

Conservation issues are often complicated by sociopolitical controversies that reflect competing philosophies and values regarding natural systems, animals, and people. Effective conservation outcomes require managers to engage myriad influences (social, cultural, political, and economic, as well as ecological). The contribution of conservation scientists who generate the information on which solutions rely is constrained if they are unable to acknowledge how personal values and disciplinary paradigms influence their research and conclusions. Conservation challenges involving controversial species provide an opportunity to reflect on the paradigms and value systems that underpin the discipline and practice of conservation science. Recent analyses highlight the ongoing reliance on normative values in conservation. We frame our discussion around controversies over feral horses (Equus ferus caballus) in the Canadian West and New Zealand and suggest that a lack of transparency and reflexivity regarding normative values continues to prevent conservation practitioners from finding resilient conservation solutions. We suggest that growing scrutiny and backlash to many normative conservation objectives necessitates formal reflexivity methods in conservation biology research, similar to those required of researchers in social science disciplines. Moreover, given that much conservation research and action continues to prioritize Western normative values regarding nature and conservation, we suggest that adopting reflexive methods more broadly is an important step toward more socially just research and practice. Formalizing such methods and requiring reflexivity in research will not only encourage reflection on how personal and disciplinary value systems influence conservation work but could more effectively engage people with diverse perspectives and values in conservation and encourage more novel and resilient conservation outcomes, particularly when dealing with controversial species.     

Tutor choice and imitation accuracy during song learning in a wild population of the Puget Sound white-crowned sparrow

Songbirds learn to sing by modeling their songs on the songs of other males through a process of social learning. Models of social learning predict that animals should be selective in what and when they learn. In this study, we asked whether young males in a wild population of the Puget Sound white-crowned sparrow, Zonotrichia leucophrys pugetensis, were selective in their choice of tutor models and what factors influenced how accurately they imitated tutors’ songs. We first examined two strategies for tutor choice: whether pupils have a conformity bias and/or a preference for high-quality tutors. In keeping with a conformity bias, tutors that sang song types that were relatively common within a radius of about 500 m of their territory were more likely to be imitated than were tutors that sang rarer song types. Most potential tutors were not imitated by pupils. Aspects of tutor quality, such as age, pairing status, and survival to the next year had no effect on whether a tutor’s song was imitated. Secondly, we tested whether pupil repertoire size, pupil quality, and local abundance of tutor models affected the accuracy of song imitations. We found a trade-off between repertoire size and tutor imitation accuracy with males that sang two or more song types developing significantly poorer imitations than males that sang one type. We discuss possible functions of a conformity learning strategy and factors that could produce a trade-off between imitation accuracy and repertoire size.     

Determining association networks in social animals: choosing spatial–temporal criteria and sampling rates

Social Network Analysis has become an important methodological tool for advancing our understanding of human and animal group behaviour. However, researchers tend to rely on arbitrary distance and time measures when defining ‘contacts’ or ‘associations’ between individuals based on preliminary observation. Otherwise, criteria are chosen on the basis of the communication range of sensor devices (e.g. bluetooth communication ranges) or the sampling frequencies of collection devices (e.g. Global Positioning System devices). Thus, researchers lack an established protocol for determining both relevant association distances and minimum sampling rates required to accurately represent the network structure under investigation. In this paper, we demonstrate how researchers can use experimental and statistical methods to establish spatial and temporal association patterns and thus correctly characterise social networks in both time and space. To do this, we first perform a mixing experiment with Merino sheep (Ovis aries) and use a community detection algorithm that allows us to identify the spatial and temporal distance at which we can best identify clusters of previously familiar sheep. This turns out to be within 2–3 m of each other for at least 3 min. We then calculate the network graph entropy rate—a measure of ease of spreading of information (e.g. a disease) in a network—to determine the minimum sampling rate required to capture the variability observed in our sheep networks during distinct activity phases. Our results indicate the need for sampling intervals of less than a minute apart. The tools that we employ are versatile and could be applied to a wide range of species and social network datasets, thus allowing an increase in both the accuracy and efficiency of data collection when exploring spatial association patterns in gregarious species.     

Individual specific contact calls of pallid bats (<Emphasis Type="Italic">Antrozous pallidus</Emphasis>) attract conspecifics at roosting sites

Contact calls are utilized by several bird and mammal species to maintain group cohesion and coordinate group movement. From a signal design perspective, contact calls typically exhibit acoustic features that make them easily localizable and encode information about individual or group identity. Pallid bats (Antrozous pallidus) are unusual among vespertilionids in that they often emit a loud, partially audible frequency-modulated social call several times in rapid succession while in flight. This call appears to function as a contact call in that it is frequently given when bats return from foraging and perform circular flights before entering a crevice roost. However, the degree to which pallid bats respond to the calls of conspecifics and what information is provided in the call is unknown. Thus, the goal of this study was to investigate pallid bat calling behavior to determine if calls attract roostmates or elicit responses from them and provide sufficient information for individual recognition. In playback studies, we found that contact calls, elicit calls, and approaches and that free-flying bats respond more to familiar than unfamiliar calls. In addition, analysis of frequency and temporal measurements of calls collected from multiple sites and spectral cross correlation analysis of calls recorded from the same radio-tagged bats on multiple evenings revealed that the frequency pattern of contact calls is highly repeatable over time within individuals but exhibits significant differences among individuals. Thus, contact call structure appears to be unique to individuals and stable through time, which makes these calls well-suited for roostmate recognition.     

Sensory ecology in a changing world: salinity alters conspecific recognition in an amphidromous fish, Pseudomugil signifer

Group fission and fusion processes are driven by state dependence, risk and the availability of information from others. Yet the availability of information changes under different environmental conditions, thus aiding or inhibiting group formation and maintenance. Chemical cues provide information on the location of individuals and can act as a mechanism for individuals to group together, although they can be greatly affected by environmental conditions. Using a flow channel, we studied how one shoaling fish species, the Pacific blue-eye (Pseudomugil signifer), responds to conspecific chemical cues (CCCs) in different environmental conditions (salinities). This species lives in estuarine environments, ranging in salinity from fresh to fully marine. P. signifer responded to CCCs in freshwater but not in saltwater. Furthermore, P. signifer did not respond to saltwater with CCCs added from freshwater. It took significantly longer for fish in saltwater, than in freshwater, to locate and join a shoal when only CCCs from the shoal were present. Finally, fish formed more cohesive shoals in freshwater than in brackish or saltwater. These results suggest that these fish do not rely on chemical cues in saltwater to locate conspecific shoals. Furthermore, the reduced amounts of these cues in saltwater may inhibit the maintenance of tight shoal structures. We suggest that fish utilise different sensory modalities in fresh or saltwater in order to locate one another, or the social structure of these groups is fundamentally different between these two water types. The importance of this study in relation to understanding how animals utilise and change different sensory modalities in varying environmental conditions is discussed.     

Maintenance of familiarity and social bonding via communal latrine use in a solitary primate (Lepilemur leucopus)

Latrine use (i.e., the repeated use of specific defecation/urination sites) has been described for several mammals, including carnivores, ungulates, and primates. However, the functional significance of latrine use in primates has not been studied systematically yet. We, therefore, followed 14 radio-collared individuals of the pair-living white-footed sportive lemur (Lepilemur leucopus) for 1097 hours of continuous focal observations to investigate latrine distribution, seasonality of latrine use, as well as age and sex of users to test various hypotheses related to possible functions of latrine use, including territory demarcation, resource defense, signaling of reproductive state, social bonding, and mate defense. All individuals of a social unit exhibited communal use of latrines located in the core area of their territory, supporting the social boding hypothesis. Latrine use seems to facilitate familiarity and social bonding within social units via olfactory communication in this primate that lives in family units but exhibits low levels of spatial cohesion and direct social interactions. In addition, frequency of latrine visitation was higher during nights of perceived intruder pressure, supporting the mate defense hypothesis. However, animals did not react to experimentally introduced feces from neighboring or strange social units, indicating that urine may be the more important component of latrines than feces in this arboreal species. Based on a survey of latrine use and function in other mammals, we conclude that latrines facilitate communication particularly in nocturnal species with limited habitat visibility and in species where individuals are not permanently cohesive because they constitute predictable areas for information exchange.     

The link between social network density and rank-order consistency of aggressiveness in juvenile eels

The emergence of an animal’s personality is the result of interactions between genetics, environment and experience. It is known that individuals are able to modulate their behaviour according to the context or the social environment. Many studies have shown for example, that familiarity among conspecifics diminishes aggressiveness, although little is known about the underlying processes. Nevertheless, personality traits have long been determined while ignoring the social context, especially in lower vertebrates such as fish. In the present experiment, we hypothesize that group connections (network density) may be positively correlated to consistency of aggressiveness by avoiding over-aggressive acts in further encounters. To test this hypothesis, we used eels (Anguilla anguilla) as a model species and monitored both aggressiveness and sociability in 64 individuals over their first 7 months of growth from the glass eel stage. As expected, social fish were less aggressive than their non-social counterparts at all times, highlighting the existence of a behavioural syndrome in eels. Additionally, rank-order consistency of aggressiveness was higher in groups of fish with high social connectivity, compared to those in less-connected fish groups. While aggressiveness must be consistent to be considered a personality trait, our results suggest that both aggressiveness and its consistency are influenced by initial social context.     

Complete reproductive skew within white-browed sparrow weaver groups despite outbreeding opportunities for subordinates of both sexes

The distribution of reproductive success within societies is a key determinant of the outcomes of social evolution. Attempts to explain social diversity, therefore, require that we quantify reproductive skews and identify the mechanisms that generate them. Here, we address this priority using life history and genotypic data from >600 individuals in 40 wild groups of the cooperatively breeding white-browed sparrow weaver, Plocepasser mahali. We show that groups comprise up to six males and seven females, but within-group reproduction is completely monopolised by a single dominant male and female, while extra-group males sire 12–18 % of offspring. Strong within-group kin structure could frequently explain these monopolies, as subordinates had typically delayed dispersal from their natal groups and so frequently (1) lacked within-group outbreeding partners, and/or (2) stood to gain little from contesting dominant reproduction, being almost as related to the dominant’s young as they would have been to their own. Kin structure alone cannot account entirely for these monopolies, however, as they remained complete following the immigration of unrelated males and females. That subordinate females remain reproductively quiescent despite also showing comparable body condition to dominants, overlapping them substantially in age, and showing no evidence of elevated stress hormone levels raises the possibility that they exercise reproductive restraint due instead to a threat of action by dominants and/or deficits in offspring fitness that might arise if subordinates bred. Our findings highlight the complexity of the mechanisms that generate reproductive disparities in animal societies and the challenge of identifying them when skews are complete.     

Sequence of quorums during collective decision making in macaques

Synchronization of activity is one of the major challenges of any society, and to what extent social animals reach a consensus still remains to be established. In the case of group movements, recent studies have underlined the importance of the pre-departure period and suggested that some individuals in a group express their motivation to move by showing a preference for a specific direction. However, how do other group members really choose the time and direction of movement? This study shows that in two semi-free ranging Tonkean macaque (Macaca tonkeana) groups, several individuals propose different directions for movement by displaying unique behavior. The whole group eventually moves in the choice of direction supported by the majority of individuals according to a sequence of three quorum rules. Moreover, when the number of individuals choosing another direction is higher than their own group, individuals that proposed alternative directions eventually renounce and follow the majority. Despite conflict of interests, group members reach a consensus before the actual start of group movement. This demonstrates that processes of this type, which can be considered to be voting processes, are not exclusive to human societies and may be explained by a complex sequence of simple rules.     

From eavesdropping on performance to copying the behavior of others: a review of public information use

Public information (PI), a form of indirect social information, is used by individuals to estimate the quality of environmental parameters. It can be acquired in two ways. One way is by noting the performance of others. The other way it can be acquired is by noting the behavioral decisions of other individuals. Performance-based PI has been observed most often in the context of food and breeding patch estimation, as well as by individuals eavesdropping on contests between others in the context of assessing the fighting ability of opponents and the quality of mates. Evidence for the acquisition of PI from behavioral decisions of others comes mostly from studies of copying behavior, although recent work suggests that it also occurs when individuals estimate the tendency of others to be altruistic. PI use appears to be widespread across many taxa, although most work has been conducted on birds and fish. Absent from the literature are clear examples of PI use in mammals. The use of PI appears to often depend on its cost of acquisition and whether it contradicts an individual’s personal prior experience. PI can be an important benefit of associating with others.     

The effect of dominance hierarchy on the use of alternative foraging tactics: a phenotype-limited producing-scrounging game

Group living is thought to be advantageous for animals, though it also creates opportunities for exploitation. Using food discovered by others can be described as a producer-scrounger, frequency-dependent game. In the game, scroungers (parasitic individuals) do better than producers (food finders) when scroungers are rare in the group, but they do worse when scroungers are common. When the individuals' payoffs do not depend on their phenotype (i.e. a symmetric game), this strong negative frequency dependence leads to a mixed stable solution where both alternatives obtain equal payoffs. Here, we address the question of how differences in social status in a dominance hierarchy influence the individuals' decision to play producer or scrounger in small foraging groups. We model explicitly the food intake rate of each individual in a dominance-structured foraging group, then calculate the Nash equilibrium for them. Our model predicts that only strong differences in competitive ability will influence the use of producing or scrounging tactics in small foraging groups; dominants will mainly play scrounger and subordinates will mostly use producer. Since the differences in competitive ability of different-ranking individuals likely depend on the economic defendability of food, our model provides a step towards the integration of social foraging and resource defence theories. Received: 30 July 1997 / Accepted after revision: 15 November 1997     

Increased behavioural lateralization in parasitized coral reef fish

Preferential use of one side of the body for cognitive or behavioural tasks (lateralization) is common in many animals, including humans. However, few studies have demonstrated whether lateralization is phenotypically plastic, and varies depending on the ecological context. We studied lateralization (measured as a turning preference) in the bridled monocle bream (Scolopsis bilineatus). This coral reef fish is commonly infected by a large, ectoparasitic isopod (Anilocra nemipteri) that attaches to the left or right side of its host’s head. Fish that were parasitized showed no turning bias with respect to the side on which the parasite had attached. On average, however, parasitised fish were significantly more lateralized (i.e. had a strong side bias) than unparasitized fish. The extent of lateralization declined significantly when we experimentally removed the parasite. Our results indicate that lateralization can vary with the ecological context. One possible explanation is that lateralization shortens the response time until fish flee after encountering a predator. A stronger side bias might be advantageous for parasitized individuals to overcome their recently documented lower maximum swimming speed.     

Tactics, effectiveness and avoidance of mate guarding in the blue-footed booby (Sula nebouxii)

We examined the dynamics and avoidance of mate guarding, by males and females, in the blue-footed booby, in which the two social mates are usually simultaneously present on the territory but each of them is unmonitored by the other for one-quarter of its time. Both sexes were promiscuous and liable to switch mates. Cuckolded individuals did not increase their overall presence on the territory, but in response to the extra-pair (EP) courtships of their mates, both sexes doubled their rate of intra-pair (IP) courtship and sometimes showed aggression. The male or female's presence depressed the social mate's EP activity, but intra-pair courtship had no such effect, tending even to propitiate that EP activity. Similarly, when females responded to their social mates' EP courtship with approach or aggression, disruption of EP activity was short-lived. Promiscuous females modified their diurnal pattern of attendance, as if attempting to sidestep monitoring by their mates, but cuckolded males matched the modification. Both sexes tended to perform their EP activities at a distance when their mates were present, possibly to evade monitoring or disruption by their mates. Male and female boobies cannot monitor their mates continuously, they do little to facultatively adjust their presence on territory to the risk of infidelity, and their immediate responses to overt infidelity have only the briefest impact; but the information they acquire while monitoring their mates may be critical to constraining their mates' infidelity and also to calibrating their own reproductive investment.