Effects of Group Size on Approach to Novel Objects in Ravens (Corvus corax)

Neophobia may constrain explorative behaviour, learning and innovation, while social context may facilitate approach to novel objects and acceptance of novel food. We examined the effects of neophobia on the exploration of novel objects in relation to social context in ravens (Corvus corax). Ravens are suitable subjects for studying effects of social context, as they are highly neophobic scavengers that recruit conspecifics to food. We tested two groups of six and 11 hand‐raised birds in three conditions: single‐bird, in dyadic combinations and sibling sub‐groups of three to six birds. Contrary to expectation, individuals of both groups were quicker to approach novel objects when tested alone than when tested with conspecifics. However, they spent more time close to and manipulating the novel objects in the social conditions (dyadic and group) than when being alone. We discuss the possibility that the higher latencies of dyads and groups to approach novel objects may reflect a ‘negotiation’ process in a ‘war of attrition’ between the individuals over risk‐taking.     

Impact of Environmental Disturbance on the Stability and Benefits of Individual Status within Dominance Hierarchies

Changes in environmental conditions affect social interactions and thus may modify an individual's competitive ability within a social group. We subjected three‐spined sticklebacks, Gasterosteus aculeatus, housed in groups of four individuals, to environmental perturbations to assess the impact on dominance hierarchy stability. Hierarchy stability decreased during increased turbulence or lowered water levels (‘simulated drought’) whereas control hierarchies became more stable in a constant environment. The dominant individual either became more aggressive and remained dominant during the environmental manipulation or was usurped by a lower rank member. Only simulated drought affected rates of aggression where levels of aggression were higher after the water level was dropped which may be the result of an increased encounter rate in these conditions. When there were large size differences between the group members, the dominant individual performed the greatest amount of aggression and ate the largest proportion of food and there was little aggressive behaviour from the lower ranks. In groups of similar‐sized individuals, aggression was much higher. The benefit of being dominant was to gain weight over the experimental period whereas ranks 2 and 3 lost weight. The lowest rank, 4, actually gained weight over the experimental period. This study suggests that it would benefit an individual to be dominant, highly aggressive and gain weight or be submissive, avoid aggressive interactions and, by sneakily obtaining access to food, also gain weight. Altering environmental conditions has a profound effect on social behaviour in this study.     

Social Learning in the Caracara Chimango,Milvago chimango (Aves: Falconiformes): an Age Comparison

Milvago chimango is a gregarious raptor showing great ecological plasticity. Their ability to explore new resources has allowed them to survive in areas with increasing human modification. In this study, we evaluated the social learning ability in wild‐caught individuals of M. chimango. In particular, we tested whether an ‘observer’ individual could improve the acquisition of a novel behaviour by watching a ‘demonstrator,’ and we examined the effects of age of both observers and demonstrators on social learning. We measured the ability of 18 observers to open an opaque Plexiglas box containing food, and we compared their performance to that of 10 control birds who did not watch a demonstrator solve the task. Prior to watching a demonstrator, only two of the observers and two of the control birds were able to open the box. After watching a demonstrator, 67% of observers were able to open the box, outperforming control birds in speed and success. Juvenile observers were more successful and faster than adults at contacting and opening the box. The age of the demonstrator did not influence the observers’ likelihood of success. These results showed that M. chimango are able to learn a box‐opening task with a hidden food reward by observing the behaviour of a conspecific and that this behaviour persisted over several days. Social learning ability in M. chimango might allow certain behavioural patterns, such as those related to novel resource acquisition in modified environments, to be socially transmitted among individuals in a population.     

Space competition in the acorn woodpecker: Power struggles in a cooperative breeder

During a 5-year study of the cooperatively living acorn woodpecker Melanerpes formicivorus, I observed 26 irruptions of agonistic behaviour involving large numbers of intruders invading a territory. Evidence from several sources suggests that these invasions are usually contests among potential immigrants for the opportunity to replace a missing group member (hence, ‘power struggles’). Thus, power struggles indicate intense competition among individuals to transfer out of their natal group. Such dispersal is not necessarily excluded by the hypothesis that individuals gain directly by living in cooperative groups through either increased foraging efficiency or better predator defence. However, birds involved in power struggles expend considerable energy and incur considerable risks. These costs are likely to be an acceptable part of dispersal only if cooperative living is a result of birds being forced to remain in groups as a result of resource localization.     

The influence of flock size and geometry on the scanning behaviour of spotted turtle doves,Streptopelia chinensis

Abstract The negative correlation between the time individuals spend scanning the environment for predators and group size is usually explained by the benefit of corporate vigilance. However, this negative correlation may be explained additionally in terms of the ‘dilution effect’ and ‘selfish herd geometry’. Our experimental investigation of the scanning behaviour of free-living spotted turtle doves foraging at different shaped feeders revealed that flock geometry influenced individual scanning rates. The time spent scanning declined with group size less rapidly among birds foraging in linear flocks than among those foraging in more two-dimensional flocks. These results were not confounded by aggressive behaviour, and indicate that the benefits of foraging in groups include the so-called selfish herd geometry.     

Individuality in Problem Solving: String Pulling in Two Carduelis Species (Aves: Passeriformes)

The research reported here was designed to study the individual peculiarities of birds in solving a problem. Goldfinches Carduelis carduelis and siskins C. spinus were tested with the string‐pulling task: sitting on a perch from which a small food container is suspended by a string the test bird had to lift the container, using the bill to pull the string stepwise up and a foot to hold it, and repeat that until they could reach the food. Fifty‐two goldfinches and 29 siskins raised under controlled conditions were tested individually. Three groups became apparent: ‘inventors’ (23% of goldfinches; 62% of siskins) solved the problem by themselves; ‘imitators’ (25% of goldfinches; 10% of siskins) succeeded after seeing a performing conspecific; ‘duffers’ (52% of goldfinches, 28% of siskins) did not succeed either way. The species – but not the sexes – differed significantly in string‐pulling ability. The results of our experiments indicate that string pulling is an acquired combination of innate behaviour elements. An individual's string‐pulling competence may depend on prior experience of handling branchlets, on trial‐and‐error learning and on social learning (emulation). However, some individuals succeeded without these facilitating factors, while others did not succeed at all despite all of them present. Although functionally and motivationally related to feeding, the learned string pulling is often shown as a playful activity without an obvious reward.     

Individual differences in the use of social information in foraging by captive great tits

We investigated individual differences in copying behaviour of captive great tits, Parus major, by analysing their response to a tutor indicating a new feeding site. We used two groups, each of seven male birds, labelled 'fast' and 'slow' explorers based on previous studies in which consistent individual differences in the speed of exploring were found. The birds were trained to search for food hidden in a number of differently coloured and shaped feeders, and later to search in only one type of feeder. During the tests, food was absent and the birds were observed in two different situations: alone or in the presence of a tutor, a bird that had been trained to feed in a different kind of feeder. When alone, slow birds readily extended their search to other feeders while fast birds did not change their routine of visiting the previously rewarded ones. In the presence of the tutor, the opposite occurred: slow birds did not change their behaviour while fast birds significantly increased their visits to the feeders indicated by the tutor. Fast and slow individuals thus differ in their foraging and copying behaviour, consistent with the producer-scrounger model. Copyright 2000 The Association for the Study of Animal Behaviour.     

The effects of spatial food distribution and group size on foraging behaviour in a benthic fish

Animals foraging in heterogeneous environments benefit from information on local resource density because it allows allocation of foraging effort to rich patches. In foraging groups, this information may be obtained by individuals through sampling or by observing the foraging behaviour of group members. We studied the foraging behaviour of goldfish (Carassius auratus) groups feeding in pools on resources distributed in patches. First, we determined if goldfish use sampling information to distinguish between patches of different qualities, and if this allowed goldfish to benefit from a heterogeneous resource distribution. Then, we tested if group size affected the time dedicated to food searching and ultimately foraging success. The decision of goldfish to leave a patch was affected by whether or not they found food, indicating that goldfish use an assessment rule. Giving-up density was higher when resources were highly heterogeneous, but overall gain was not affected by resource distribution. We did not observe any foraging benefits of larger groups, which indicate that grouping behaviour was driven by risk dilution. In larger groups the proportion searching for food was lower, which suggests interactions among group members. We conclude that competition between group members affects individual investments in food searching by introducing the possibility for alternative strategies, such as scrounging or resource monopolisation.     

The Social Facilitation of Drinking: What is Facilitated,and Who is Affected?

The effect of a drinking companion on conspecifics was investigated using domestic hens (Callus gallus domesticus). The two aims of the investigation were firstly to see whether drinking is sensitive to social facilitation, and if so, to see whether it follows the same pattern as that of feeding, i.e. whether it acts primarily on the drinking behaviour rather than the amount ingested. The second aim was to see whether differences in how influenced the birds were by a drinking companion correlated with one or more social or non-social traits. Social facilitation was measured by comparing the intake and number of ‘head-ups’ (i.e. swallowing) of an individual in the presence of a thirsty or non-thirsty companion. The traits measured were as follows: the rank within each pair, social dependence/fear, reaction to a novel object, reaction in a feeding-inhibition test, and finally weight (as a possible index of the overall rank of the birds in the group). Social facilitation of drinking occurs but acts primarily on drinking behaviour (number of ‘head ups’); birds do not swallow as much per ‘head up’ as when they themselves are thirsty. The lightest (possibly most subdominant) birds were also the ones most influenced by the drinking conspecific. Furthermore, the heaviest (possibly most dominant) individuals were the ones with the shortest social distance and the longest latency in approaching a novel object. This meant that the birds that were most influenced by a drinking companion were also the individuals that were least fearful and showed the highest degree of exploration.     

Collective Waves of Sleep in Gulls (Larus spp.)

How should animals sleep in groups? Because sleeping reduces the ability of an individual to detect potential threats, not all individuals should sleep at the same time. The obvious solution of taking turns to sleep is not documented in animal groups. Individuals can also organize their sleeping bouts independently of each other but this simple strategy can be dangerous if too many individuals happen to sleep at the same time. One solution to this problem is to monitor the behaviour of other group members and adjust sleeping bouts accordingly. For instance, as the number of sleeping individuals increases, companions may decide that it must be a safe time to sleep. However, when fewer group members are sleeping, an individual may benefit by curtailing sleep, given that it would be more vulnerable than vigilant group members should an attack occur. Such monitoring can therefore lead to contagious behaviour in the group, which can be detected in a group by collective waves of activities through time. Using spectral analysis, I investigated the proportion of sleeping birds in loafing gulls (Larus spp.) as a function of time over 2 yr and found that in many groups, the proportion of sleeping birds rises and decreases in a systematic and statistically significant fashion. These results add more weight to the now increasingly supported view that vigilance in general is a social phenomenon and suggest that adaptive behaviour at the level of the individual can lead to collective phenomena such as waves of sleep in animal groups.     

Piping as a display of dominance by wintering Oystercatchers Haematopus ostralegus

Although Oystercatchers Haematopus ostralegus wintering in the Exe Estuary performed the piping display in any type of aggressive encounter, it was predominantly observed in two contexts. ‘Solitary piping’ occurred when one individual piped without an obvious opponent being in the immediate vicinity. In ‘piping ceremonies’, two or more individuals piped at each other for prolonged periods at close distance. Prolonged contests over mussels were usually won by the individual that had initially started piping. The most aggressive and successful birds (the dominants), engaged most often in solitary piping and piping ceremonies, using up to 10% of their time this way. This suggests that the behaviour was associated with dominance rank. The hypothesis is that the two birds are likely to start piping if their relative status has not already been established, so that a quick end to a dispute over a mussel or feeding site is not achieved. A dominant individual performs solitary piping from a distance to suppress the aggressive activities of other birds within its feeding range. If this signal fails, the dominant may join the ceremony, explaining why three was the most frequent number of birds involved in piping ceremonies. The ‘butterfly’ flight, used mainly by apparently dominant birds as they arrived on the feeding grounds, may signal their high status to birds already there. The outcome of several piping ceremonies involving birds of known dominance was consistent with this explanation. The location of piping ceremonies involving only two birds (interpreted as piping ceremonies not joined by dominants) at the extremities of the feeding ranges of the most dominant individuals is also consistent with this hypothesis, because this is where their dominance would be expected to decline and would therefore be more likely to be contested. This explanation of piping and butterfly displays assumes that individuals can recognize other individuals or quickly assess their dominance. Since dominant individuals were attacked less often than subdominants and attacking birds were rebuffed less than would be expected if recognition did not occur, we suggest that some form of dominance or individual recognition did exist. The width of the white collar did not correlate with dominance, but the individual variations in this feature may enable birds to distinguish each other and thus learn their relative status. It is intriguing that the diplomatist posture, which is typically used to rebuff misguided attackers, seems almost designed to give the opponent a particularly clear view of the white collar.     

Behavioural and hormonal effects of social isolation and neophobia in a gregarious bird species, the European starling (Sturnus vulgaris)

Separating gregarious individuals from their group members often results in behavioural and physiological changes, like increased levels of corticosterone. Testosterone and corticosterone, in particular, have been implicated in the response of mammals to novelty. Data in birds are, however, rare. The presence or absence of group members may also influence an individual's response to novel stimuli. We assessed the behaviour and hormonal response of European starlings (Sturnus vulgaris) to a novel object in two different situations and seasons: each starling was tested when separated and when in contact with its group members in May/June (breeding season) and again in September/October (non-breeding season). Starlings are gregarious throughout the year, but as foraging flocks are small during the breeding season and large during the non-breeding season, we assumed that non-breeding starlings would be more affected by social isolation. Overall, starlings had higher levels of corticosterone, lost more body mass, and were more active when they were separated from their group. Isolated individuals, however, did not show a greater neophobic response than individuals in the presence of their group members in either season. Circulating levels of testosterone and corticosterone were higher after a test with novel object than after a test with only the familiar feeding dish in both sexes and seasons. However, control tests for handling effects confirmed only the increase in testosterone. Our study shows that social isolation is stressful for unrelated and unpaired members of a wild flocking bird species and demonstrates that novelty can lead to a rise in testosterone in birds.     

Predator Avoidance and Alarm-response Behaviour in Kin-discriminating Tadpoles (Rana cascadae)

When captured by predators, the tadpoles of some species of frogs and toads may release an alarm substance that alters the behaviour of conspecifics. Such ‘alarm response’ behaviour has been proposed as a potential mechanism whereby related conspecifics may ‘warn’ relatives of a predator's presence and thus, improve their inclusive fitness. We examined predator avoidance and alarm response behaviour in tadpoles of the Cascades frog (Rana cascadae) and tested whether such behaviour is influenced by kinship factors. Tadpoles reduced activity when in the presence of a predatory newt (Taricha granulosa). Individuals in sibling groups were more active than both solitary tadpoles and individuals in mixed groups of siblings and nonsiblings. However, we found no evidence of an alarm response in R. cascadae. Behaviour of tadpoles in groups exposed only to predators was not different from that of tadpoles in groups exposed to predators plus crushed conspecifics. Tadpoles in groups exposed to crushed tadpoles were as active as tadpoles in groups exposed to water controls, and some test individuals fed upon the dead tadpoles. Thus, while R. cascadae tadpoles reduce activity in response to newt predators, crushed tadpoles appear to initiate a feeding response rather than an alarm response as has been previously proposed.     

Division of labour and specification of castes in the red imported fire ant Solenopsis invicta buren

Division of labour in Solenopsis invicta follows a familiar pattern: younger, smaller ants tend toward brood care while older, larger ants tend toward foraging. However, long-term observations of marked individuals reveal that length of nursing and foraging ‘careers’ and the age of transition between these activities vary considerably between and within size groups, and are related to length of life. Experiments with entire colonies show that larger ants are more likely than smaller ants to forage for insect prey. There are two main worker castes, ‘nurses’ and ‘foragers’, whose members span a wide age-size range, and a large ‘reserve’ subcaste, heterogeneous in age, size, and behaviour: reserves may nurse, forage, store liquid food, or relay food from nurses to foragers. The proportion of ants engaged in foraging decreases with colony size because many ants in large colonies are not exposed to recruitment signals.     

The Use and Transfer of Information About Predation Risk in Flocks of Wintering Finches

Several studies in behavior have focused in some way on how groups of prey gather and use information about predation risk. Although asymmetries in information about risk exist among members of real groups, we know little about how such uneven information might affect individual or group antipredator decisions. Hence, we studied the use and transfer of information about the risk of predation in small flocks of wintering birds. House finches (Carpodacus mexicanus; 28 groups of three) were held in large enclosures divided into safe and risky patches. We controlled the information about risk available to each individual by conducting attacks with a model hawk that was visible to only a single (informed) bird. Repeated attacks on a single individual did not reduce the amount of feeding by other birds in that patch, although the time to resume feeding after observing a response to an attack event was somewhat longer than after a no attack event. These results suggest that informed individuals impart some information to naïve (uninformed) birds, but this effect was not strong. In fact, the frequent return of informed individuals to feeders after observing uninformed individuals feed suggests that finches relied more on public information regarding safety than their own personal information in deciding when to feed. Group patch choice appeared to be based on a majority‐rules decision, although an effect of dominance status was apparent. Our results suggest that subordinate flock members may exert a large influence over group decision‐making by acting as spatial ‘anchors’.     

Status Signalling by Parus major: An Experiment in Deception

The ‘social control’ and ‘incongruence’ hypotheses, first put forward by ROHWER (1977) to explain how attempted ‘deceit’ status signalling is kept in check among winter-flocking birds, were tested under semi-natural conditions for Parus major. This species signals its social status by the width of its breast stripe. The lowest-ranked male in experimental flocks, each made up of four individuals, was manipulated in one of three ways: 1) the status signal was altered by painting the breast stripe to make it broader; 2) agonistic behaviour was altered by injecting testosterone; 3) both status signal and behaviour were manipulated. A study of the outcome of subsequent agonistic encounters by these birds revealed that the status of the manipulated individuals only rose when both their behaviour and status signal were altered. This indicates that the ‘social control’ hypothesis must be rejected, but not the ‘incongruence’ hypothesis.     

Do broiler chicks have a cognitive representation of food quality?. Appetitive, behavioural and ingestive responses to a change in diet quality

It has been observed that when a new diet formulation, or a new batch of the same diet formulation, is presented to poultry, there is a transitory suppression of feeding. It appears that the birds do not recognise the food as being edible, or classify it as being 'unknown'. In order to understand more about food recognition and rejection, the aim of this experiment was to determine whether cognitive processes are involved. Sixteen groups of four broiler chicks were used, and were fed a low quality diet in their home pens. The groups of chicks were trained to run a winding maze to gain access to a high quality diet in a test trough for 15min per day. When training was completed, the feed in the test trough for eight of the groups was changed to that which they received in the home pen, while the other eight groups received no change as a control. Time to traverse the runway did not show an immediate decrease on the day after the change (P>0.05) as would be expected if the birds used a cognitive comparison to determine speed of approach to the test trough. However, the experimental groups were significantly slower compared to the control groups after 4 days (P<0.05). The behaviour observed on the day of the change was indicative of frustration with more scratching and hurried movements shown (P<0.05), and less pecking at feed (P<0.01). Food consumption was lower for experimental groups compared to control groups on all days from the day of change onward (P<0.001). It was concluded that although there was no definitive evidence for the presence of a cognitive representation of food, this may have been due in part to the testing of groups of birds rather than individuals, and the way in which food quality is perceived. The occurrence of behaviours indicative of frustration suggest that a cognitive expectation may have been present.     

Mallards Feed Longer to Maintain Intake Rate under Competition on a Natural Food Distribution

Animals foraging in groups may benefit from a faster detection of food and predators, but competition by conspecifics may reduce intake rate. Competition may also alter the foraging behaviour of individuals, which can be influenced by dominance status and the way food is distributed over the environment. Many studies measuring the effects of competition and dominance status have been conducted on a uniform or highly clumped food distribution, while in reality prey distributions are often in‐between these two extremes. The few studies that used a more natural food distribution only detected subtle effects of interference and dominance. We therefore conducted an experiment on a natural food distribution with focal mallards Anas platyrhynchos foraging alone and in a group of three, having a dominant, intermediate or subordinate dominance status. In this way, the foraging behaviour of the same individual in different treatments could be compared, and the effect of dominance was tested independently of individual identity. The experiment was balanced using a 4 × 4 Latin square design, with four focal and six non‐focal birds. Individuals in a group achieved a similar intake rate (i.e. number of consumed seeds divided by trial length) as when foraging alone, because of an increase in the proportion of time feeding (albeit not significant for subordinate birds). Patch residence time and the number of different patches visited did not differ when birds were foraging alone or in a group. Besides some agonistic interactions, no differences in foraging behaviour between dominant, intermediate and subordinate birds were measured in group trials. Possibly group‐foraging birds increased their feeding time because there was less need for vigilance or because they increased foraging intensity to compensate for competition. This study underlines that a higher competitor density does not necessarily lead to a lower intake rate, irrespective of dominance status.     

Response to a Dangerous Enemy: Should a Brood Parasite be Mobbed?

Models of a long-tailed cuckoo and a song thrush (as a control) were presented 1.5 to 3 m from the nests of whiteheads, a species that the cuckoo is known to parasitize. Tests were conducted early in incubation (days 3 to 8) and during the nestling period. Most breeding females responded to the cuckoo by remaining inconspicuous or hiding. Other members of the cooperatively breeding group mobbed the cuckoo if they saw it. This difference in behaviour resulted in differences in overall response between incubation and nestling periods; the cuckoo was only occasionally mobbed during incubation because birds other than the female rarely approached the nest, whereas the cuckoo was almost always mobbed after eggs hatched. Cuckoos benefit from group breeding by whiteheads because chicks receive more food if reared by larger groups. I suggest that inconspicuous behaviour by the female in the presence of a cuckoo results in at least two effects: first, females can determine what the intentions of the cuckoo are, particularly if it intends to lay in the nest; second, no cues about size of breeding group are given to the cuckoo. Comparisons are drawn with the recent study of PAYNE et al. (1985) on splendid wrens and Chrysococcyx cuckoos. Despite similar social behaviour, splendid wrens and whiteheads behaved differently in the presence of a cuckoo. Mobbing may not always be the most appropriate tactic when faced with a brood parasite.     

The Effect of Within-Flock Spatial Position on the Use of Social Foraging Tactics in Free-Living Tree Sparrows

The benefit of producer (searches for own food) or scrounger (exploits the others’ food discoveries) foraging tactic in a group of socially feeding animals may depend on where the individual searches for food within the group. Scrounging may be more advantageous in the centre of the group, having more individuals around to join, while producing may be more beneficial at the edges, where more unexplored food patches may be found. This study shows within‐flock position correlates with foraging tactic use of feeding birds in socially foraging tree sparrows, Passer montanus. Sparrows staying closer to the centre of the flock found their food patches more frequently by joining (i.e. use more frequently the scrounging tactic) than those staying toward the edges. To our knowledge this is the first field study demonstrating the relationship between spatial position and foraging tactic use. We investigated this relationship under different perceived predation hazard, and found that under elevated risk of predation, central individuals may increase their use of joining more than individuals on the periphery of the flock. Moreover, we show that extremely specialized use of searching tactics may be very infrequent in tree sparrows. As both within‐flock position and search tactic use can be altered very quickly and without leaving the flock, individuals may easily alter them in order to adjust their behaviour.