Opening the Social Gateway: Early Vocal and Social Sensitivities in Brown-Headed Cowbirds (Molothrus ater)

The organization of cowbird ( Molothrus ater ) social groups affords individuals living in the groups different opportunities for learning and also structures trajectories of social development. Here, we studied the influence of adults on social organization of very young cowbirds. In three experiments, we housed juvenile birds in large, seminatural environments that either contained or did not contain adult conspecifics. We then observed the social associations and vocalizations of juveniles in each environment. The presence of adults in the social environment influenced the assortment and singing patterns of juveniles, although throughout the three experiments adults rarely interacted physically with juveniles. Juveniles housed with adults interacted with other juveniles more often and sang significantly less often than juveniles housed without adults. Effects of adult presence or absence on social organization and singing patterns emerged extremely rapidly and could be reversed quickly. Taken as a whole, the experiments revealed that very young cowbirds, in the first days of independence from their hosts, were sensitive to, and reacted rapidly to, the composition of their social environment. Specifically, presence of other age classes of individuals within the group increased juvenile associations and delayed production of vocalizations by juvenile males. Self-organization within the social group produced different social environments, which could in turn create different gateways for social learning and vocal development.     

The importance of growing up: juvenile environment influences dispersal of individuals and their neighbours

Dispersal is a key ecological process that is strongly influenced by both phenotype and environment. Here, we show that juvenile environment influences dispersal not only by shaping individual phenotypes, but also by changing the phenotypes of neighbouring conspecifics, which influence how individuals disperse. We used a model system (Tribolium castaneum, red flour beetles) to test how the past environment of dispersing individuals and their neighbours influences how they disperse in their current environment. We found that individuals dispersed especially far when exposed to a poor environment as adults if their phenotype, or even one‐third of their neighbours’ phenotypes, were shaped by a poor environment as juveniles. Juvenile environment therefore shapes dispersal both directly, by influencing phenotype, as well as indirectly, by influencing the external social environment. Thus, the juvenile environment of even a minority of individuals in a group can influence the dispersal of the entire group.     

Detection of the Sexual Identity of Conspecifics through Volatile Chemical Signals in a Territorial Salamander

Territorial red‐backed salamanders (Plethodon cinereus) have been shown to use nonvolatile chemical signals in both territorial defense and to convey a variety of information to conspecifics. We investigated whether or not red‐backed salamanders could determine the sexual identity of conspecifics through volatile chemical signals, and we explored their use in the context of territorial defense. We exposed male and female red‐backed salamanders to four experimental treatments (i.e. filter papers that had been scent marked by male or female conspecifics for 1 and 5 d) and two control treatments (i.e. unscented filter papers for 1 and 5 d tests). The focal salamanders were prevented from physically accessing the scent marked filter papers and, presumably, some of the substrate scent marks had volatile components that were detected and interpreted by the focal salamanders. Both male and female red‐backed salamanders spent significantly more time in threat displays when they were exposed to volatile chemical signals from same‐sex conspecifics than they did toward similar signals from opposite‐sex conspecifics. A similar statistical pattern was observed for the amount of chemosensory sampling exhibited by focal red‐backed salamanders. From these results, we infer that red‐backed salamanders can determine the sexual identity of conspecifics through volatile chemical signals, some of which may be used in territorial defense. Further, such airborne pheromones may influence the spatial organization of salamander territories on the forest floor.     

An experimental analysis of social interaction in the common marmoset (Callithrix jacchus jacchus)

Adult and juvenile common marmosets were introduced to unfamiliar conspecifics individually and in whole groups. In introductions using animals of the same sex, adults were mutually hostile unless they were related or socially familiar; juveniles behaved submissively to adults and little aggression was observed between juveniles. In introductions using animals of the opposite sex, males of all ages solicited females but females did not reciprocate. When whole groups were introduced, in two experiments most of the elder group members behaved aggressively and younger animals did not interact frequently; but in a third experiment, in which the adult males were related, little aggression was observed and younger animals behaved amicably. It is argued that the behavioral reactions shown by individual marmosets are related to territorial hostility and that the behavior of juveniles to adults may aid assimilation of younger animals into new groups.     

Context-specific territorial behavior in urban birds: no evidence for involvement of testosterone or corticosterone

Testosterone produced by the gonads is a primary mediator of seasonal patterns of territoriality and may directly facilitate territorial behavior during an encounter with a potential intruder. Costs and benefits associated with territoriality can vary as a function of habitat, for example through differences in resource distribution between areas occupied by different individuals. We investigated behaviors in response to simulated territorial intrusions (hereafter territorial behaviors) in urban (Phoenix, Arizona) and nearby desert populations of two Sonoran Desert birds (Curve-billed Thrasher and Abert's Towhee). We also examined the degree to which these behaviors are mediated by testosterone (T) and the adrenal steroid, corticosterone (CORT), which can interact with T in territorial contexts. In both species, urban birds displayed more territorial behaviors than their desert conspecifics, but this difference was not associated with variation in either plasma total or in plasma free (i.e., unbound to binding globulins) T or CORT. In addition, neither plasma T nor plasma CORT changed as a function of duration of the simulated territorial intrusion. Urban Abert's Towhees displayed more territorial behaviors in areas where their population densities were high than in areas of low population densities. Urban Curve-billed Thrashers displayed more territorial behaviors in areas with a high proportion of desert-type vegetation, particularly in areas that differed in vegetation composition from nearby randomly sampled areas, than in areas with a high proportion of exotic or non-desert type vegetation. Associations between territorial behavior and habitat characteristics were not related to plasma T or CORT. Understanding the hormonal processes underlying these associations between behavior and habitat may provide insight into how free-ranging animals assess territorial quality and alter their defensive behavior accordingly.     

Tests of aggregative preferences of wandering salamanders (Aneides vagrans)

Plethodontid salamanders usually exhibit territorial spacing patterns and behaviors. Previous works suggest that plethodontids from Vancouver Island, Canada lack strong territoriality. In one Vancouver Island species, Aneides vagrans, salamanders may even be found in the field in small aggregations. We investigated preferences of A. vagrans for occupied vs unoccupied cover objects and for cover objects occupied by conspecifics of different age/size classes. We also tested A. vagrans preferences for conspecific vs heterospecific salamanders in an attempt to determine if the lack of territoriality by A. vagrans is species-specific and perhaps related to habitat segregation. A. vagrans did not avoid occupied cover objects or exhibit a preference for occupied vs unoccupied sites. Age class did not affect cover object choice. A. vagrans did not avoid heterospecifics that exhibit the highest degree of resource overlap (Plethodon vehiculum), which we believe might be attributable to a benign and resource-rich habitat, although alternative explanations exist. A. vagrans did avoid syntopic newts (Taricha granulosa), though this is probably due to toxic newt skin secretions.     

The effects of conspecifics on burrow selection in juvenile spotted salamanders (<Emphasis Type="Italic">Ambystoma maculatum)</Emphasis>

Spotted salamanders (Ambystoma maculatum) are pond-breeding amphibians that disperse into terrestrial habitat from natal wetlands after undergoing metamorphosis, relying on small-mammal burrows and coarse woody debris for refugia. The effect of conspecifics on burrow use in juvenile salamanders is poorly understood. Determining how the presence of conspecifics influences the settlement decisions of juvenile salamanders can increase our understanding of amphibian dispersal and our ability to predict population dynamics. We conducted behavioral laboratory trials using 58 recently metamorphosed salamanders to examine how salamanders selected burrows in the presence of conspecifics. Salamanders were more likely to settle in a burrow that was occupied by a conspecific versus an unoccupied burrow. Our results indicate that juvenile salamanders may show conspecific attraction and/or trailing behavior during the dispersal phase.     

Early‐life foraging: Behavioral responses of newly fledged albatrosses to environmental conditions

In order to survive and later recruit into a population, juvenile animals need to acquire resources through the use of innate and/or learnt behaviors in an environment new to them. For far‐ranging marine species, such as the wandering albatross Diomedea exulans, this is particularly challenging as individuals need to be able to rapidly adapt and optimize their movement strategies in response to the highly dynamic and heterogeneous nature of their open‐ocean pelagic habitats. Critical to this is the development and flexibility of dispersal and exploratory behaviors. Here, we examine the movements of eight juvenile wandering albatrosses, tracked using GPS/Argos satellite transmitters for eight months following fledging, and compare these to the trajectories of 17 adults to assess differences and similarities in behavioral strategies through time. Behavioral clustering algorithms (Expectation Maximization binary Clustering) were combined with multinomial regression analyses to investigate changes in behavioral mode probabilities over time, and how these may be influenced by variations in day duration and in biophysical oceanographic conditions. We found that juveniles appeared to quickly acquire the same large‐scale behavioral strategies as those employed by adults, although generally more time was spent resting at night. Moreover, individuals were able to detect and exploit specific oceanographic features in a manner similar to that observed in adults. Together, the results of this study suggest that while shortly after fledging juvenile wandering albatrosses are able to employ similar foraging strategies to those observed in adults, additional skills need to be acquired during the immature period before the efficiency of these behaviors matches that of adults.     

Effects of conditionally expressed phenotypes and environment on amphibian dispersal in nature

Individuals vary greatly in the distance they disperse, and in doing so, strongly affect ecological and evolutionary processes. Dispersal, when viewed as a component of phenotype, can be affected independently or jointly by environment. However, among taxa with complex life cycles that occupy different habitats over ontogeny, the effects of environment on dispersal and the interaction between environment and phenotype remains poorly understood. Here, we conducted a field experiment to measure how dispersal distance was affected by phenotype, environment experienced before and after metamorphosis, and their interaction. We manipulated the environment encountered by a pond‐breeding salamander Ambystoma annulatum during the aquatic larval stage and again as dispersing terrestrial juveniles. After assaying juvenile phenotype (exploration behavior, body condition, and morphology), we then measured the initial distance dispersed by juveniles. The distance moved by dispersing salamanders was affected by attributes of both larval and juvenile habitat, with salamanders that encountered low quality habitat in either life stage moving the farthest. However, we did not find support for an interactive effect of phenotype and environment affecting the distance moved by dispersers. Interestingly, exploration behavior explained the distance moved by philopatric animals but not dispersing ones. Our findings indicate that the environment experienced before metamorphosis can affect juvenile dispersal behavior, and demonstrates the need to consider dispersal in species with complex life cycles to understand the coupling between local and regional population dynamics.     

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.     

Harem structure and female territoriality in the dwarf hawkfish <Emphasis Type="Italic">Cirrhitichthys falco</Emphasis> (Cirrhitidae)

We investigated the territoriality and the spatial and mating relationships of the haremic hawkfish, Cirrhitichthys falco, on a reef off Kuchierabu-jima Island in southern Japan. Each individual maintained a territorial home range which was defended against same-sex conspecifics at the boundary of the home range. The territory of each male encompassed the territories of 2–3 females, allowing the male to completely monopolize mating opportunities with those females. Based on our observations, we classified the harem type of C. falco as a territorial female type. Large juveniles maintained independent home ranges outside the female territories. In contrast, small juveniles were allowed to cohabit within the territory of an adult female. Stomach contents analysis revealed that the smallest size class of C. falco fed primarily on copepods. In contrast, all other size classes fed primarily on decapods. Together, these results suggest that female territoriality plays an important role in defending food resources.     

Giant chacoan peccary: Feeding and social behavior of a captive group in natural habitat

Social behavior of the giant Chacoan peccary, also known as the taguá, is relatively unknown due to the harsh environment and endangered status of the population. In order to characterize feeding and social behaviors, a group of 7 adults, 2 juveniles, and 3 infants was observed in their feeding area for 99.25 hours over 18 days during October and November 1989. All animals were housed in a 2 ha natural-habitat enclosure in the Chaco of Paraguay. It was determined that these peccaries fed in intervals throughout daylight hours and that juveniles spent more time per approach in the feeding area than adult males. It was also observed that infants were accepted and well tolerated by the entire group, including adult males. These observations have relevance to taguá behavior in the wild and provide new insights into social patterns of this recently discovered species.     

Social Reversal of Sex‐Biased Aggression and Dominance in a Biparental Cichlid Fish (Julidochromis marlieri)

In biparental species, aggression, dominance, and parental care are typically sexually dimorphic. While behavioral dimorphism is often strongly linked to gonadal sex, the environment—either social or ecological—may also influence sex‐biased behavior. In the biparental cichlid fish Julidochromis marlieri, the typical social environment for breeding pairs consists of large females paired with smaller males. While both sexes are capable of providing territory defense and parental care, the larger female provides the majority of defense for the pair, while the smaller male remains in the nest guarding their offspring. We examine the contributions of sex and relative mate size to these sex‐biased behaviors in monogamous J. marlieri pairs. Both female‐larger and male‐larger pairs were formed in the laboratory and were observed for territorial aggression (against conspecifics and heterospecifics), dominance, and parental care. In female‐larger pairs, territorial aggression and intra‐pair dominance were female‐biased, while in male‐larger pairs this bias was reversed. For both pairing types, the presence of an intruder amplified sex differences in territorial aggression, with the larger fish always attacking with greater frequency than its mate. Though less robust, there was evidence for plasticity of sex‐bias for some egg care related behaviors in the inverse direction. Our study suggests that relative mate size strongly influences the sex bias of aggression and dominance in J. marlieri and that this aspect of the social environment can override the influence of gonadal sex on an individual's behavior. The remarkable plasticity of this species makes Julidochromis an exciting model that could be used to address the relationship between proximate and ultimate mechanisms of behavioral plasticity.     

Factors affecting reproduction in zoo‐housed geoffroy's tamarins(Saguinus geoffroyi)

The captive population of Geoffroy's tamarins (Saguinus geoffroyi) has suffered a severe decline over the past 10 years. This population decline is attributed not to a failure to produce offspring, but rather to a failure to successfully rear offspring. To date, no studies have quantitatively examined the behaviors and hormones of this tamarin species in captivity. Therefore, this study was conducted to determine whether there were any discernable factors that could be correlated with failure to rear offspring in S. geoffroyi. Fifteen adult Geoffroy's tamarins (GTs) at the Cleveland Metroparks Zoo (CMZ) were observed by means of instantaneous sampling on a focal animal. In addition, all instances of social behaviors were recorded. A factorial arrangement of treatments was used, as animals were divided between a colony‐housing situation and a non‐colony situation with hand‐reared and mother‐reared animals in both treatments. Repeated‐measures analysis of variance (ANOVA) showed no interactions between the treatments, and no differences between rearing histories for the behaviors studied. However, animals housed in a colony setting exhibited higher levels of aggressive behaviors and lower activity levels compared to those in a non‐colony setting. There also was a trend for colony‐housed animals to huddle, scent‐mark, and exhibit sexual behaviors more often than non‐colony animals. First‐morning‐void urine samples were collected once per week, and assays for E₁C and PdG indicated that reproduction was not being suppressed. ANOVA conducted on samples assayed for excreted cortisol showed no differences in mean cortisol concentration by rearing histories or housing conditions. While these physiological indicators reveal no signs of stress, high levels of aggressive and territorial behaviors indicate social unrest in the colony‐housed condition, which may be contributing to the poor reproductive success of those individuals. Zoo Biol 22:545–559, 2003. © 2003 Wiley‐Liss, Inc.     

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.     

Noninvasive genetic sampling reveals intrasex territoriality in wolverines

Due to its conspicuous manifestations and its capacity to shape the configuration and dynamics of wild populations, territorial behavior has long intrigued ecologists. Territoriality and other animal interactions in situ have traditionally been studied via direct observations and telemetry. Here, we explore whether noninvasive genetic sampling, which is increasingly supplementing traditional field methods in ecological research, can reveal territorial behavior in an elusive carnivore, the wolverine (Gulo gulo). Using the locations of genotyped wolverine scat samples collected annually over a period of 12 years in central Norway, we test three predictions: (1) male home ranges constructed from noninvasive genetic sampling data are larger than those of females, (2) individuals avoid areas used by other conspecifics of the same sex (intrasexual territoriality), and (3) avoidance of same‐sex territories diminishes or disappears after the territory owner's death. Each of these predictions is substantiated by our results: sex‐specific differences in home range size and intrasexual territoriality in wolverine are patently reflected in the spatial and temporal configuration of noninvasively collected genetic samples. Our study confirms that wildlife monitoring programs can utilize the spatial information in noninvasive genetic sampling data to detect and quantify home ranges and social organization.     

Temporal constraints on the potential role of fry odors as cues of past reproductive success for spawning lake trout

Deciding where to reproduce is a major challenge for most animals. Many select habitats based upon cues of successful reproduction by conspecifics, such as the presence of offspring from past reproductive events. For example, some fishes select spawning habitat following odors released by juveniles whose rearing habitat overlaps with spawning habitat. However, juveniles may emigrate before adults begin to search for spawning habitat; hence, the efficacy of juvenile cues could be constrained by degradation or dissipation rates. In lake trout (Salvelinus namaycush), odors deposited by the previous year's offspring have been hypothesized to guide adults to spawning reefs. However, in most extant populations, lake trout fry emigrate from spawning reefs during the spring and adults spawn during the fall. Therefore, we postulated that the role of fry odors in guiding habitat selection might be constrained by the time between fry emigration and adult spawning. Time course chemical, physiological, and behavioral assays indicated that the odors deposited by fry likely degrade or dissipate before adults select spawning habitats. Furthermore, fry feces did not attract wild lake trout to constructed spawning reefs in Lake Huron. Taken together, our results indicate fry odors are unlikely to act as cues for lake trout searching for spawning reefs in populations whose juveniles emigrate before the spawning season, and underscore the importance of environmental constraints on social cues.     

Observations of false mating behavior in entocytherid ostracods from the northwestern United States

Five male entocytherid ostracods of four different species recovered from signal crayfish Pacifastacus leniusculus were observed exhibiting false mating behaviors with either another adult male, an A1 sub‐adult male, or an A3 juvenile of indeterminate species and sex. In all five instances, the ostracod being mated was not sclerotized, indicating recent molt. In one case, a male of Uncinocythere thektura was observed in false copulation with an A1 male of a different species, U. occidentalis, suggesting that taxonomic designations of females in mixed populations of Uncinocythere can be problematic. Possible reasons for false mating behavior include stress of being confined post‐collection, lack of available females, or possibly males mistaking non‐sclerotized partners for A1 females.     

Adult Male Density Influences Juvenile Microhabitat Use in a Territorial Lizard

Habitat choice often has strong effects on performance and fitness. For many animals, optimal habitats differ across age or size classes, and individuals shift habitat use through ontogeny. Although many studies document ontogenetic habitat shifts for various taxa, most are observational and do not identify the causal factor of size‐specific habitat variation. Field observations of the brown anole lizard (Anolis sagrei) show that juveniles perch on shorter and thinner vegetation than adults. We hypothesized that this variation is due to adult males forcing smaller juveniles to less preferred habitat. To test this assertion, we manipulated adult male densities in mesh enclosures with artificial trees to examine the response of juvenile microhabitat choice. We found that adult male density had strong effects on juvenile perch height, perch width, and substrate use, suggesting that age‐class competition contributes to the observed ontogenetic differences in habitat choice. We also found that time of day significantly affected juvenile perch height and substrate use. In many cases, our results suggest that juveniles distance themselves from adults using different microhabitats from those used in our control ‘no‐adult’ treatment. However, these findings were often body size dependent and varied depending upon time of day. This study highlights the complexity of juvenile perching behavior and demonstrates the role of intraspecific interactions in shaping habitat use by juvenile animals.     

Agonistic Behaviour and Development of Territoriality During Ontogeny of the Sea Anemone Dwelling Crab Allopetrolisthes Spinifrons (H. Milne Edwards, 1837) (Decapoda: Anomura: Porcellanidae)

Allopetrolisthes spinifrons is an ectosymbiotic crab of the sea anemone Phymactis clematis. As a consequence of low host abundance, these represent a scarce and limited resource for the crab. Additionally, the relatively small size of the sea anemone host suggests that few symbiotic crabs can cohabit on one host individual, forcing crabs to adopt a territorial behaviour. In order to examine the potential presence and ontogenetic development of territoriality, the agonistic behaviour between crabs of various ontogenetic stages (adults, juveniles, and recruits) was studied in the laboratory. Laboratory experiments demonstrated that adult or juvenile crabs aggressively defended their sea anemone hosts against adult or juvenile intruders, respectively, but both adult and juvenile crabs tolerated recruits. Adult crabs behaved indifferently towards juvenile crabs, sometimes tolerating them, sometimes expelling them. Recruits never showed agonistic behaviour among themselves. The agonistic interactions observed in the laboratory and the uniform population distribution pattern on sea anemones recently described for A. spinifrons indicate that this species exhibits territorial behaviour, which develops during ontogeny. Territoriality in this species and other symbiotic decapods may function as a density-dependent mechanism of population regulation, being mediated by the availability of hosts. Resource monopolisation behaviours may be common among other symbiotic and free-living marine invertebrates inhabiting discrete habitats that represent a limiting resource.