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Journal: Animal behaviour

253

Body size and metabolic rate both fundamentally constrain how species interact with their environment, and hence ultimately affect their niche. While many mechanisms leading to these constraints have been explored, their effects on the resolution at which temporal information is perceived have been largely overlooked. The visual system acts as a gateway to the dynamic environment and the relative resolution at which organisms are able to acquire and process visual information is likely to restrict their ability to interact with events around them. As both smaller size and higher metabolic rates should facilitate rapid behavioural responses, we hypothesized that these traits would favour perception of temporal change over finer timescales. Using critical flicker fusion frequency, the lowest frequency of flashing at which a flickering light source is perceived as constant, as a measure of the maximum rate of temporal information processing in the visual system, we carried out a phylogenetic comparative analysis of a wide range of vertebrates that supported this hypothesis. Our results have implications for the evolution of signalling systems and predator-prey interactions, and, combined with the strong influence that both body mass and metabolism have on a species' ecological niche, suggest that time perception may constitute an important and overlooked dimension of niche differentiation.

Concepts: Psychology, Evolution, Metabolism, Energy, Organism, Ecology, Information, Niche differentiation

37

Following human gaze in dogs and human infants can be considered a socially facilitated orientation response, which in object choice tasks is modulated by human-given ostensive cues. Despite their similarities to human infants, and extensive skills in reading human cues in foraging contexts, no evidence that dogs follow gaze into distant space has been found. We re-examined this question, and additionally whether dogs' propensity to follow gaze was affected by age and/or training to pay attention to humans. We tested a cross-sectional sample of 145 border collies aged 6 months to 14 years with different amounts of training over their lives. The dogs' gaze-following response in test and control conditions before and after training for initiating eye contact with the experimenter was compared with that of a second group of 13 border collies trained to touch a ball with their paw. Our results provide the first evidence that dogs can follow human gaze into distant space. Although we found no age effect on gaze following, the youngest and oldest age groups were more distractible, which resulted in a higher number of looks in the test and control conditions. Extensive lifelong formal training as well as short-term training for eye contact decreased dogs' tendency to follow gaze and increased their duration of gaze to the face. The reduction in gaze following after training for eye contact cannot be explained by fatigue or short-term habituation, as in the second group gaze following increased after a different training of the same length. Training for eye contact created a competing tendency to fixate the face, which prevented the dogs from following the directional cues. We conclude that following human gaze into distant space in dogs is modulated by training, which may explain why dogs perform poorly in comparison to other species in this task.

Concepts: Time, Human, Training, Ageing, Explanation, Face, Dog breed, Border Collie

7

Animals can insure themselves against the risk of starvation associated with unpredictable food availability by storing energy reserves or gathering information about alternative food sources. The former strategy carries costs in terms of mass-dependent predation risk, while the latter trades off against foraging for food; both trade-offs may be influenced by an individual’s developmental history. Here, we consider a possible role of early developmental experience in inducing different mass regulation and foraging strategies in European starlings. We measured the body mass, body condition, foraging effort, food consumption and contrafreeloading (foraging for food hidden in sand when equivalent food is freely available) of adult birds (≥10 months old) that had previously undergone a subtle early life manipulation of food competition (cross-fostering into the highest or lowest ranks in the brood size hierarchy when 2-12 days of age). We found that developmentally disadvantaged birds were fatter in adulthood and differed in foraging behaviour compared with their advantaged siblings. Disadvantaged birds were hyperphagic compared with advantaged birds, but only following a period of food deprivation, and also spent more time contrafreeloading. Advantaged birds experienced a trade-off between foraging success and time spent contrafreeloading, whereas disadvantaged birds faced no such trade-off, owing to their greater foraging efficiency. Thus, developmentally disadvantaged birds appeared to retain a phenotypic memory of increased nestling food competition, employing both energy storage and information-gathering insurance strategies to a greater extent than their advantaged siblings. Our results suggest that subtle early life disadvantage in the form of psychosocial stress and/or food insecurity can leave a lasting legacy on foraging behaviour and mass regulation even in the absence of food insufficiency during development or adulthood.

Concepts: Energy, Mass, Food, Food security, Foraging, Famine, European Starling, The Brood

5

Host-parasite interactions are a key determinant of the population dynamics of wild animals, and behaviours that reduce parasite transmission and infection may be important for improving host fitness. While antiparasite behaviours have been demonstrated in laboratory animals and domesticated ungulates, whether these behaviours operate in the wild is poorly understood. Therefore, examining antiparasite behaviours in natural populations is crucial for understanding their ecological significance. In this study, we examined whether two wild rodents (white-footed mice, Peromyscus leucopus, and deer mice, Peromyscus maniculatus), selectively foraged away from conspecific faeces or avoided faeces altogether, and whether faecal gastrointestinal parasite status affected their behaviour. We also tested whether wild mice, when nesting, avoided using material that had previously been used by healthy or parasite-infected conspecifics. Our results, in contrast to laboratory mouse studies, suggest that wild mice do not demonstrate faecal avoidance, selective foraging or selective use of nesting material; they preferred being near faeces and did not differentiate between faeces from parasitized and uninfected conspecifics. Behavioural avoidance to reduce parasite infection may still represent an important strategy; however, mice in our study population appeared to favour the opportunity to feed and nest over the risks of coming into contact with faecal-transmitted parasites. Furthermore, the presence of conspecific faeces may actually provide a positive cue of a good foraging or nesting location. Ultimately, balancing the trade-off of performing antiparasite behaviours to reduce infection with missing an important feeding or nesting opportunity may be very different for animals in the wild facing complex and stochastic environments.

Concepts: Population, Behavior, Human behavior, Mouse, Peromyscus, Peromyscus maniculatus, Neotominae, White-footed Mouse

4

•We previously selected for large and small brain size in guppies.•Large-brained females outperformed small-brained females in a learning task.•Healy and Rowe challenged our interpretations of larger brains = better learning.•Here we argue why we think they are mistaken.

Concepts: Psychology, Brain, Cognition, Cognitive science, Mind, Thought, Benefit of clergy

4

Recent evidence has shown that humans are remarkably sensitive to artificial cues of conspecific observation when making decisions with potential social consequences. Whether similar effects are found in other great apes has not yet been investigated. We carried out two experiments in which individual chimpanzees, Pan troglodytes, took items of food from an array in the presence of either an image of a large conspecific face or a scrambled control image. In experiment 1 we compared three versions of the face image varying in size and the amount of the face displayed. In experiment 2 we compared a fourth variant of the image with more prominent coloured eyes displayed closer to the focal chimpanzee. The chimpanzees did not look at the face images significantly more than at the control images in either experiment. Although there were trends for some individuals in each experiment to be slower to take high-value food items in the face conditions, these were not consistent or robust. We suggest that the extreme human sensitivity to cues of potential conspecific observation may not be shared with chimpanzees.

Concepts: Human, Hominidae, Chimpanzee, Gorilla, Ape, Bonobo, Common Chimpanzee, Homininae

2

In most primate societies, strong and enduring social bonds form preferentially among kin, who benefit from cooperation through direct and indirect fitness gains. Chimpanzees, Pan troglodytes, differ from most species by showing consistent female-biased dispersal and strict male philopatry. In most East African populations, females tend to forage alone in small core areas and were long thought to have weak social bonds of little biological significance. Recent work in some populations is challenging this view. However, challenges remain in quantifying the influence of shared space use on association patterns, and in identifying the drivers of partner preferences and social bonds. Here, we use the largest data set on wild chimpanzee behaviour currently available to assess potential determinants of female association patterns. We quantify pairwise similarities in ranging, dyadic association and grooming for 624 unique dyads over 38 years, including 17 adult female kin dyads. To search for social preferences that could not be explained by spatial overlap alone, we controlled for expected association based on pairwise kernel volume intersections of core areas. We found that association frequencies among females with above-average overlap correlated positively with grooming rates, suggesting that associations reflected social preferences in these dyads. Furthermore, when available, females preferred kin over nonkin partners for association and grooming, and variability was high among nonkin dyads. While variability in association above and below expected values was high, on average, nonkin associated more frequently if they had immature male offspring, while having female offspring had the opposite effect. Dominance rank, an important determinant of reproductive success at Gombe, influenced associations primarily for low-ranking females, who associated preferentially with each other. Our findings support the hypothesis that female chimpanzees form well-differentiated social relationships that are of potential adaptive value to females and their offspring.

Concepts: Human, Male, Reproduction, Female, Sociology, Sex, Chimpanzee, Common Chimpanzee

2

Friend of a friend relationships, or the indirect connections between people, influence our health, well-being, financial success and reproductive output. As with humans, social behaviours in other animals often occur within a broad interconnected network of social ties. Yet studies of animal social behaviour tend to focus on associations between pairs of individuals. With the increase in popularity of social network analysis, researchers have started to look beyond the dyad to examine the role of indirect connections in animal societies. Here, I provide an overview of the new knowledge that has been uncovered by these studies. I focus on research that has addressed both the causes of social behaviours, i.e. the cognitive and genetic basis of indirect connections, as well as their consequences, i.e. the impact of indirect connections on social cohesion, information transfer, cultural practices and fitness. From these studies, it is apparent that indirect connections play an important role in animal behaviour, although future research is needed to clarify their contribution.

Concepts: Psychology, Sociology, Social relation, Social network, Knowledge management, Network theory, Mathematical sociology, Structural cohesion

2

For many long-lived mammalian species, extended maternal investment has a profound effect on offspring integration in complex social environments. One component of this investment may be aiding young in aggressive interactions, which can set the stage for offspring social position later in life. Here we examined maternal effects on dyadic aggressive interactions between immature (<12 years) chimpanzees. Specifically, we tested whether relative maternal rank predicted the probability of winning an aggressive interaction. We also examined maternal responses to aggressive interactions to determine whether maternal interventions explain interaction outcomes. Using a 12-year behavioural data set (2000-2011) from Gombe National Park, Tanzania, we found that relative maternal rank predicted the probability of winning aggressive interactions in male-male and male-female aggressive interactions: offspring were more likely to win if their mother outranked their opponent's mother. Female-female aggressive interactions occurred infrequently (two interactions), so could not be analysed. The probability of winning was also higher for relatively older individuals in male-male interactions, and for males in male-female interactions. Maternal interventions were rare (7.3% of 137 interactions), suggesting that direct involvement does not explain the outcome for the vast majority of aggressive interactions. These findings provide important insight into the ontogeny of aggressive behaviour and early dominance relationships in wild apes and highlight a potential social advantage for offspring of higher-ranking mothers. This advantage may be particularly pronounced for sons, given male philopatry in chimpanzees and the potential for social status early in life to translate more directly to adult rank.

Concepts: Sociology, Emergence, Chimpanzee, Mother, Social status, Epistasis, Jane Goodall, Gombe Stream National Park

2

Although in anurans the predominant mode of intra- and intersexual communication is vocalization, modalities used in addition to or instead of acoustic signals range from seismic and visual to chemical. In some cases, signals of more than one modality are produced through or by the anuran vocal sac. However, its role beyond acoustics has been neglected for some time and nonacoustic cues such as vocal sac movement have traditionally been seen as an epiphenomenon of sound production. The diversity in vocal sac coloration and shape found in different species is striking and recently its visual properties have been given a more important role in signalling. Chemosignals seem to be the dominant communication mode in newts, salamanders and caecilians and certainly play a role in the aquatic life phase of anurans, but airborne chemical signalling has received less attention. There is, however, increasing evidence that at least some terrestrial anuran species integrate acoustic, visual and chemical cues in species recognition and mate choice and a few secondarily mute anuran species seem to fully rely on volatile chemical cues produced in glands on the vocal sac. Within vertebrates, frogs in particular are suitable organisms for investigating multimodal communication by means of experiments, since they are tolerant of disturbance by observers and can be easily manipulated under natural conditions. Thus, the anuran vocal sac might be of great interest not only to herpetologists, but also to behavioural biologists studying communication systems.

Concepts: Species, Sexual selection, Sound, Tetrapod, Amphibian, Frog, Salamander