Spatial and foraging behaviour, diet and morphogenesis of postlarvae and juveniles of Pempheris vanicolensis in the Gulf of Aqaba, Red Sea

Post-settlement ecology, ontogenesis of morphology and behaviour were studied in juvenile Pempheris vanicolensis (Pempheridae, Teleostei) in the Gulf of Aqaba, Red Sea. Postflexion larvae of 5.6-6.0 mm total length settled around breakwater heads and natural rocky outcrops of the subtidal zone, widely separated from adults dwelling in groups in coral reef crevices and caves. Settling juveniles formed groups of uniform age and foraged on zooplankton around their habitats. Their circadian behaviour was regulated by levels of illumination in the water, differing slightly in the various age groups. At sunset, juvenile P. vanicolensis migrated and dispersed in deeper water, returning before sunrise to their rocky habitats. Their patterns of foraging and migration behaviours were analysed. At 28.0-32.0 mm total length and 500 mg body weight, the increase in height and length ( H/L ratio) of the juvenile changed from monoto heterochronous, after which a rhomboid body shape, typical for this species, was formed. Concomitantly, the retina of the juvenile changed from a photopic to a scotopic one, the body became heavily pigmented, and the fish left the subtidal rocks, migrating to crevices in coral formations where they settled in close proximity to groups of adult P. vanicolensis .     

Effects of age and sex on intra-group spacing behaviour in juvenile savannah baboons,Papio cynocephalus cynocephalus

Twenty juvenile members of known genealogies in two baboon groups were studied over a 16-month period to evaluate a number of predictions about juvenile spacing behaviour based on the natural history of savannah baboons. Young juveniles (1–2·5 years old) approached more frequently and spent more time in proximity to other group members than did old juveniles (3–5·5 years old). In particular, young juveniles associated more closely with their mothers, particular adult males (possible fathers) and age-peers than did old juveniles. Approaches of young juveniles towards unrelated, high-ranking adults were more likely to occur during feeding than were those of old juveniles. Also, following such an approach, young juveniles were more likely than old juveniles to begin feeding immediately. The overall rates of feeding of old juveniles were depressed when they were in proximity to unrelated, high-ranking adults, whereas the feeding rates of young juveniles were not. Juvenile males approached adult males more often than did juvenile females. Juvenile females approached unrelated adult females more often than did juvenile males. Sex differences also existed in juveniles' choices of unrelated adult female neighbours. Juvenile females associated most often with lactating females, whereas juvenile males associated primarily with cycling females. During group resting, juvenile females approached adult females from higher-ranking matrilines more often than they approached adult females from lower-ranking matrilines. Juvenile males did not exhibit this attraction. Also, among old juveniles, females associated closely with their mothers, whereas males did not. Taken together, the results support the hypotheses that juvenile baboons associate with group members in ways that (1) enhance the probability of surviving an early period of high mortality, (2) create opportunities for social learning of sex-typical behaviours/skills, and, for females, (3) facilitate acquisition of familial dominance status.     

Social learning of hunting skills in juvenile marsh harriers <Emphasis Type="Italic">Circus aeruginosus</Emphasis>

The impact of social factors on the improvement of hunting skills of juvenile marsh harriers during their first autumnal migration were studied in SE Poland. While foraging with adult birds, juveniles performed more dives on prey both in terms of number of trials and rates. Hunting sessions of juveniles were more efficient in the presence of adults than in the absence of adults. Juveniles hunting with adults and other juveniles could select adequate habitat patches in which access to prey is easier. The role of vertical and horizontal transmission of information in the development of hunting skills in juvenile marsh harrier were confirmed because faster development of hunting ability was achieved in the social hunting after the end of their postfledging dependency period.     

From cradle to early grave: juvenile mortality in European shags Phalacrocorax aristotelis results from inadequate development of foraging proficiency

In most long-lived animal species, juveniles survive less well than adults. A potential mechanism is inferior foraging skills but longitudinal studies that follow the development of juvenile foraging are needed to test this. We used miniaturized activity loggers to record daily foraging times of juvenile and adult European shags Phalacrocorax aristotelis from fledging to the following spring. Juveniles became independent from their parents 40 days post-fledging. They compensated for poor foraging proficiency by foraging for approximately 3 h d(-1) longer than adults until constrained by day length in early November. Thereafter, juvenile foraging time tracked shortening day length up to the winter solstice, when foraging time of the two age classes converged and continued to track day length until early February. Few individuals died until midwinter and mortality peaked in January-February, with juvenile mortality (including some of the study birds) five times that of adults. In their last two weeks of life, juveniles showed a marked decline in foraging time consistent with individuals becoming moribund. Our results provide compelling evidence that juveniles compensate for poor foraging proficiency by increasing foraging time, a strategy that is limited by day length resulting in high winter mortality.     

Wild fledgling tits do not mob in response to conspecific or heterospecific mobbing calls

Mobbing, where birds harass a predator through a combination of vocalizations and stereotyped behaviours, is an effective anti-predator behaviour for many species. Mobbing may be particularly important for juveniles, as these individuals are often more vulnerable than adults. Although the component behaviours of mobbing are often considered to be un-learned, there are few confirmatory data, and the developmental trajectory of mobbing is unknown. In this study, we tested whether conspecific or heterospecific mobbing calls initiated mobbing behaviour in juvenile Blue Tits Cyanistes caeruleus. We located wild adult and recently fledged juvenile Blue Tits and presented them with playback recordings of adult conspecific (Blue Tit) and heterospecific (Great Tit Parus major) mobbing alarm calls. Although adult birds readily mob in response to these types of playbacks, juveniles did not exhibit characteristic mobbing behaviour. Some juveniles did, however, exhibit individual components of mobbing behaviour found in mobbing, despite not producing adult-like mobbing behaviour in response to either conspecific or heterospecific playback. These results suggest that, although birds might be capable of mobbing as juveniles, the associations between the non-vocal stereotyped mobbing behaviours and mobbing calls may be learned.     

Social enhancement can create adaptive,arbitrary and maladaptive cultural traditions

Many animals are known to learn socially, i.e. they are able to acquire new behaviours by using information from other individuals. Researchers distinguish between a number of different social-learning mechanisms such as imitation and social enhancement. Social enhancement is a simple form of social learning that is among the most widespread in animals. However, unlike imitation, it is debated whether social enhancement can create cultural traditions. Based on a recent study on capuchin monkeys, we developed an agent-based model to test the hypotheses that (i) social enhancement can create and maintain stable traditions and (ii) social enhancement can create cultural conformity. Our results supported both hypotheses. A key factor that led to the creation of cultural conformity and traditions was the repeated interaction of individual reinforcement and social enhancement learning. This result emphasizes that the emergence of cultural conformity does not necessarily require cognitively complex mechanisms such as ‘copying the majority’ or group norms. In addition, we observed that social enhancement can create learning dynamics similar to a ‘copy when uncertain’ learning strategy. Results from additional analyses also point to situations that should favour the evolution of learning mechanisms more sophisticated than social enhancement.     

Nocturnal relocation of adult and juvenile coral reef fishes in response to reef noise

Juvenile and adult reef fishes often undergo migration, ontogenic habitat shifts, and nocturnal foraging movements. The orientation cues used for these behaviours are largely unknown. In this study, the use of sound as an orientation cue guiding the nocturnal movements of adult and juvenile reef fishes at Lizard Island, Great Barrier Reef was examined. The first experiment compared the movements of fishes to small patch reefs where reef noise was broadcast, with those to silent reefs. No significant responses were found in the 79 adults that were collected, but the 166 juveniles collected showed an increased diversity each morning on the reefs with broadcast noise, and significantly greater numbers of juveniles from three taxa (Apogonidae, Gobiidae and Pinguipedidae) were collected from reefs with broadcast noise. The second experiment compared the movement of adult and juvenile fishes to reefs broadcasting high (>570 Hz), or low (<570 Hz) frequency reef noise, or to silent reefs. Of the 122 adults collected, the highest diversity was seen at the low frequency reefs; and adults from two families (Gobiidae and Blenniidae) preferred these reefs. A similar trend was observed in the 372 juveniles collected, with higher diversity at the reefs with low frequency noises. This preference was seen in the juvenile apogonids; however, juvenile gobiids were attracted to both high and low sound treatments equally, and juvenile stage Acanthuridae preferred the high frequency noises. This evidence that juvenile and adult reef fishes orientate with respect to the soundscape raises important issues for management, conservation and the protection of sound cues used in natural behaviour.     

Inter-population variation in the behaviour of adult and juvenile Red-footed Boobies Sula sula

Early life is a critical phase of the life cycle of animals and is attracting increased attention because little information is available on the behaviour of young individuals during this period. Behaviour during early life is probably influenced by the environmental conditions encountered by young animals, but data on intraspecific variation between breeding sites during this crucial period of life are limited. Here we study variability in the foraging behaviour of juveniles and adults in three colonies of a pantropical seabird, the Red-footed Booby Sula sula. Both adults and juveniles were measured and fitted with GPS loggers in three remote islands: Genovesa (Galapagos, Eastern Pacific Ocean), Europa (Western Indian Ocean) and Surprise (New Caledonia, Western Pacific Ocean). Foraging behaviour was compared between age-classes, sex and colonies by examining trip characteristics, different behaviours at sea, potential associations between individuals and morphological characteristics. Compared with adults, juveniles conducted shorter trips that were restricted to around the colony, especially on Genovesa (max. range: 203.4 ± 125.1 km and 3.6 ± 3.1 km, respectively). Juveniles appeared more constrained by poor flight skills and experience rather than by their morphology. Adults travelled 45% of the time during at-sea trips, whereas juveniles spent a a lower proportion of time travelling but foraged more often using an ‘area-restricted search’ behaviour, potentially training to catch prey. Associations between juveniles were commonly detected in the three colonies and occurred mostly during foraging, suggesting that social learning is an important strategy. Variability of morphometric measurements in both adults and juveniles was high between sites, with larger birds found on Genovesa. These results suggest that adaptations to local environmental conditions are already visible in their early life. Future studies should continue to investigate the behavioural flexibility of juvenile birds to better understand the effect of local environmental conditions during this critical stage of life.     

Do naive juvenile seabirds forage differently from adults?

Foraging skills of young individuals are assumed to be inferior to those of adults. The reduced efficiency of naive individuals may be the primary cause of the high juvenile mortality and explain the deferment of maturity in long-lived species. However, the study of juvenile and immature foraging behaviour has been limited so far. We used satellite telemetry to compare the foraging movements of juveniles, immatures and breeding adult wandering albatrosses Diomedea exulans, a species where foraging success is positively influenced by the distance covered daily. We showed that juveniles are able to use favourable winds as soon as the first month of independence, but cover shorter distances daily and spend more time sitting on water than adults during the first two months after fledging. These reduced movement capacities do not seem to be the cause of higher juvenile mortality. Moreover, juveniles almost never restrict their movement to specific areas, as adults and immatures frequently do over shelf edges or oceanic zones, which suggest that the location of appropriate areas is learned through experience. Immatures and adults have equivalent movement capacities, but when they are central place foragers, i.e. when adults breed or immatures come to the colony to display and pair, immatures make shorter trips than adults. The long duration of immaturity in this species seems to be related to a long period of learning to integrate the foraging constraints associated with reproduction and central place foraging. Our results indicate that foraging behaviour of young albatrosses is partly innate and partly learned progressively over immaturity. The first months of learning appear critical in terms of survival, whereas the long period of immaturity is necessary for young birds to attain the skills necessary for efficient breeding without fitness costs.     

The role of parent–offspring interactions during and after fledging in the Black-legged Kittiwake

Most bird species endure a high mortality at fledging, and selection should favour parental behaviour diminishing these costs. Post-fledging parental care varies greatly among species and is often linked to parent–offspring recognition. In the Black-legged Kittiwake (Rissa tridactyla), fledglings need to return to the natal nest to be fed by their parents until independence. Rejections of fledglings by non-parent adults may be fairly violent, and parents are expected to recognize and help their chicks at the time of first return. However, previous cross-fostering experiments pointed out that parents are not able to recognize their chicks up to 15 days before fledging. In this paper, we study the behaviour of both parents and juveniles at fledging. We found that parents answered significantly more to their fledgling's calls than to those of others. Compared to silent juveniles, juveniles that called before landing were more likely to be accepted by their parents. No such pattern was observed with foreign juveniles, indicating that fledglings’ voice may carry individual identity. Furthermore, fledglings found their way back to the natal nest faster when parents attended the natal nest and reacted to their offspring's calls than when they were absent or inactive. Such interactions may therefore diminish juvenile mortality at fledging.     

Female Bechstein's Bats Share Foraging Sites with Maternal Kin but do not Forage Together with them – Results from a Long‐Term Study

In many social animals, group members exchange information about where to feed. Thereby, they may gain direct benefits, for example, if social hunting enhances individual foraging success. Alternatively, individuals may receive indirect fitness benefits by preferentially sharing information about suitable feeding sites with kin. Indeed, in some species, a positive correlation between the degree of relatedness among individuals and the overlap among their foraging areas was found. However, sharing foraging sites with kin can also have costs if it increases food competition, which is not compensated by direct benefits. The goal of this study was to investigate whether sharing of individual foraging areas in female Bechstein's bats is best explained by kin selection or by direct benefits through social foraging. To assess their individual foraging behaviour, we analysed radio‐tracking data of 22 members of one maternity colony, including nine mother–daughter pairs, seven pairs of less closely related individuals and six pairs of unrelated bats. We examined the bats' fidelity to specific foraging areas during several years and quantified the influence of kinship on the overlap among individual foraging areas. By measuring how close to each other the bats foraged, we assessed whether individuals with overlapping areas are likely to forage together. Our study confirms previous findings that Bechstein's bats show high fidelity to foraging areas across years. Moreover, we found that relatives share foraging areas significantly more often compared with unrelated colony members. Finally, our data reveal for the first time that most colony members that share foraging areas are unlikely to forage together. This suggests that female Bechstein's bats gain no direct benefits from sharing foraging areas with members of the same maternal lineage. Our findings also have implications for conservation as habitat loss within a colony's home range might expose entire matrilines to high risks.     

Ecological Risk Aversion and Foraging Behaviors of Juvenile Squirrel Monkeys (Saimiri sciureus)

The ‘ecological risk aversion hypothesis’ [C.H. Janson and C.P. van Schaik, Juvenile Primates, Oxford Univ. Press, New York (1993), pp. 57–74] proposes that the pattern of slow growth characteristic of juvenile primates is a response to ecological risks (predation and starvation) experienced by juveniles. Juveniles are thought to avoid predation risk by positioning themselves near conspecifics, therefore experiencing high levels of feeding competition with older individuals, reduced access to resources and, consequently, high starvation risks during periods of food scarcity. The present study compared the foraging behaviors of juvenile and adult squirrel monkeys, a small neotropical primate characterized by a long juvenile period, to determine how predation and starvation risks affected juvenile behaviors. The study was conducted in Eastern Amazonia, in a seasonal environment. Due to their slow development, small body size and large group sizes, it was expected that juveniles in this species would behave in a manner consistent with the risk aversion hypothesis. However, age differences in foraging efficiency and foraging success were smaller than predicted. There was also no evidence that juveniles sacrificed access to food for predator protection. Adults did not have preferential access to fruit patches and direct competition was rare. Feeding competition for prey, the most common resource in the troop's diet, was negligible. Therefore, the slow growth and long juvenile period of squirrel monkeys do not correspond with evidence of predation or starvation risk, as predicted by the risk aversion hypothesis.     

Foraging Ontogeny is not Linked to Delayed Maturation in Squirrel Monkeys (Saimiri sciureus)

The acquisition of complex foraging behaviors by young is a proposed cause of a prolonged juvenile phase in many vertebrates, including primates. I compared the foraging behaviors of infant, juvenile and adult squirrel monkeys to determine if significant age‐related differences in foraging behavior and efficiency were present. Infants and juveniles differed from each other in patterns of prey and fruit foraging, but few differences existed between juveniles and adults. Despite differing in the use of foraging substrates, young juveniles (8–12 mo) were as efficient as older juveniles (1–4 yr) and adults at capturing and processing large prey. Young juveniles (<1 yr) were limited in their ability to consume husked palm fruits due to an inability to peel them to obtain pulp. By 1 yr of age, however, foraging behaviors of adults and juveniles were nearly indistinguishable. The absence of meaningful differences between adults and juvenile foraging is not consistent with the hypothesis that the need to develop foraging skills accounts for the pattern of extended juvenility in squirrel monkeys.     

Behavioural and Genetic Data Suggest that Bechstein's Bats Predominantly Mate Outside the Breeding Habitat

Investigating mating systems of species with a cryptic lifestyle often requires a combination of behavioural and genetic data. We used such a combined approach to investigate the mating system of the communal breeding Bechstein's bat (Myotis bechsteinii). Although females of this species are philopatric, gene flow among colonies is high. Gene flow occurs if dispersing males mate within the colony to which they moved. Males could gain local matings by defending resources or females in their breeding habitat. Alternatively, mating may take place at swarming sites, apart from the breeding habitat of the females, where males and females of several colonies meet. Whether or not males defend resources or females in the breeding habitat is of importance for understanding the mating system. Detailed observations of individual foraging and roosting behaviour over 4 yr suggest that males do not defend resources or females to gain matings. Moreover, paternity assignment based on microsatellite data of four complete juvenile cohorts showed that local males fathered less than 25% of the juveniles born in the colony where they settled. Even more striking, none of the males that had immigrated into our study area reproduced with the local females.     

Prolonged Parental Feeding in Tool‐Using New Caledonian Crows

According to life‐history theory, the duration of extended parental feeding is determined by the costs and benefits of maximising reproductive success. Therefore, the length of regular parental provisioning should be correlated with the time required for juveniles to acquire the skills that they need to be independent. The relatively few cases of extremely prolonged parental feeding in both land and sea birds appear to be consistent with this prediction because they are associated with learning‐intensive foraging techniques. New Caledonian crows have the most intricate tool manufacture techniques amongst non‐human animals and juveniles take over 1 yr to reach adult‐like proficiency in their tool skills. We investigated the prediction that this species also should have prolonged parental provisioning. We found that these crows have one of the longest known periods of regular extended parental provisioning in birds. Some parents regularly fed juveniles for up to 10 mo post‐fledging. Humans also stand out amongst primates because of their learning‐intensive foraging strategies and an extended period of juvenile dependence. The independently evolved association between a relatively high level of technological skill in foraging and prolonged juvenile provisioning in both humans and New Caledonian crows raises the possibility that these two characteristics might be causally related.     

Conformist learning in nine-spined sticklebacks' foraging decisions

Theoretical analyses have reported that in most circumstances where natural selection favours reliance on social learning, conformity (positive frequency-dependent social learning) is also favoured. These findings suggest that much animal social learning should involve a copy-the-majority strategy, yet there is currently surprisingly little evidence for conformist learning among animals. Here, we investigate this possibility in the nine-spined stickleback (Pungitius pungitius) by manipulating the number of demonstrator fish at two feeders, one rich and one poor, during a demonstration phase and evaluating how this affects the likelihood that the focal fish copy the demonstrators'' apparent choices. As predicted, we observed a significantly increased level of copying with increasing numbers of demonstrators at the richer of the two feeders, with copying increasing disproportionately, rather than linearly, with the proportion of demonstrators at the rich foraging patch. Control conditions with non-feeding demonstrators showed that this was not simply the result of a preference for shoaling with larger groups, implying that nine-spined sticklebacks copy in a conformist manner.     

Social interactions of juvenile brown boobies at sea as observed with animal-borne video cameras

While social interactions play a crucial role on the development of young individuals, those of highly mobile juvenile birds in inaccessible environments are difficult to observe. In this study, we deployed miniaturised video recorders on juvenile brown boobies Sula leucogaster, which had been hand-fed beginning a few days after hatching, to examine how social interactions between tagged juveniles and other birds affected their flight and foraging behaviour. Juveniles flew longer with congeners, especially with adult birds, than solitarily. In addition, approximately 40% of foraging occurred close to aggregations of congeners and other species. Young seabirds voluntarily followed other birds, which may directly enhance their foraging success and improve foraging and flying skills during their developmental stage, or both.     

Conformity and differentiation are two sides of the same coin

Variation between individuals is a key component of selection and hence evolutionary change. Social interactions are important drivers of variation, potentially making behaviour more similar (i.e., conform) or divergent (i.e., differentiate) between individuals. While documented across a wide range of animals, behaviours and contexts, conformity and differentiation are typically considered separately. Here, we argue that rather than independent concepts, they can be integrated onto a single scale that considers how social interactions drive changes in interindividual variance within groups: conformity reduces variance within groups while differentiation increases it. We discuss the advantages of placing conformity and differentiation at different ends of a single scale, allowing for a deeper understanding of the relationship between social interactions and interindividual variation.     

Social behaviour of juvenile red sea urchins, Strongylocentrotus franciscanus (Agassiz)

This study examined social behaviour of juvenile red sea urchins, Strongylocentrotus franciscanus (Agassiz). At four coastal locations in British Columbia, Canada, distributions of juvenile sea urchins were examined in relation to adults. One-third of all juveniles were found underneath an adult “spine canopy”, another third outside the spine canopy but close to an adult, and the remainder were found apart from adults. Juveniles associated with adults were contagiously distributed with each other. In the laboratory, juveniles chose adults over other target locations. They selected adults in particular locations over others, and in one case showed selection among adults; but they did not select between fed and non-fed adults, nor between adults that had been “protective” and “non-protective” of juveniles in the field. In time-lapse film studies, starved juveniles showed little change in social behaviour when food was introduced, but showed a dramatic change in behaviour when predators were active. We suggest that juvenile red sea urchins are found under the spine canopies of adults as a result of juvenile behaviour; and that this interaction functions to protect juveniles from predation. The strong gregarious nature of juveniles among themselves may have evolved in a different selective situation as a defence against predation and to ensure reproductive success.     

Paternal signature in kin recognition cues of a social insect: concealed in juveniles,revealed in adults

Kin recognition is a key mechanism to direct social behaviours towards related individuals or avoid inbreeding depression. In insects, recognition is generally mediated by cuticular hydrocarbon (CHC) compounds, which are partly inherited from parents. However, in social insects, potential nepotistic conflicts between group members from different patrilines are predicted to select against the expression of patriline-specific signatures in CHC profiles. Whereas this key prediction in the evolution of insect signalling received empirical support in eusocial insects, it remains unclear whether it can be generalized beyond eusociality to less-derived forms of social life. Here, we addressed this issue by manipulating the number of fathers siring clutches tended by females of the European earwig, Forficula auricularia, analysing the CHC profiles of the resulting juvenile and adult offspring, and using discriminant analysis to estimate the information content of CHC with respect to the maternal and paternal origin of individuals. As predicted, if paternally inherited cues are concealed during family life, increases in mating number had no effect on information content of CHC profiles among earwig juveniles, but significantly decreased the one among adult offspring. We suggest that age-dependent expression of patriline-specific cues evolved to limit the risks of nepotism as family-living juveniles and favour sibling-mating avoidance as group-living adults. These results highlight the role of parental care and social life in the evolution of chemical communication and recognition cues.