The relationship between social behaviour and habitat familiarity in African elephants (Loxodonta africana)

Social associations with conspecifics can expedite animals'' acclimation to novel environments. However, the benefits gained from sociality may change as the habitat becomes familiar. Furthermore, the particular individuals with whom animals associate upon arrival at a new place, familiar conspecifics or knowledgeable unfamiliar residents, may influence the type of information they acquire about their new home. To examine animals'' social dynamics in novel habitats, we studied the social behaviour of African elephants (Loxodonta africana) translocated into a novel environment. We found that the translocated elephants'' association with conspecifics decreased over time supporting our hypothesis that sociality provides added benefits in novel environments. In addition, we found a positive correlation between body condition and social association, suggesting that elephants gain direct benefits from sociality. Furthermore, the translocated elephants associated significantly less than expected with the local residents and more than expected with familiar, but not necessarily genetically related, translocated elephants. The social segregation between the translocated and resident elephants declined over time, suggesting that elephants can integrate into an existing social setting. Knowledge of the relationship between sociality and habitat familiarity is highly important in our constantly changing world to both conservation practice and our understanding of animals'' behaviour in novel environments.     

Space use and social association in a gregarious ungulate: Testing the conspecific attraction and resource dispersion hypotheses

Animals use a variety of proximate cues to assess habitat quality when resources vary spatiotemporally. Two nonmutually exclusive strategies to assess habitat quality involve either direct assessment of landscape features or observation of social cues from conspecifics as a form of information transfer about forage resources. The conspecific attraction hypothesis proposes that individual space use is dependent on the distribution of conspecifics rather than the location of resource patches, whereas the resource dispersion hypothesis proposes that individual space use and social association are driven by the abundance and distribution of resources. We tested the conspecific attraction and the resource dispersion hypotheses as two nonmutually exclusive hypotheses explaining social association and of adult female caribou (Rangifer tarandus). We used location data from GPS collars to estimate interannual site fidelity and networks representing home range overlap and social associations among individual caribou. We found that home range overlap and social associations were correlated with resource distribution in summer and conspecific attraction in winter. In summer, when resources were distributed relatively homogeneously, interannual site fidelity was high and home range overlap and social associations were low. Conversely, in winter when resources were distributed relatively heterogeneously, interannual site fidelity was low and home range overlap and social associations were high. As access to resources changes across seasons, caribou appear to alter social behavior and space use. In summer, caribou may use cues associated with the distribution of forage, and in winter caribou may use cues from conspecifics to access forage. Our results have broad implications for our understanding of caribou socioecology, suggesting that caribou use season‐specific strategies to locate forage. Caribou populations continue to decline globally, and our finding that conspecific attraction is likely related to access to forage suggests that further fragmentation of caribou habitat could limit social association among caribou, particularly in winter when access to resources may be limited.     

Local enhancement promotes cockroach feeding aggregations

Communication and learning from each other are part of the success of animal societies. Social insects invest considerable effort into signalling to their nestmates the locations of the most profitable resources in their environment. Growing evidence also indicates that insects glean such information through cues inadvertently provided by their conspecifics. Here, we investigate social information use in the foraging decisions by gregarious cockroaches (Blattella germanica L.). Individual cockroaches given a simultaneous choice in a Y-olfactometer between the odour of feeding conspecifics and the mixed odour of food plus non-feeding conspecifics showed a preference for the arm scented with the odour of feeding conspecifics. Social information (the presence of feeding conspecifics) was produced by cockroaches of all age classes and perceived at short distance in the olfactometer arms, suggesting the use of inadvertently provided cues rather than signals. We discuss the nature of these cues and the role of local enhancement (the selection of a location based on cues associated with the presence of conspecifics) in the formation of feeding aggregations in B. germanica. Similar cue-mediated recruitments could underpin a wide range of collective behaviours in group-living insects.     

Visual Cues Given by Humans Are Not Sufficient for Asian Elephants (Elephas maximus) to Find Hidden Food

Recent research suggests that domesticated species – due to artificial selection by humans for specific, preferred behavioral traits – are better than wild animals at responding to visual cues given by humans about the location of hidden food. \Although this seems to be supported by studies on a range of domesticated (including dogs, goats and horses) and wild (including wolves and chimpanzees) animals, there is also evidence that exposure to humans positively influences the ability of both wild and domesticated animals to follow these same cues. Here, we test the performance of Asian elephants (Elephas maximus) on an object choice task that provides them with visual-only cues given by humans about the location of hidden food. Captive elephants are interesting candidates for investigating how both domestication and human exposure may impact cue-following as they represent a non-domesticated species with almost constant human interaction. As a group, the elephants (n = 7) in our study were unable to follow pointing, body orientation or a combination of both as honest signals of food location. They were, however, able to follow vocal commands with which they were already familiar in a novel context, suggesting the elephants are able to follow cues if they are sufficiently salient. Although the elephants’ inability to follow the visual cues provides partial support for the domestication hypothesis, an alternative explanation is that elephants may rely more heavily on other sensory modalities, specifically olfaction and audition. Further research will be needed to rule out this alternative explanation.     

Perception of infection: disease-related social cues influence immunity in songbirds

While avoidance of sick conspecifics is common among animals, little is known about how detecting diseased conspecifics influences an organism''s physiological state, despite its implications for disease transmission dynamics. The avian pathogen Mycoplasma gallisepticum (MG) causes obvious visual signs of infection in domestic canaries (Serinus canaria domestica), including lethargy and conjunctivitis, making this system a useful tool for investigating how the perception of cues from sick individuals shapes immunity in healthy individuals. We tested whether disease-related social information can stimulate immune responses in canaries housed in visual contact with either healthy or MG-infected conspecifics. We found higher complement activity and higher heterophil counts in healthy birds viewing MG-infected individuals around 6–12 days post-inoculation, which corresponded with the greatest degree of disease pathology in infected stimulus birds. However, we did not detect the effects of disease-related social cues on the expression of two proinflammatory cytokines in the blood. These data indicate that social cues of infection can alter immune responses in healthy individuals and suggest that public information about the disease can shape how individuals respond to infection.     

Indirect Information Transfer: Three‐Spined Sticklebacks Use Visual Alarm Cues From Frightened Conspecifics About an Unseen Predator

Animals can attempt to reduce uncertainty about their environment by gathering information personally or by observing others' interactions with the environment. There are several sensory modalities that can be used to transmit social information from chemical to visual to audible cues. When predation risk is variable, visual cues of conspecific behavior might be especially telling about the presence of a potential threat; however, most studies couple visual and chemical cues together. Here, we tested whether visual behavioral cues from frightened conspecifics were sufficient to indirectly transfer information about the presence of an unseen predator in three‐spined sticklebacks. Our results demonstrate that visual behavioral cues from conspecifics about the presence of a predator are sufficient to induce an antipredator response. This suggests that information transfer can occur rapidly in the absence of chemical cues and that some individuals weigh social information more heavily than others.     

Conspecific and heterospecific information use in bumblebees

Heterospecific social learning has been understudied in comparison to interactions between members of the same species. However, the learning mechanisms behind such information use can allow animals to be flexible in the cues that are used. This raises the question of whether conspecific cues are inherently more influential than cues provided by heterospecifics, or whether animals can simply use any cue that predicts fitness enhancing conditions, including those provided by heterospecifics. To determine how freely social information travels across species boundaries, we trained bumblebees (Bombus terrestris) to learn to use cues provided by conspecifics and heterospecific honey bees (Apis mellifera) to locate valuable floral resources. We found that heterospecific demonstrators did not differ from conspecifics in the extent to which they guided observers'' choices, whereas various types of inorganic visual cues were consistently less effective than conspecifics. This was also true in a transfer test where bees were confronted with a novel flower type. However, in the transfer test, conspecifics were slightly more effective than heterospecific demonstrators. We then repeated the experiment with entirely naïve bees that had never foraged alongside conspecifics before. In this case, heterospecific demonstrators were equally efficient as conspecifics both in the initial learning task and the transfer test. Our findings demonstrate that social learning is not a unique process limited to conspecifics and that through associative learning, interspecifically sourced information can be just as valuable as that provided by conspecific individuals. Furthermore the results of this study highlight potential implications for understanding competition within natural pollinator communities.     

Unusually extensive networks of vocal recognition in African elephants

Research on acoustic communication has often focused on signalling between territorial individuals or static neighbouring groups. Under these circumstances, receivers have the opportunity to learn to recognize the signals only of the limited number of conspecifics with which they are in auditory contact. In some mammals, however, social units move freely with respect to one another and range widely, providing individuals with opportunities to learn to recognize the signals of a wide range of conspecifics in addition to those of their immediate neighbours. We conducted playback experiments on African elephants, Loxodonta africana, in Amboseli National Park, Kenya, to determine the extent to which adult female elephants, which have a highly fluid social system, can recognize others in the population through infrasonic contact calls. Female elephants could distinguish the calls of female family and bond group members from those of females outside of these categories; moreover, they could also discriminate between the calls of family units further removed than bond group members, on the basis of how frequently they encountered them. We estimated that subjects would have to be familiar with the contact calls of a mean of 14 families in the population (containing around 100 adult females in total), in order to perform these discriminations. Female elephants thus appear to have unusually extensive networks of vocal recognition, which may prove to be typical of long-lived species that have both fluid social systems and the means for long-distance vocal communication. Copyright 2000 The Association for the Study of Animal Behaviour.     

Female German cockroaches join conspecific groups based on the incidence of auditory cues

Insects deciding whether to approach and join a group of conspecifics may utilize olfactory and auditory signals, or cues, from that group as indicators of its size or the suitability of its shelter. Here we show (1) that German cockroaches, Blattella germanica L. (Dictyoptera: Blattellidae), in groups wing‐fan; (2) that the incidence of wing‐fanning (WF) increases proportionately to group size; and (3) that gravid females utilize auditory cues associated with WF behaviour when they decide whether or not to enter a shelter. In binary choice arena bioassays, proportionately more gravid females avoided shelters associated with play‐back of high‐incidence male‐ or female‐produced WF sound, indicating a high‐density group of conspecifics, but sought shelters associated with play‐back of low‐incidence WF sound, indicating a low‐density group of conspecifics. These auditory cues seem to convey information on group size or density, avoid sensory fatigue in enclosed environments, and allow sonotactic orientation to the group's location.     

Can a solitary avian species use collective detection? An assay in semi-natural conditions

Collective detection (e.g., enhanced predator detection through the vigilance of conspecifics) is expected to have evolved particularly in social species. However, we assessed the degree to which an avian territorial species (California towhee Pipilo crissalis) would use social cues about predation in a semi-natural assay. We also exposed a social species (house finch Carpodacus mexicanus) to similar conditions. California towhees increased scanning rates when foraging with conspecifics, whereas house finches increased scanning rates when foraging solitarily, suggesting that vigilance in these species is regulated mostly through interference competition and through predation risk, respectively. California towhees did not show early detection, and actually the last detector in the group delayed detection in relation to solitary individuals. House finches benefited from early detection, but the second and last detectors maintained detection at the level of solitary individuals. California towhees increased the chances of fleeing when in groups in relation to solitary conditions, but this effect was less pronounced in the last detector. House finches always fled across conditions. Overall, an asocial avian species may use collective detection, but limited to certain types of cues: responses were more pronounced to overt (conspecifics walking or fleeing) rather than subtle (conspecifics becoming alert or crouching) social cues.     

Learned Recognition of Intraspecific Predators in Larval Long-Toed Salamanders Ambystoma macrodactylum

The ability of prey to detect predators and respond accordingly is critical to their survival. The use of chemical cues by animals in predator detection has been widely documented. In many cases, predator recognition is facilitated by the release of alarm cues from conspecific victims. Alarm cues elicit anti‐predator behavior in many species, which can reduce their risk of being attacked. It has been previously demonstrated that adult long‐toed salamanders, Ambystoma macrodactylum, exhibit an alarm response to chemical cues from injured conspecifics. However, whether this response exists in the larval stage of this species and whether it is an innate or a learned condition is unknown. In the current study, we examined the alarm response of naïve (i.e. lab‐reared) larval long‐toed salamanders. We conducted a series of behavioral trials during which we quantified the level of activity and spatial avoidance of hungry and satiated focal larvae to water conditioned by an injured conspecific, a cannibal that had recently been fed a conspecific or a non‐cannibal that was recently fed a diet of Tubifex worms. Focal larvae neither reduced their activity nor spatially avoided the area of the stimulus in either treatment when satiated, and exhibited increased activity towards the cannibal stimulus when hungry. We regard this latter behavior as a feeding response. Together these results suggest that an anti‐predator response to injured conspecifics and to cannibalistic conspecifics is absent in naïve larvae. Previous studies have shown that experienced wild captured salamanders do show a response to cannibalistic conspecifics. Therefore, we conducted an additional experiment examining whether larvae can learn to exhibit anti‐predator behavior in response to cues from cannibalized conspecifics. We exposed larvae to visual, chemical and tactile cues of stimulus animals that were actively foraging on conspecifics (experienced) or a diet of Tubifex (naïve treatment). In subsequent behavioral treatments, experienced larvae significantly reduced their activity compared to naive larvae in response to chemical cues of cannibals that had recently consumed conspecifics. We suggest that this behavior is a response to alarm cues released by consumed conspecifics that may have labeled the cannibal. Furthermore, over time, interactions with cannibals may cause potential prey larvae to learn to avoid cannibals regardless of their recent diet.     

The role of biotic and abiotic cues in stimulating aggregation by larval cane toads (Rhinella marina)

Tadpoles of the cane toad (Rhinella marina) form dense aggregations in the field, but the proximate cues eliciting this behavior are not well understood. We sampled water‐bodies in the Northern Territory of Australia, finding that the density of cane toad tadpoles increased with increasing temperature. Furthermore, we conducted laboratory experiments to explore the roles of biotic factors (attraction to conspecifics; chemical cues from an injured conspecific; food) and spatially heterogeneous abiotic factors (light levels, water depth, physical structure) to identify the cues that induce tadpole aggregation. Annulus and binary choice trials demonstrated weak but significant attraction between conspecifics. Tadpoles decreased swimming speeds, but did not increase grouping in response to cues from an injured conspecific. Larvae aggregated in response to abiotic cues (high levels of illumination and proximity to physical structures) and were strongly attracted to feeding conspecifics. Overall, aggregation by cane toad tadpoles is likely driven by weak social attraction coupled with a shared preference for specific abiotic features, creating loose aggregations that are then reinforced by movement toward feeding conspecifics.     

Impaired social contacts with familiar anesthetized conspecific in CA3‐restricted brain‐derived neurotrophic factor knockout mice

Familiarity is conveyed by social cues and determines behaviors toward conspecifics. Here, we characterize a novel assay for social behaviors in mice—contacts with anesthetized conspecific—which eliminates reciprocal interactions, including intermale aggression and shows behaviors that are independent of the demonstrator's activity. During the initial 10 minutes (phase‐1), the wild‐type (WT) subjects contacted the anesthetized conspecifics vigorously regardless of familiarity. During the subsequent 80 minutes (phase‐2), however, they contacted more with familiar than unfamiliar conspecifics. We then applied this test to highly aggressive mice with a hippocampal CA3‐restricted knockout (KO) of brain‐derived neurotrophic factor (BDNF), in which aggression may mask other social behaviors. The KO mice showed less preference for contacting familiar conspecifics than did WT mice during phase‐2 but no differences during phase‐1. Among nonsocial behaviors, WT mice also spent less time eating in the presence of familiar than with unfamiliar conspecifics, which was not seen in KO mice. In addition, KO mice exhibited reduced pain sensitization. Altogether, these findings suggest that CA3‐specific deletion of BDNF results in deficits in circuits that process social cues from familiar conspecifics as well as pain and may underlie empathy‐like behaviors.     

Social Context Influences the Antipredator Behaviour of Fathead Minnows to Chemical Alarm Cues

Prey fishes, like many organisms under fluctuating predation threat, rely on multiple sources of information to accurately gauge current risk. This includes the use of chemical cues such as alarm cues from damaged conspecifics or familiar heterospecifics, as well as the odour of known predators. While each fish is well equipped with its own array of sensory abilities, they should also be alert to the behaviours of nearby neighbours who may have information they lack. In the present study, we tested the ability of fathead minnows to use social cues in combination with the odour of damaged conspecifics and heterospecifics to mediate the assessment of predation risk. Specifically, we tested whether the presence of a shoal of conspecifics or familiar heterospecifics would significantly change a minnow's antipredator behaviour when exposed to the odour of a damage‐release cue from a conspecific or ecologically similar heterospecific. The results of our study showed a significant interaction between the damage‐release cues to which the minnows were exposed and the presence/absence of shoalmates. These findings have important implications for the design of future investigations of antipredator responses because most studies of group‐living prey have been conducted on solitary subjects.     

In Space and Time: Territorial Animals are Attracted to Conspecific Chemical Cues

Territorial animals lay scent marks around their territories to broadcast their presence, but these olfactory signals can both attract and repel conspecifics. Attraction or aversion can have a profound impact in terms of space use and thereby influence an individual's access to resources and mates. Here, we test the impact of chemical signals on the long‐term space use and activity of receivers, comparing the response of males and females, territory holders, and temporary visitors in Sceloporus undulatus lizards in the field. We placed either male femoral gland secretions (chemical) or blank (control) cues on resident male landmarks, repeatedly over 5 d, while monitoring the activity and location of all lizards in the vicinity. We found that resident males and females, but not non‐resident males, were active on more days near landmarks treated with chemical cues than landmarks treated with control cues. Non‐resident males remained closer to chemical than control cues. These results suggest that territorial scent marks are attractive to conspecifics and impact space use, but that the specific effects depend on receiver sex and residency status. Such subtle or gradual changes in behavior may frequently be overlooked by short‐term choice experiments. Future studies investigating the behavioral significance of a communicative signal should consider these finer details of behavior for a more comprehensive assessment.     

The Effects of Density on the Learned Recognition of Heterospecific Alarm Cues

Numerous species, both aquatic and terrestrial, use alarm cues to mediate predation risk. These cues may be either intentionally or inadvertently released, and may be received by either conspecifics or heterospecifics. In aquatic systems, alarm cues are often chemical in nature and are released when an organism is disturbed or damaged by a predator. In some cases the recognition of alarm cues from conspecifics, or closely related heterospecifics, is innate, while the recognition of alarm cues from distantly related species must be learned. Many studies have documented the use of heterospecific alarm cues, but few have explored the manner in which these cues come to be recognized as an indication of predation. In the current study, we examined the fathead minnow (Pimephales promelas)/brook stickleback (Culaea inconstans) alarm system. We tested the effect of density on the ability of minnows to learn to recognize stickleback alarm cues as a threat. We hypothesized that the ability of minnows to learn to recognize stickleback alarm cues should increase with increasing stickleback density because there would be more opportunity for minnows to associate the heterospecific alarm cue with the threat. To test this hypothesis we stocked minnows into large outdoor pools with no stickleback, low numbers of stickleback, or high numbers of stickleback. All pools contained a predator (pike, Esox lucius) known to the minnows. Following a 14 d conditioning period, minnows were tested for a response to skin extract from stickleback, minnow, and an unknown heterospecific (swordtail, Xiphophorus helleri). Minnows from pools with large numbers of stickleback learned to respond to stickleback alarm cues while minnows from pools with low numbers of stickleback, or no stickleback, did not.     

Oestrous state dynamics in chemical communication by captive female Asian elephants

In many mammals, reproductive status is revealed through chemical cues in urine. The reproductive status of receivers may influence their interest in such signals. For social mammals that live in matrilineal groups, females may benefit by detecting the reproductive condition of herdmates. Responses to urine during oestrous cycles of senders and receivers are potential indicators of signal functions. We examined the chemosensory responses, first by four captive female Asian elephants,Elephas maximus , over their oestrous cycles to familiar follicular and luteal phase urine and second by 14 different female Asian elephants to unfamiliar conspecific follicular and luteal phase urine. We asked whether females could distinguish the reproductive state of another female as measured by their differential response to luteal- and follicular-phase urine. We further examined the influence of the receiver's reproductive status on response levels. Females responded more with specific tactolfactory trunk behaviours to follicular- than to luteal-phase urine, but only when the receiving female was in her follicular phase. Like their male conspecifics, Asian elephant females can detect changes in the reproductive state of conspecifics. The functional significance of this ability has yet to be determined but may be related more to the resource holding power of females in follicular phase than to a means for females to synchronize oestrous cycles. Such female-female communication may have important effects on social group dynamics. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Do anuran larvae respond behaviourally to chemical cues from an invasive crayfish predator? A community-wide study

Antipredator behaviour is an important fitness component in most animals. A co-evolutionary history between predator and prey is important for prey to respond adaptively to predation threats. When non-native predator species invade new areas, native prey may not recognise them or may lack effective antipredator defences. However, responses to novel predators can be facilitated by chemical cues from the predators’ diet. The red swamp crayfish Procambarus clarkii is a widespread invasive predator in the Southwest of the Iberian Peninsula, where it preys upon native anuran tadpoles. In a laboratory experiment we studied behavioural antipredator defences (alterations in activity level and spatial avoidance of predator) of nine anurans in response to P. clarkii chemical cues, and compared them with the defences towards a native predator, the larval dragonfly Aeshna sp. To investigate how chemical cues from consumed conspecifics shape the responses, we raised tadpoles with either a tadpole-fed or starved crayfish, or dragonfly larva, or in the absence of a predator. Five species significantly altered their behaviour in the presence of crayfish, and this was largely mediated by chemical cues from consumed conspecifics. In the presence of dragonflies, most species exhibited behavioural defences and often these did not require the presence of cues from predation events. Responding to cues from consumed conspecifics seems to be a critical factor in facilitating certain behavioural responses to novel exotic predators. This finding can be useful for predicting antipredator responses to invasive predators and help directing conservation efforts to the species at highest risk.     

Scents and scents-ability: pollution disrupts chemical social recognition and shoaling in fish

Chemical cues are of enormous importance in mediating the behaviour of animals, enabling them to navigate throughout their habitats, to detect the presence of predators or prey and for social recognition-identifying and discriminating between conspecifics. In many species of freshwater fish, social recognition is known to be based primarily on chemical cues. Such recognition mechanisms are vulnerable to disruption by the presence of anthropogenic contaminants in the aquatic environment. Here we show that acute exposure to low, environmentally relevant dosages of the ubiquitous contaminant, 4-nonylphenol, can seriously affect social recognition and ultimately social organization in fishes. A 1 hour 0.5 microgl-1 dose was sufficient to alter the response of members of a shoaling fish species (juvenile banded killifish, Fundulus diaphanus) to conspecific chemical cues. Dosages of 1-2 microgl-1 caused killifish to orient away from dosed conspecifics, in both a flow channel and an arena. Given the overall importance of shoaling as an adaptive strategy against predators and for locating food, it is likely that its disruption by anthropogenic contaminants would have serious implications for fishes' fitness.     

The Costs of Conspecifics: Are Social Distractions or Environmental Distractions More Salient?

Animals have limited attention that predisposes them to distraction, but the impact of different types of distraction is relatively unknown. We first conducted a discrimination experiment to ensure brown anoles (Anolis sagrei) responded appropriately to model predators (a rubber snake) and model conspecifics. We found anoles responded to the snake by suppressing looking and increasing locomotion, a marked difference than their response when presented with a conspecific. Next, we designed a prime and probe experiment to test the salience of social and environmental distractions on brown anoles. The social distraction consisted of a conspecific exemplar that was presented to the focal individual. Environmental distractions were vegetation that was moved at different speeds (ambient or vigorous) near the focal individual. Following 30 seconds of distraction with one of the three treatments, we presented the model snake, which was initially moved to within 1 m of a subject, and then moved closer until the subject looked in response to the approaching threat, and then fled. There was no effect of distractor on alert distance, but anoles distracted with the conspecific tolerated a closer approach with the model predator before fleeing than they did to either of the vegetative movements. These results cannot be explained by three other models of risk assessment and suggest that social cues distracted brown anoles more than environmental cues. These results may be generalizable to other social species that must simultaneously monitor conspecifics and assess predation risk.