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Animal Behaviour
Journal Prestige (SJR): 1.58
Citation Impact (citeScore): 3
Number of Followers: 205  
  Hybrid Journal Hybrid journal (It can contain Open Access articles)
ISSN (Print) 0003-3472 - ISSN (Online) 1095-8282
Published by Elsevier Homepage  [3158 journals]
  • Can behaviour explain invasion success' A comparison between sympatric
           invasive and native lizards
    • Abstract: Publication date: Available online 16 April 2019Source: Animal BehaviourAuthor(s): Isabel Damas-Moreira, Julia L. Riley, D. James Harris, Martin J. WhitingTo reduce the impact of biological invasions, we need to understand the behavioural mechanisms that enable some species to be successful invaders. Testing differences in behaviour between sympatric congeneric species with different invasive potential is an opportunity to study specific behavioural traits associated with invasion success. Using the invasive Italian wall lizard, Podarcis sicula, and a noninvasive congeneric, the green Iberian wall lizard, Podarcis virescens, which live in sympatry in a location that is novel for P. sicula, we tested their exploratory behaviour, neophobia and boldness: all traits that should promote invasion success. The invasive P. sicula was more exploratory, bold and neophilic than the sympatric native P. virescens. Native lizards had highly repeatable behaviour, whereas in P. sicula boldness was the only behavioural trait that was repeatable. The behavioural traits of the native species, but not the invasive species, were correlated. A lack of correlation between behavioural traits, as well as a lack of repeatability in two of the three behavioural traits, suggests higher levels of behavioural plasticity in P. sicula, which may also explain the success of this lizard during invasions. Our experiment highlights the potential importance of behavioural traits in invasions and provides insight into why P. sicula is such a successful invader.
  • Affiliation and disease risk: social networks mediate gut microbial
           transmission among rhesus macaques
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Krishna N. Balasubramaniam, Brianne A. Beisner, Josephine A. Hubbard, Jessica J. Vandeleest, Edward R. Atwill, Brenda McCowanIn social animals, affiliative behaviours bring many benefits, but also costs such as disease risk. The ways in which affiliation may affect the risk of infectious agent transmission remain unclear. Moreover, studies linking variation in affiliative interactions to infectious agent incidence/diversity have speculated that disease transmission may have occurred, rather than revealing that transmission did occur. We address these gaps using the phylogenetics of commensal gut Escherichia coli to determine whether affiliative grooming and huddling social networks mediated microbial transmission among rhesus macaques. We collected behavioural and microbial data from adult macaques across a 12-week period that was split into two 6-week phases to better detect dyadic transmission. We reconstructed undirected social networks from affiliative interactions and reconstructed microbial transmission networks from the pairwise phylogenetic similarity of E. coli pulsotypes from macaques within and across adjacent sampling events. Macaque E. coli pulsotypes were more phylogenetically similar to each other than to environmental isolates, which established a premise for socially mediated transmission. Dyadic grooming and huddling frequencies strongly influenced the likelihood of E. coli transmission during the second data collection phase, but not the first. Macaques that were more central/well connected in both their grooming and huddling networks were also more central in the E. coli transmission networks. Our results confirmed that affiliative grooming and huddling behaviours mediate the transmission of gut microbes among rhesus macaques, particularly among females and high-ranking individuals. The detectability of socially mediated E. coli transmission maybe partially masked by environmental acquisition in males, or by high frequencies of interactions in captivity. Predicting the potential transmission pathways of gastrointestinal parasites and pathogens, our findings add to current knowledge of the coevolutionary relationships between affiliative behaviour and health and may be used to identify ‘superspreader’ individuals as potential targets for disease control strategies.
  • Maternal age influences offspring behaviour and growth efficiency during
           provisioning in northern elephant seals
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Amanda W. Hooper, Ryan W. Berger, Lovisa S. Rubin, Birgitte I. McDonald, Daniel E. CrockerOffspring growth rates impact fitness and can be influenced by maternal effects. Despite efforts to understand the influence of maternal traits (e.g. age, size, body condition) on reproductive effort, much less is known about how maternal traits and environment influence the behaviour of offspring and ultimately, how offspring behaviour may influence the efficiency of the translation of maternal investment into offspring growth. Offspring of capital breeders, such as the northern elephant seal, Mirounga angustirostris, are nursed exclusively from maternal body reserves and investment is limited by the resources acquired prior to parturition. Combined new and previously published milk energy intake and offspring storage data (N = 38) suggest impacts of maternal age on growth efficiency that are independent of rates of energy delivery. To determine the mechanisms underlying this effect of maternal age, behavioural data were collected from pups of 46 known-age females, from parturition to weaning, across 6 years and three different sites along the central California coast, representing 3954 seal-hours of observation. Pup behaviours were divided into five mutually exclusive categories that potentially impacted pup energetics. The offspring of older females spent more time resting, while offspring of younger females spent more time locomoting and distant from their mother. As pups developed, they spent more time suckling and locomoting and less time resting. Pup behaviour showed strong diel patterns, with activity decreasing over the day. The magnitude of these relationships varied between rookeries, suggesting influences of harem size, topography or environmental features on pup behaviour. Together these findings suggest direct impacts of maternal age and breeding experience on pup behaviour and growth efficiency.
  • Anthropogenic noise pollution reverses grouping behaviour in hermit crabs
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Svenja Tidau, Mark BriffaNoise is a form of human-induced rapid environmental change, and mounting evidence suggests that it can affect the sensory environment and consequently the decision-making ability of animals. However, while the effects of anthropogenic noise on individual organisms in the context of movement patterns, foraging and predation risk have been reported, relatively little is known about how noise impacts groups and intraspecific interactions. Here we investigated the effects of anthropogenic noise on grouping preference (i.e. being with conspecifics or alone) in the European hermit crab, Pagurus bernhardus. Hermit crabs live in empty gastropod shells and frequently fight with each other to gain an optimal-fitting shell. Thus, crabs' grouping preference may depend on the optimality of their own shell and thus on their motivation to gain another. To test the effect of shell size and its interaction with noise exposure on grouping preferences, crabs were housed in either suboptimal or optimal shells before being exposed to playbacks of either ship noise or an ambient sound (control) and given the choice to group with one or five conspecifics or to remain alone in a neutral zone. Crabs occupying suboptimal shells had a longer latency to enter the zone with a single crab than crabs in optimal shells. This difference was only seen in the ambient sound treatment, disappearing completely under ship noise. Under ambient sound, crabs in optimal shells spent most of their time close to a single crab, while crabs in suboptimal shells showed no clear preference. However, exposure to ship noise reversed the effect of shell quality on grouping preference. Our results demonstrate that exposure to anthropogenic noise can alter not only individual behaviour but also social behaviour.
  • Keep calm, we know each other: kin recognition affects aggressiveness and
           conflict resolution in a solitary parasitoid
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Anthony G.E. Mathiron, Patrice Pottier, Marlène GoubaultIntraspecific competition for indivisible resources can trigger the expression of agonistic behaviour in individuals of many animal species. Aggressiveness and conflict resolution may be influenced by the value individuals place on the resource (subjective resource value) but also by genetic relatedness between competitors. The ability to differentiate genetically related from unrelated individuals (i.e. kin recognition) can play a key role in the dynamics of agonistic interactions between individuals. In this context, the theory of kin selection predicts that competitors should display fewer aggressive behaviours towards closely related individuals. Recognition of kin can be driven by the perception of (1) genetically linked phenotypic cues (phenotypic matching) and/or (2) environmental cues (familiarity). In the hymenopteran solitary parasitoid Eupelmus vuilleti, individuals develop on larvae and pupae of their host, Callosobruchus maculatus, which infest cowpea seeds, Vigna unguiculata. Eupelmus vuilleti females can fight for the host on which they lay their eggs. Here, we investigated the effect of genetic relatedness (genotype) and familiarity (the seed containing the host on which they develop as juveniles) on aggressiveness and contest outcome over hosts in E. vuilleti females. We first demonstrated that the probability of a conflict escalating was affected by the interaction between genetic relatedness and familiarity among females. We then found that familiarity alone affected the likelihood of contest resolution. The occurrence of escalated conflicts was reduced between related and familiar females, and contests were more likely to be clearly resolved when occurring between familiar competitors. Our results highlight a parasitoid wasp's abilities to identify and discriminate kin, showing for the first time that two kin recognition components can interact in mediating competition avoidance for resource access in a solitary insect species.
  • Evolutionary roads to syntax
    • Abstract: Publication date: Available online 10 April 2019Source: Animal BehaviourAuthor(s): Klaus ZuberbühlerSyntax is habitually named as what sets human language apart from other communication systems, but how did it evolve' Comparative research on animal behaviour has contributed in important ways, with mainly three sets of data. First, animals have been subjected to artificial grammar tasks, based on the hypothesis that human syntax has evolved through advanced computational capacity. In these experiments humans generally outperform animals, but there are questions about validity, as experimental stimuli are (deliberately) kept devoid of semantic content. Second, animal communication has been compared in terms of the surface structures with the aim of developing a typology of animal syntax, based on the hypothesis that syntax is an evolutionary solution to the constraints of small signal repertoires. A wide range of combinatorial phenomena has been described, mainly in nonhuman primates, but there is little support for the hypothesis that syntax has emerged due to repertoire size constraints. A third way of studying the evolution of syntax is to compare how animals perceive and communicate about external events, the mental deep structure of syntax. Human syntax is closely aligned with how we perceive events in terms of agency, action and patience, each with subsidiary functions. The event perception hypothesis has been least explored in animals and requires a serious research programme.
  • Male body size, dominance rank and strategic use of aggression in a
           group-living mammal
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Edward Wright, Jordi Galbany, Shannon C. McFarlin, Eric Ndayishimiye, Tara S. Stoinski, Martha M. RobbinsBody size is a key determinant of male fighting ability and reproductive success in many animal species, but relationships between these variables have only rarely been examined in group-living animals in which body size often correlates with dominance rank. We examined the relationships between body size (crest height, back breadth and body length), dominance rank, alpha male tenure length, number of adult females and patterns of aggression in 26 wild adult male mountain gorillas, Gorilla beringei beringei, living in multimale groups. A composite measure combining crest height and back breadth (variables were highly correlated and combined into a crest–back score), but not body length, significantly correlated with dominance rank, alpha male tenure length and number of adult females per group. The alpha male had the largest crest–back score in six of the seven groups, and in the majority of dyads the male with the higher crest–back score was higher ranking. The frequency (and intensity on mating days) of aggressive contests was higher between males close in rank. Additionally, aggression occurred more frequently when the initiator was larger than the recipient. Our results suggest that factors other than body size are likely to influence dominance rank, but large size helps males attain and retain high dominance rank, probably leading to greater reproductive success. Further studies on how the timing and intensity of male–male competition influences life history trade-offs between investment in secondary sexual characteristics, body condition and survival may explain variance in lifetime reproductive success within and between species.
  • Nutrition during sexual maturation and at the time of mating affects
           mating behaviour in both sexes of a burying beetle
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Jon Richardson, Per T. SmisethTheory predicts that the outcome of mating interactions should be influenced by the condition of both males and females. First, females should base their mating decisions on reliable cues about male quality, which are often condition dependent. Second, the costs and/or benefits of being choosy during mating may depend on the female's own condition. Finally, when males divide their time between different mating tactics, investment in alternative mating tactics may depend on male condition. Here we examined the effects of male and female nutritional condition on mating behaviour in the burying beetle Nicrophorus vespilloides. We manipulated male and female nutritional condition either during sexual maturation or at the time of mating and monitored female mate choice and male mating tactics. We found that females in poor condition (i.e. starved either during sexual maturation or at the time of mating) preferred to mate with males in good condition over males that were starved at the time of mating. In contrast, well-fed females showed no such preference. Furthermore, males that were starved during sexual maturation increased their investment in alternative mating tactics by spending more time signalling for females. Our results add to evidence suggesting that females in poor condition bias mating towards males in good condition although it is currently unclear why these females are choosier in this species. Ours is the first study to demonstrate that nutritional condition during sexual maturation can influence mating behaviour, which may have implications for the rate and direction of sexual selection.
  • Chemical profiles reflect heterozygosity and seasonality in a tropical
           lekking passerine bird
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Danielle J. Whittaker, Meredith Kuzel, Mikayla J.E. Burrell, Helena A. Soini, Milos V. Novotny, Emily H. DuValMany lekking systems exhibit highly skewed male reproductive success, but traits preferred by females are not always evident. In the lance-tailed manakin, Chiroxiphia lanceolata, male reproductive success is correlated with age, experience and heterozygosity, but mechanisms by which females might detect these qualities are unknown. Avian chemical signals, such as volatile compounds present in preen oil secreted by the uropygial gland, have been shown to predict reproductive success, correlate with genetic diversity and indicate body condition and immune function in temperate passerines. Here we tested whether preen oil volatile compounds contain information about individual characteristics related to mate choice in male and female lance-tailed manakins, including social and breeding status, heterozygosity and age. In males, preen oil volatile profiles reflected heterozygosity but not social status or age. In females, we found a relationship between volatile profiles and age but not breeding status or heterozygosity. Volatile compounds covaried with sampling date in both sexes. The information reflected in manakin volatile profiles suggests that they have the potential to serve as mate assessment cues in this species.
  • Animal communication, cognition, and the evolution of language
    • Abstract: Publication date: Available online 3 April 2019Source: Animal BehaviourAuthor(s): William A. Searcy
  • Association Page
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s):
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    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s):
  • Impact of long-term behavioural studies in the wild: the blue petrel,
           Halobaena caerulea, case at Kerguelen
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Matthieu Bergès, Rémi Choquet, Francesco BonadonnaThe study of animal behaviour contributes to our understanding of how individuals or populations live and survive. To this purpose researchers often manipulate, in a broad sense, wild animals in sensitive periods, such as breeding, when animals are accessible/available. Few studies, however, show the impact of these manipulations on the survival of individuals. In this study we measured short- and long-term impacts of experimental manipulations on a small colony of blue petrels at Kerguelen archipelago where behavioural studies have been performed since 2006. We developed two models to measure potential impacts of manipulation: (1) a new multievent model that allowed us to account for different severity classes of experimental manipulation independently and (2) a model that allowed us to consider the cumulative impact over several years of these experimental manipulations. We found no evidence that our experimental manipulations have negatively affected survival or breeding probabilities either in the short or in the long term. Conversely, similarly to other capture–recapture studies on blue petrels, survival was shown to be dependent on the birds' experience (birds that probably bred for the first time versus birds that had already bred several times before) and the breeding probability to be dependent on the year, possibly because of environmental conditions.
  • Wild zebra finches choose neighbours for synchronized breeding
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Hanja B. Brandl, Simon C. Griffith, Wiebke SchuettOrganisms should aim to time their reproduction to match the optimal ecological conditions and thus maximize their fitness. However, social cues have been identified as determinants of reproductive decisions and might also be involved in coordinating the timing of reproduction. Breeding synchronously with other individuals can bring several advantages, including a reduced individual predation risk and an increased opportunity for social foraging. The behavioural mechanisms underlying reproductive synchrony are versatile and not well understood, particularly in species inhabiting unpredictable environments. In contrast to highly seasonal environments, more variable and unpredictable environments can support periods of extended breeding with lower levels of synchronous breeding overall, but opportunities for individuals to breed synchronously at a finer temporal and spatial scale. Zebra finches, Taeniopygia guttata, are a highly social species, naturally inhabiting the ecologically unpredictable arid zone of Australia. In the wild, reproduction at a broad population level is not highly synchronized and, at any time during a period of breeding activity, reproductive attempts can be found at different stages. However, previous work has suggested that at a finer spatial scale neighbours tend to breed at approximately the same time. Using nestboxes, we tested whether wild zebra finches preferentially seek to settle and initiate a breeding attempt adjacent to conspecifics at an early stage of breeding (nest building), as opposed to others at later stages of breeding and with which the opportunity to breed synchronously was reduced or absent. Pairs were more likely to initiate egg laying in nestboxes close to conspecifics at an early stage of breeding, suggesting that they do try to maximize the level of synchronicity with neighbours. Our results indicate the importance of social effects on both the phenology and spatial distribution of breeding.
  • Early vocal ontogeny in a polytocous mammal: no evidence of social
           learning among sibling piglets, Sus scrofa
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Marek Špinka, Michaela Syrová, Richard Policht, Pavel LinhartAnimals living in social proximity often have similar vocalizations. For many bird and several mammal species, at least part of the vocal similarity is socially learned during ontogeny. Little is known, however, about the ontogenetic origin of vocal similarities among siblings in polytocous mammals. We investigated the influence of social environment and genetic relatedness on the development of acoustic similarities among suckling piglets. To examine whether the common acoustic features are innate or learned by postnatal vocal convergence in the same litter, we cross-fostered piglets among pairs of mother sows immediately after birth and recorded contact calls (grunts) of both the cross-fostered and the noncross-fostered piglets during the suckling period. Acoustic distances of the cross-fostered piglets to their new littermates remained longer than those among noncross-fostered siblings and were as long as those between piglets from different litters. The results show that after being neonatally cross-fostered to another litter, the piglets did not converge acoustically with their new littermates even after several weeks of cohabitation. This is in contrast to the presence of vocal plasticity during the ontogeny of other mammals including other ungulates, indicating that use of vocal learning may vary even in closely related species, perhaps in relation to its adaptive utility within the life history and social organization of the species.
  • The effect of experience and olfactory cue in an inhibitory control task
           in guppies, Poecilia reticulata
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Maria Santacà, Melania Busatta, Beste Başak Savaşçı, Tyrone Lucon-Xiccato, Angelo BisazzaBehavioural responses to the environment often require the suppression of strong internal predispositions or the overriding of external lures, tasks performed by a cognitive function called inhibitory control. Inhibitory control of nonhuman animals is generally measured with the cylinder task: subjects are presented with food inside a transparent cylinder and must inhibit their tendency to reach the food directly and instead detour round the cylinder to solve the task. However, several studies have raised concerns about the validity of this test to compare different species. Recently, a tiny teleost fish, the guppy, Poecilia reticulata, was tested with the cylinder task and scored higher than many mammals and birds. Before considering that guppies display unpredictably high inhibitory control, it is important to exclude that they were advantaged by some noncognitive factors. In particular, guppies could enjoy three advantages: experience with transparent surfaces (i.e. the walls of maintenance aquaria), experience with detouring round see-through obstacles (vegetation in the maintenance aquaria) and the spread of food odour in the water from the sides of the cylinder. We tested whether these factors affect guppies' performance in the cylinder task by manipulating both their experience with transparent surfaces before the task and the diffusion of food odour cues from the cylinder. Guppies raised in transparent aquaria or with transparent panels placed inside the tank did not show advantages over guppies with no experience with transparent surfaces. Furthermore, the guppies’ performance was not reduced when the cylinder was pierced in the middle, so that both visual and olfactory cues lured them in the same direction. These results seem to exclude methodological explanations for the high inhibitory control score of guppies, and they indicate that even teleost fish can display efficient inhibitory control.
  • Group-enhanced predator detection and quality of vigilance in a social
           ground squirrel
    • Abstract: Publication date: May 2019Source: Animal Behaviour, Volume 151Author(s): Annemarie van der Marel, Marta López-Darias, Jane M. WatermanAnimals may form groups for different reasons, and one major benefit of grouping in many species is reduced predation risk. In diurnal species, vigilance is used to detect predators, resulting in a trade-off between feeding activity and predation risk. Species can reduce the cost of this trade-off with low-quality vigilance – performing another behaviour while vigilant – in comparison to high-quality vigilance (only being vigilant). Two nonmutually exclusive hypotheses explaining an inverse relationship between individual vigilance and group size are the dilution effect, where predation risk decreases in larger groups, and collective detection, where larger groups have more individuals that may detect a predator. Two predictions that support collective detection but not the dilution effect are that (1) overall group (collective) vigilance will increase with increasing group size, even while individual vigilance decreases, and (2) at least one group member must be vigilant to detect potential danger and communicate that information to group members. To test these predictions, we recorded behavioural data on low- and high-quality vigilance and alarm calling in the gregarious Barbary ground squirrel, Atlantoxerus getulus. Barbary ground squirrels allocated more time to high-quality vigilance than low-quality vigilance. The collective detection hypothesis was partly supported: as group size increased, individual low- and high-quality vigilance did not decrease, but collective high-quality vigilance did increase. Furthermore, we found that repetitive alarm calling warned group members of terrestrial threats. Our results show that this invasive species displays specific antipredator behaviours to different aerial and terrestrial predators compared to predators in their endemic range. The low level of time allocated to low-quality vigilance indicates that natural selection strongly favours high-quality vigilance in this species despite the trade-off with foraging. Our study broadens our understanding of antipredator and risk-sensitive behaviour.
  • The speech-like properties of nonhuman primate vocalizations
    • Abstract: Publication date: Available online 29 March 2019Source: Animal BehaviourAuthor(s): Thore J. Bergman, Jacinta C. Beehner, Melissa C. Painter, Morgan L. GustisonThe origins of speech, the most complex form of animal communication, remain a puzzle. Human speech and nonhuman primate vocalizations have traditionally been viewed dichotomously, with several aspects of speech having no clear analogues in the calls of our primate relatives. The putative unique aspects of speech include a diverse array of learned sounds that are rapidly produced in rhythmic strings and continuously recombined in new sequences. However, recent research challenges the idea that these features are indeed unique to humans and suggests more continuity between nonhuman and human primates than was previously appreciated. Here we review recent findings in four areas of this emerging continuity. In light of these studies, we argue that the evolution of human speech abilities most likely originated in a primate ancestor capable of (1) producing a ‘speech-ready’ range of vowel-like sounds, (2) vocalizing with simultaneous rhythmic mouth movements, (3) combining long strings of varied and structured sounds and (4) exercising some volitional control over calls that were modified based on experience. Taken together, these results suggest that the considerable latent vocal ability that we observe in nonhuman primates is consistent with the hypothesis that a key step towards human speech was the evolution of greater cognitive control of the vocal apparatus (and not the evolution of speech-specific anatomical adaptations). By shifting research emphasis away from the mechanics of how speech is produced to the conditions that favoured more diverse, open-ended and imitative vocal systems, we hope to encourage new avenues for future comparative research.
  • Personality types vary in their personal and social information use
    • Abstract: Publication date: Available online 27 March 2019Source: Animal BehaviourAuthor(s): Judith A.H. Smit, Kees van OersGathering information about the environment, such as the location and quality of food, is crucial for an animal's survival, particularly in a changing environment. An animal can collect ‘personal information’ by interacting with the environment itself, or it can collect ‘social information’ by observing the behaviour of others. The use of these two types of information varies across different situations and between individuals. Personality is a concept that captures consistent interindividual differences in behaviour and could be one of the factors driving interindividual variation in information use. We tested this by conducting behavioural experiments based on a colour association task in captive great tits, Parus major, originating from lines bidirectionally selected for high and low exploratory behaviour. We quantified personal information use by measuring to what extent a bird relied on previously rewarded options instead of novel options. Social information use was measured by recording how birds chose according to social information provided by video playbacks of a conspecific. Here, we demonstrate that variation in the use of both personal and social information is indeed personality related. In their decision making, slow explorers relied more on prior knowledge, from both personal and social origins, whereas fast explorers tended to ignore the available information and chose more randomly. The differences between the personality types imply different costs or constraints in acquiring and/or applying the two types of information, possibly due to variation in, for example, cognitive styles. In conclusion, we demonstrate that personality types have different strategies to cope with environmental uncertainty.
  • Social context modulates how the winner effect restructures territorial
           behaviour in free-living woodpeckers
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Meredith C. Miles, Matthew J. FuxjagerAnimal interactions such as competition are mediated by complex social strategies, which consist of behaviours and cognitive mechanisms that guide their production. As a result, behaviour is highly flexible. This poses a challenge to understanding how competition plays out in natural systems, because the course of a contest can essentially be rewritten by prior experience and/or changes in social context. Here we addressed this gap by studying how both of these factors interact to reconfigure competitive strategies used in territorial defence by a wild bird. After experimentally inducing the winner effect, a cognitive-behavioural phenomenon in which winning a contest increases the probability of winning again in the future, we found that male red-bellied woodpeckers, Melanerpes carolinus, adopted a new social strategy marked by more flexible transitions between different aggressive displays, as well as increasing overall aggressive output. However, this effect was mitigated by the arrival of the female social mate; in response to this momentary shift in social context, males decreased their use of territorial drum displays and became less likely to move around the territory or switch display modes during competition. In other words, the winner effect increased spatiotemporal diversity of territorial strategies, such that males frequently changed their location and display output. A female's arrival, however, reversed this effect. More specifically, males tended to revert to advertisement and social vocalizations on female arrival, which suggests that the need to attend to the social mate may supersede the threat of territorial intrusion. This is consistent with a model in which competitive outcomes are impacted by the interactive effects of an individual's past experiences and shifts in present-day social context, which may allow monogamous animals to effectively manage the competing demands of driving off intruders and attending to the social mate.
  • Behavioural syndromes as a link between ecology and mate choice: a field
           study in a reef fish population
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Sophie Picq, Marco Scotti, Oscar PueblaThe link between ecology and reproductive isolation constitutes the cornerstone of the ecological hypothesis of speciation. Such a link can arise when traits under ecologically based selection are also used as cues for mating (‘magic traits’) or as a by-product of habitat choice when mating takes place within habitats. Here, we propose that behavioural syndromes may also constitute such a link. We illustrate this mechanism in the butter hamlet, Hypoplectrus unicolor, a reef fish from the wider Caribbean, with aggressive mimicry as the focal ecological trait. Aggressive mimicry is of particular interest in hamlets since it has been proposed to play a key role in the radiation of Hypoplectrus. Individuals from a natural population in Bocas del Toro, Panama, were tagged and their diurnal and spawning behaviours observed over 2 years. The results indicate that aggressive mimicry behaviour differed consistently between individuals and formed two discrete behavioural types that also differed with respect to territoriality. Differences in territoriality between the two behavioural types translated into different use of space in spawning contexts, which generated a tendency for assortative mating by behavioural type. This case study illustrates how behavioural syndromes may form a link between ecologically relevant behavioural traits and mate choice, suggesting that they might play an underappreciated role in the early stages of speciation.
  • No evidence that male sexual experience increases mating success in a
           coercive mating system
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Maider Iglesias-Carrasco, Rebecca J. Fox, Alan Vincent, Megan L. Head, Michael D. JennionsSeveral studies have shown that sexual experience can alter a male's mating behaviour to increase his future mating success. One explanation is that experienced males are better at courting females and inducing them to mate. Experienced males might also be better at identifying higher quality mates, although fewer studies have tested for this benefit. In both cases, however, these potential benefits of sexual experience might be partially offset by the energetic costs of courting and mating, which tend to reduce a male's subsequent ability to invest in sexually selected traits, and thereby reduce his future attractiveness and mating success (i.e. hasten the onset of reproductive senescence). Here we used the eastern mosquitofish, Gambusia holbrooki, to test whether sexual experience elevates male mating success. We housed recently matured males either with full mating access to females (experienced males) or in the visual and olfactory presence of females with which they could not mate (naïve males). We then measured the strength of male mate choice for larger, more fecund, females, male mating behaviour (time spent chasing females and the number of copulation attempts) and insemination success. Experienced and naïve males did not differ significantly in their mating behaviour and there was no effect of sexual experience on the likelihood of mating or on the number of sperm inseminated (although experienced males had a tendency to be less successful when performing gonopodial thrusts). Experienced males in two-choice trials were, however, significantly more likely to ‘inspect’ both females and had a significantly stronger preference for larger females. Finally, we measured male immune response and growth to test for any costs of the increased mating effort that is concomitant with greater experience. Experienced males had significantly slower postmaturation growth and a significantly weaker immune response than naïve males.
  • Corrigendum to “Giraffe social preferences are context dependent”
           [Animal Behaviour 146 (2018) 37–49]
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Zoe Muller, Mauricio Cantor, Innes C. Cuthill, Stephen Harris
  • Mixed support for state maintaining risky personality traits in
           yellow-bellied marmots
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Matthew B. Petelle, Julien G.A. Martin, Daniel T. BlumsteinIn a variety of taxa, individuals behave in consistently different ways. However, there are relatively few studies that empirically test the potential mechanisms underlying the causes and maintenance of these personality differences. Several hypotheses for the causes and maintenance of risky personality traits have been suggested but all have received mixed support. Both the pace-of-life hypothesis and state-dependent safety hypothesis propose that differences in internal state cause and maintain personality traits. Formally, the pace-of-life hypothesis states that differences in life-history traits including productivity (growth) and residual reproductive value (age) create initial differences in individual behaviour that is later maintained by positive feedback, while the state-dependent safety hypothesis suggests that body condition (mass) is responsible for causing and maintaining behavioural differences. We tested and evaluated whether either of these two hypotheses explained the causes or maintenance of variation in risk-related personality traits –defensive aggression, activity and exploration– in yellow-bellied marmots, Marmota flaviventer. We found little support overall for these hypotheses in explaining maintenance in activity or exploration. However, for defensive aggression, we found positive feedback for both mass and age.
  • Black-tailed prairie dog, Cynomys ludovicianus (Sciuridae), metapopulation
           response to novel sourced conspecific signals
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Lauren C. Connell, Lauren M. Porensky, Anna D. Chalfoun, John D. ScastaAggregation of territorial individuals within a species can be facilitated via conspecific signals, wherein settlement implies habitat suitability, ease of resource acquisition and/or increased predator detection. The black-tailed prairie dog is a colonial small mammal with alarm vocalizations that confer benefits via group vigilance against predators and increased foraging time. Although prairie dog alarm calls are relatively well understood, the information embedded in their jump-yip call, which includes both a distinct cry and a bodily gesture, remains less clear. We evaluated prairie dog behaviour in response to conspecific acoustic signals using playbacks of alarm and jump-yip calls at 26 sites in northeastern Wyoming, U.S.A. Recorded calls from an isolated colony were broadcast to a mean of five individuals per site, and behavioural responses were compared against uninfluenced behaviour and a control playback of ambient sounds. The alarm playback caused prairie dogs to increase vigilance 122% and decrease foraging time 23%, demonstrating prairie dogs will shift behaviour based on signals from individuals of an unfamiliar colony. However, the alarm call playback reduced frequency of the jump-yip behaviour only at colonies nearest the recording source. The jump-yip playback caused unfamiliar prairie dogs to display 339% more jump-yips than uninfluenced behaviour. The jump-yip playback did not alter recipients' foraging or vigilance behaviours relative to control treatments, suggesting that although prairie dogs can understand and reciprocate an unfamiliar, single modality signal, they may not shift other behaviours based on this stimulus. As such, the purpose and benefits of the jump-yip call remain unclear. Playback efficacy also had a nonlinear relationship with distance from recording source. Our work improves understanding of communication at the metapopulation level, examines the potential role of the jump-yip and provides insights for how conspecific signals might be used as a management tool.
  • The scent of symbiosis: gut bacteria may affect social interactions in
           leaf-cutting ants
    • Abstract: Publication date: Available online 18 March 2019Source: Animal BehaviourAuthor(s): Serafino Teseo, Jelle S. van Zweden, Luigi Pontieri, Pepijn W. Kooij, Søren J. Sørensen, Tom Wenseleers, Michael Poulsen, Jacobus J. Boomsma, Panagiotis SapountzisAnimal gut microbiota affect host physiology and behaviour. In social insects, where colony level integrity is preserved via a nestmate discrimination system based on cuticular hydrocarbon mixtures, microorganismal effects may therefore influence social dynamics. Although nestmate recognition has undergone a thorough exploration during the last four decades, few studies have investigated the putative role of gut microbes. Here, we integrated 16S rRNA-based microbial community profiling, chemical and behavioural approaches to test whether gut microbes affect nestmate recognition in Acromyrmex echinatior leaf-cutting ants. Treating workers with a sterile diet or with antibiotics resulted in a substantial alteration of their gut microbial communities. In pairwise social interactions, untreated versus antibiotic-treated nestmates behaved more aggressively than other nestmate and non-nestmate pairs, suggesting that bacterial suppression may alter chemical social cues and trigger aggressive behaviour. Chemical analyses of treated individuals revealed a decrease in the abundance of two metapleural gland antifungal compounds, and confirmed the correspondence between aggression levels and chemical profile differences. Feeding microbiota-remodelled ants with conspecific faecal droplets partially restored the original bacterial communities. Ants fed with faecal droplets from different colonies were unusually aggressive compared to pairs fed with faecal droplets from the same colony. We cannot exclude confounding effects resulting from the potentially harmful action of antibiotics on ant hosts. However, our results suggest a correlation between chemical profiles and the presence of certain microbial species in the gut, which may affect nestmate recognition and division of labour. This opens novel questions about the role of symbiotic microorganisms in the evolution of social insect behaviour.
  • Evidence of high individual variability in seed management by
           scatter-hoarding rodents: does ‘personality’ matter'
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Mariano Feldman, Mariona Ferrandiz-Rovira, Josep Maria Espelta, Alberto MuñozThe predation and dispersal of seeds by scatter-hoarding animals is one of the most studied processes in the context of animal–plant interactions. Seed management by these animals has been traditionally approached at the population level: the patterns documented in the field are assumed to be similar for all individuals of the population and the variability within the population is considered to be random noise. However, little is known about to what extent this variability responds to different and consistent behaviours between individuals. The aim of this study was to analyse the individual variation and consistency in behaviour of scatter-hoarding rodents within a population. As our model we used the wood mouse, Apodemus sylvaticus, a key disperser of holm oak acorns, Quercus ilex, which, in turn, suffers high predation pressure by the common genet, Genetta genetta. In two sets of laboratory experiments, we compared the variance and consistency in behaviours and acorn management due to individual differences with that due to manipulation, using genet scents, of the perceived predation risk. Genet scents reduced the activity (i.e. time out of the refuge) in all wood mice, but the differences and consistency in activity between individuals accounted for most of the variance. Also, mice showed different and consistent stress or relaxed behaviours. Most of the variance in seed management variables, such as dispersal distance and seed size selection, was explained by consistent differences between individuals across scent treatments. The increase in stress behaviours and decrease in relaxed behaviours were positively related to dispersal ability (i.e. longer distances and larger acorns). Our study highlights the importance of considering the individual component of behaviour in scatter-hoarding rodents. This fine-scale level, largely overlooked in the ecological framework, will help to increase our understanding of seed management by scatter-hoarding animals.
  • The good fathers: efficiency of male care and the protective role of
           foster parents in a Neotropical arachnid
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Rosannette Quesada-Hidalgo, Diego Solano-Brenes, Gustavo S. Requena, Glauco MachadoThe evolution of exclusive paternal care in arthropods is influenced by both natural and sexual selection. Male care may simultaneously increase egg protection against natural enemies and male attractiveness to ovipositing females. When caring males desert or die, their clutches may be adopted either by females that provide flexible compensation of parental care or by males that may increase their own attractiveness caring for unrelated eggs. Whether foster parents are as efficient as the original owner males in protecting the clutch is a question that has rarely been addressed. Here we experimentally evaluated the efficiency of egg attendance provided by males of the mud-nest harvestman Quindina limbata. We also tested whether unattended nests are adopted by females and/or unrelated males, and compared the efficiency of the protection provided by foster parents with that provided by the original owner males. We found that when males were present inside the nest, nest visits by egg predators were much lower than in experimentally unattended nests. Ten conspecifics (8 males, 2 females) adopted experimentally unattended nests. Foster males were as efficient as the original owner males in decreasing nest visits by egg predators. The most important conclusions of our study are: (1) male protection is crucial for egg survival because unattended nests are promptly attacked by predators; (2) flexible compensation of parental care by females is rare; (3) males adopt unrelated nests and protect the eggs as efficiently as original owner males, probably because egg attendance is a sexually selected behaviour.
  • Dismantling Babel: creation of a universal calibration for honey bee
           waggle dance decoding
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Roger Schürch, Keiran Zwirner, Bethany J. Yambrick, Tiffanie Pirault, James M. Wilson, Margaret J. CouvillonNoisy animal communication presents a unique challenge for biologists interpreting a produced signal precisely and accurately. One past approach has been to reduce noise. Lately, with the advance of quantitative tools, an alternative is to use the noise to improve model predictions. Here we take the latter approach to improve our ability to recover honey bee (Apis mellifera spp.) foraging locations, in particular the distance at which they forage, from observed waggle dance communications, which encode the vector from the hive to the forage. This vector gives both the distance from the hive to the forage, which is encoded in dance duration, and the direction from the hive to the forage, which is encoded in the angle that the bee makes while dancing. We analysed the waggle dances from individually marked honey bees (N = 859 dances from 85 bees from 3 hives) that foraged at multiple known locations to determine a distance-to-duration calibration. Next we compared this calibration to a previously published calibration, which demonstrated that while their slopes were similar (P = 0.82), their intercepts differed (P 
  • Temporal and spatial pattern of trail clearing in the Australian meat ant,
           Iridomyrmex purpureus
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Eliza J.T. Middleton, Simon Garnier, Tanya Latty, Chris R. ReidMany ant species use trails to connect important resources. In addition, some ants actively clear obstacles from their trails. Although trail clearing is thought to be beneficial in decreasing travel time, the physical process of clearing requires an investment of time and energy. Given that trail clearing is a decentralized process, how do colonies decide when to invest in clearing' In this study, we examined trail clearing in the Australian meat ant, using artificial semipermeable barriers mimicking grass. We tested the hypothesis that investment in clearing was influenced by the abundance and physical toughness of obstacles and that the selection of which grass blade to cut was a nonrandom process that decreases travel distance. We found that low abundance/low toughness treatments experienced the greatest amount of clearing and high abundance/high toughness the least. Although ants did not clear an optimally efficient trail, the results of our percolation analysis support the inference that the ants strategically deployed clearing, taking multiple factors into account when deciding to invest in this strategy. The resultant clearing patterns provided shorter travel routes for foraging ants than would be expected by the random removal of obstacles.
  • Behavioural research priorities for the study of animal response to
           climate change
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Richard Buchholz, John D. Banusiewicz, Stephanie Burgess, Sarah Crocker-Buta, Lauren Eveland, Lauren FullerBehavioural traits are thought to be important determinants of the resilience of animal species to a rapidly changing global climate. Although increasing temperature has taken centre stage in the debate over climate change, animals will have to survive more than just extreme heat to persist in the Anthropocene. The aim of this review is to stimulate interest in the opportunities for integrative and applied behavioural study of how animals can survive life-threatening weather events, in order to help achieve the societal goal of maintaining viable wildlife populations under future climate scenarios. First, using the thermoregulatory behaviour of a hypothetical ground squirrel species as an example, we explore how different scenarios of behavioural flexibility, plasticity, adaptation, exaptation and management action can lead to population persistence or extinction. Next, we propose that considering weather events such as heatwaves, storms and floods, wildfire and drought as selective pressures worthy of investigation provides a new research framework for climate-related conservation behaviour. In our review we provide examples of the responses of animals to different types of weather extremes and describe behavioural adaptations to environments with extreme climates. We give methodological recommendations to jump start climate change research by behaviourists. Finally, we conclude with suggestions for using citizen science and a public video repository to foster evidence-based decision making for managing habitats and prioritizing species conservation efforts in light of the threats to biodiversity posed by climate change.
  • Boldness at the nest predicts foraging and diving behaviour of female but
           not male African penguins
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Gwendoline Traisnel, Lorien PichegruOptimal foraging theory suggests that the environment (e.g. distribution of resource patches) will shape an individual's decision to exploit the resource available or explore other locations. Together with the environment and the social context, individual characteristics such as personality have been recently discovered to affect behaviour, offering new insights in the field of behavioural ecology. In many species, personality types differ in breeding success, and foraging behaviour could potentially mediate the influence of personality on reproductive output. A recent study conducted on African penguins, Spheniscus demersus, revealed that bold and shy individuals raised chicks with different growth rates. Here we investigated whether strategies of resource acquisition vary with personality, that is, boldness degree, which may explain the differences in breeding success previously observed in this endangered species. Over 3 years, we deployed GPS loggers, sometimes in conjunction with time–depth recorders, on chick-rearing penguins on Bird Island, Algoa Bay, South Africa, and recorded their boldness degree using a standard human approach protocol. Bolder females, but not males, foraged following a more sinuous path than shyer ones. Similarly, wiggle frequency and the total vertical distance travelled underwater increased with female boldness degree. These results suggest different resource acquisition strategies between personality types in female African penguins. However, none of the foraging or diving behaviour characteristics influenced breeding success. These findings suggest that personality may influence breeding success through other mechanisms (e.g. chick provisioning). Our study indicates that, in this species, some nest behaviours (i.e. proxy of personality) may link with foraging and diving behaviours more than others. Nevertheless, the sex-specific adaptation observed in foraging and diving behaviour may result from a differential investment in the breeding season and reduce intraspecific competition for food in this species.
  • Ephemeral temporal partitioning may facilitate coexistence in competing
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Adia R. Sovie, Daniel U. Greene, Catherine F. Frock, Alex D. Potash, Robert A. McCleeryAnimals change their daily activity patterns in response to season, food availability and the presence of competitors. Competition may be an important driver of a species’ daily activity pattern, as animals manage conflict by avoiding each other temporally. We evaluated how vegetation structure and the presence of competitors changed the daily activity patterns of closely related fox squirrels, Sciurus niger, and grey squirrels, Sciurus carolinensis. We monitored squirrel activity in north and central Florida using passive game cameras at two spatial scales: local and point. To understand how seasonality and competition interact to drive behaviour, we compared squirrel activity during the leaf-off (1 January – 15 March) and leaf-on (16 March – 1 July) seasons. We tested for a relationship between squirrel activity and canopy cover by fitting a von Mises kernel distribution. To test how season and competition affected squirrel behaviour, we compared activity by computing a kernel density overlap function, ranging from 0 (no overlap: the squirrels are never active at the same time) to 1 (complete overlap: the squirrels have identical activity patterns). We found that daily squirrel behaviour was not influenced by canopy cover (P = 0.61). Fox squirrels had a single activity peak occurring around midday. In contrast, grey squirrels had a bimodal activity pattern with peaks shortly after sunrise and before sunset. The intensity of this partitioning existed on a gradient and changed with season and the presence of competitors. Fox and grey squirrel daily patterns overlapped the most when they were allopatric in the leaf-on season (overlap = 0.70, P < 0.001) and the least while sympatric in the leaf-on season (overlap = 0.24, P < 0.08). This ephemeral response to competition highlights that various axes of resource partitioning can promote coexistence between closely related species.
  • Pair bonding in monogamously and polygynously kept African striped mice,
           Rhabdomys pumilio
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Lorène Garnier, Carsten SchradinPair bonding (i.e. individuals showing a preference for a specific opposite-sex individual) has been demonstrated in several socially monogamous species. However, social bonds also occur in nonmonogamous species, but have received less attention. Currently, we do not know whether social bonds in monogamous pairs differ from social bonds in polygynous groups. We studied the socially flexible African striped mouse in the laboratory, conducting 3 h partner preference tests typically used to measure pair bonds in socially monogamous prairie voles, Microtus ochrogaster. In the field, striped mice typically live in polygynous groups, but socially monogamous pairs have also been observed. We compared social bonds between 12 monogamous pairs and 12 polygynous groups (1 male and 2 females). The social situation (monogamous versus polygynous) did not influence social bonds. Female striped mice showed a preference for their partner. While males spent more time in body contact with their partner, they showed a sexual preference for strange females. Polygynous males did not show a preference for one of their two females. While significant preferences for partners were found in striped mice, social preference was less strong than that reported for socially monogamous prairie voles. In summary, our results suggest that opposite-sex social bonds not only occur in monogamous species but also in species that live in polygynous groups, but that these bonds might be weaker in polygynous species.
  • Slow natal dispersal across a homogeneous landscape suggests the use of
           mixed movement behaviours during dispersal in the Darwin's frog
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Andrés Valenzuela-Sánchez, Hugo Cayuela, Benedikt R. Schmidt, Andrew A. Cunningham, Claudio Soto-AzatDispersal is a key process in ecology and evolution. Both theoretical and empirical evidence in actively dispersing organisms support the general notion that the use of nearly straight dispersal paths is a highly efficient way to maximize dispersal success in heterogenous landscapes. In homogeneous landscapes, in contrast, the benefits of a straighter dispersal path could be outweighed by an increase in risk costs, favouring the evolution of tortuous dispersal paths resulting in a relatively slow dispersal. Empirical support for this theoretical prediction, however, has remained elusive. To explore this theoretical prediction, we studied the movement behaviour of the southern Darwin's frog, Rhinoderma darwinii, a fully terrestrial amphibian inhabiting a highly homogeneous environment (i.e. South American temperate forest). Using spatial capture–recapture data collected over a 4-year period in wild populations, in combination with statistical and simulation modelling, we found evidence of a slow natal dispersal lasting one year or more. In contrast, adults exhibited high site fidelity, having a median annual displacement of 3.64 m. A correlated random walk model produced synthetic distributions of juvenile annual displacement that were nearly identical to the empirical data, suggesting that a plausible explanation of juvenile dispersal is the use of routine movements (with high path tortuosity) over short temporal scales (
  • The importance of individual variation in the alarm calls of Gunnison's
           prairie dogs
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): W.J. Loughry, Mariah Oeser, Corey Devin Anderson, John L. HooglandGunnison's prairie dogs, Cynomys gunnisoni, frequently utter a repetitive series (a ‘bout’) of alarm calls (called ‘barks’) in response to a variety of terrestrial predators. Previous work has suggested that the structure of barks is finely tuned in ways that provide highly specific information about the identity of the predator. We re-evaluated these findings by presenting individually marked adult females with taxidermy mounts of three terrestrial predators of prairie dogs: a coyote, Canis latrans, a bobcat, Lynx rufus, and an American badger, Taxidea taxus, as well as one control stimulus (a cardboard box). Significant variation among individuals was evident from analyses of 10 measures associated with the general pattern of barking (e.g. latency to begin barking, bout duration, average number of barks/bout, overall bark rate, interbout interval) and also from eight structural features of individual barks (e.g. frequency and duration measures of the fundamental frequency band). When controlling for the identity of alarm callers, we found little evidence for differences in these same variables in response to different stimuli. We conclude that, like several other species of ground-dwelling squirrels, Gunnison's prairie dogs have an individually distinctive alarm call that does not differ for different predators. Our results conflict with earlier claims of extreme specificity in the alarm calls of Gunnison's prairie dogs.
  • Female and male plumage colour signals aggression in a dichromatic
           tropical songbird
    • Abstract: Publication date: Available online 4 March 2019Source: Animal BehaviourAuthor(s): Ana V. Leitão, Michelle L. Hall, Kaspar Delhey, Raoul A. MulderAnimal signals of competitive ability allow contests for limited resources to be settled without costly physical fights. Traits indicating competitive ability are diverse and span visual, acoustic or chemical modalities. Although animal signalling has been intensively studied, research has focused mainly on male traits. Little is known about the extent and functionality of competitive signals in females and whether there are sex differences in signal function. We studied whether plumage colour signals competitive ability in female and male lovely fairy-wrens, Malurus amabilis. In this species, both sexes sport elaborate but sexually dichromatic ornamental plumage. Using a mirror image stimulation test, we first assessed the relationship between male and female colour and agonistic behaviour, controlling for other physical, social and ecological variables. We then tested whether colourful plumage influenced aggressive response in both sexes by experimentally manipulating plumage colour and measuring individual responses to their mirror image. Females and males were more aggressive towards naturally less colourful reflections of the cheek patch in the mirror. However, when we manipulated plumage colour, both females and males responded more aggressively to experimentally increased cheek colour reflection in the mirror. Our findings suggest that plumage colour signals competitive ability in an aggressive context in both sexes and raises the possibility that signal reliability may be maintained by social interactions where individuals police and punish dishonest signals.
  • Communication in social insects and how it is shaped by individual
    • Abstract: Publication date: Available online 4 March 2019Source: Animal BehaviourAuthor(s): Christoph Grüter, Tomer J. CzaczkesCommunication is the foundation of all social systems, and learning is perhaps the most important cognitive tool. But how do these two critical faculties interact' With social insects being some of the best learners of the invertebrate world, and indisputably the most communicative, we examine the role of learning and experience in social insect communication. Learning plays a major role for both senders and receivers. A sender's experience can modulate what information is available for communication, whether communication is effective and whether individuals are motivated to communicate. Signalling about a resource is often modulated relative to the value of that resource and relative to the value of sharing information about it. The receiver's experience and knowledge can affect which parts of a signal's information content it attends to, how this information is acted on and, indeed, whether it attends to communication at all. Ultimately, while innate responses form the basis of social insect communication, learning is often a critical modulator of communication processes.
  • To buzz or burst-pulse' The functional role of Heaviside's dolphin,
           Cephalorhynchus heavisidii, rapidly pulsed signals
    • Abstract: Publication date: Available online 4 March 2019Source: Animal BehaviourAuthor(s): Morgan J. Martin, Simon H. Elwen, Reshma Kassanjee, Tess GridleyFour groups of toothed whales have independently evolved to produce narrowband high-frequency (NBHF) echolocation signals (i.e. clicks) with a strikingly similar waveform and centroid frequency around 125 kHz. These signals are thought to help NBHF species avoid predation by echolocating and communicating at frequencies inaudible to predators, a form of acoustic crypsis. Heaviside's dolphins produce NBHF echolocation clicks in trains and often in rapid succession in the form of buzzes. In addition, a second click type with a lower frequency and broader bandwidth was recently described, typically emitted in rapid succession in the form of burst-pulses. We investigated the relationship between buzz and burst-pulse signals and both surface behaviour (foraging, ‘interacting with the kayak’ and socializing) and group size, using a multivariable regression on the signal occurrence and signal count data. Signal occurrence and counts were not related to group size in the regression analysis. Burst-pulses were strongly linked to socializing behaviour, occurring more often and more frequently during socializing and much less during foraging. Buzz vocalizations were not strongly linked to a specific behaviour although there was some evidence of an increase in production during foraging and socializing. In addition, individual level production rates of buzzes during foraging and socializing, and burst-pulses during socializing decreased with increasing group size. Temporally patterned burst-pulse signals were also identified, often occurring within a series of burst-pulses and were directly linked to specific events such as aerial leaping, backflipping, tail slapping and potential mating. Our findings suggest Heaviside's dolphins have a more complex communication system based on pulsed vocalizations than previously understood, perhaps driven by the need to facilitate the social interactions of this species.
  • Challenges in assessing the roles of nepotism and reciprocity in
           cooperation networks
    • Abstract: Publication date: Available online 4 March 2019Source: Animal BehaviourAuthor(s): Gerald G. Carter, Gabriele Schino, Damien FarineNepotism and reciprocity are not mutually exclusive explanations for cooperation, because helping decisions can depend on both kinship cues and past reciprocal help. The importance of these two factors can therefore be difficult to disentangle using observational data. We developed a resampling procedure for inferring the statistical power to detect observational evidence of nepotism and reciprocity. We first applied this procedure to simulated data sets resulting from perfect reciprocity, where the probability and duration of helping events from individual A to B equalled that from B to A. We then assessed how the probability of detecting correlational evidence of reciprocity was influenced by (1) an increasing number of helping observations and (2) an increasing degree of simultaneous nepotism. Last, we applied the same analyses to empirical data on food sharing in common vampire bats, Desmodus rotundus, and allogrooming in mandrills, Mandrillus sphinx, and Japanese macaques, Macaca fuscata. We show that at smaller sample sizes, the effect of kinship was easier to detect and overestimated relative to the effect of reciprocal help. This bias in power was true in both empirical and simulated data, including when simulating perfect reciprocity and imperfect nepotism. We explain the causes and consequences of this difference in power for detecting the roles of kinship versus reciprocal help. When comparing the relative evidence for kin-biased help and reciprocal help, we suggest that researchers measure the relative reliability of both kinship bias and symmetry in the model by plotting the coefficients and their detection probability as a function of sampling effort. We provide R scripts to allow others to do this power analysis with their own data sets.
  • Fussing over food: factors affecting the vocalizations American crows
           utter around food
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): LomaJohn T. Pendergraft, John M. MarzluffAmerican crows, Corvus brachyrhynchos, often loudly vocalize when gathered around a food source. Because doing so would attract unwanted attention from predators and competitors, animals that have congregated around food are only expected to vocalize if the benefits (e.g. recruiting or announcing themselves to allies, deterring competitors, warning of danger, begging for a meal, appeasing a dominant) outweigh these costs. Here we demonstrate that wild crows change the quality of their calls depending on the amount of food present. The crows near a large food windfall gave shorter calls compared to their vocalizations in food's absence, and playback of these short calls only prompted a mild aggressive response from listening crows. In contrast, the calls given before the appearance of food were longer, and their playback elicited behaviours from the listening crows associated with aggression and territory defence. These findings suggest that crows avoid giving territorial calls near an exploitable food resource and vocalize for other reasons. Taken together, this study provides insights on how the caller's current context can shift the costs and benefits of vocalizing.
  • Spatial cognitive performance is linked to thigmotaxis in field crickets
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Maria D. Doria, Julie Morand-Ferron, Susan M. BertramInformation is acquired through cognitive processes and is essential for enhanced fitness in many animal species. However, little is known about intraspecific variation in cognitive performance, or about phenotypic correlates of this variation, especially in invertebrates. Here we quantified individual variation in spatial learning, exploration, boldness and thigmotaxis (wall hugging) in female Texas field crickets, Gryllus texensis. We trained females to locate a food reward (dish baited with food) located in one arm of a radial arm maze, followed by a cognitive test (probe) trial where no food reward was present. Our results suggest that female crickets could learn and remember the food location, as latency to find the reward dish, distance travelled, number of complete arm entries and number of errors made prior to finding the reward dish all declined with training trials and during the cognitive test trial. Furthermore, females performed equally well on all cognitive measures during the cognitive test trial and the last training trial, suggesting they did not rely on olfactory cues to locate the correct dish during the cognitive test trial. Individual variation in cognitive measures during the test trial were significantly negatively correlated with thigmotaxis during the trial, but were not correlated with body size, latency to leave the novel maze's acclimation zone or the percentage of the novel maze explored during initial exploration of the maze. Our findings suggest individuals with reduced thigmotaxis either (1) are better at learning a spatial task and/or (2) rely more heavily on vision than olfaction. These results contribute to the growing literature examining links between personality and cognition, and highlight the importance of quantifying correlated aspects of the phenotype when assessing individual variation in cognitive performance in a cognitive test.
  • The effect of against-background contrast on female preferences for a
           polymorphic colour sexual signal
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Jessica C.B. Lynn, Gemma L. ColeIt is thought that the more conspicuous a sexual signal, the more attractive it is; however, the role of the background in modifying the conspicuousness of a signal is not well understood, particularly for complex, highly polymorphic colour signals. The background against which a sexual signal is viewed is important because it can influence how an individual's colour pattern is perceived and therefore whether it is attractive to potential mates. Here, we tested the hypothesis that against-background contrast in hue (colour), luminance (brightness) and chroma (saturation) will influence the attractiveness of a male and we predicted that the more a male's colour pattern contrasts with the background, the more attractive he should be. We used mate choice tests to determine the effect of visual background contrast on female preferences in a freshwater fish, Poecilia reticulata, with a highly polymorphic male colour sexual signal. We placed three full-sibling males, with very similar colour patterns, in front of three different backgrounds which varied in their perceived contrast to females. We then gave females the opportunity to spend time with the male that they found most attractive. Contrary to our predictions, we found that females did not choose the males that contrasted the most with their background. More specifically, we found nonlinear relationships between female preference and the coefficient of variation (CV) of chromatic contrast, which represents the variation in colour patch saturation across the pattern; the higher the CV chroma, the more the colour patches within a pattern vary in saturation. Our results indicate possible trade-offs between natural and sexual selection and/or signatures of sensory bias. Against-background contrast may play a role in maintaining colour pattern polymorphism.
  • Developmental stage-dependent response and preference for host plant
           quality in an insect herbivore
    • Abstract: Publication date: April 2019Source: Animal Behaviour, Volume 150Author(s): Ana L. Salgado, Marjo SaastamoinenLarval-derived nutritional reserves are essential in shaping insects' adult fitness. Early larval instars of many Lepidopteran species are often sessile, and the conditions experienced by these larvae are often highly dependent on the mother's oviposition choice. Later larval stages are more mobile and therefore can choose their food whenever alternatives are available. We tested how feeding on a drought-exposed host plant impacts life history in an insect herbivore, and whether the observed responses depended on developmental stage. We used drought to alter host plant quality of the ribwort plantain, Plantago lanceolata, and assessed whether host plant preference of postdiapause larvae and adult females increased their own or their offspring's performance, respectively, in the Glanville fritillary butterfly, Melitaea cinxia. Larval response to drought-exposed host plants varied with developmental stage: early larval stages (prediapause) had decreased survival and body mass on drought-exposed plants, while later larval stages (postdiapause) developed faster, weighed more and had a higher growth rate on the drought-exposed plants. Postdiapause larvae also showed a preference for drought-exposed host plants, i.e. those that increased their performance, but only when fed on well-watered host plants. Adult females, on the other hand, showed an oviposition preference for well-watered plants, hence matching the performance of their prediapause but not their postdiapause offspring. Our results highlight how variation in environmental conditions generates stage-specific responses in insects. Individuals fine-tune their own or their offspring's diet by behavioural adjustments when variation in host plant quality is available.
  • Volition and learning in primate vocal behaviour
    • Abstract: Publication date: Available online 1 March 2019Source: Animal BehaviourAuthor(s): Asif A. Ghazanfar, Diana A. Liao, Daniel Y. TakahashiInvestigating nonhuman primate vocal communication is often with the intention of elucidating their similarities with human speech and thus reconstructing the evolutionary history of this important behaviour. However, putative parallels between primate and human vocal behaviours have, in some respects, remained elusive. Here, we review two lines of research in marmoset monkeys on volitional vocal control and vocal learning during development that could bridge our understanding of the relationship between primate vocalizations and human speech. Regarding volitional control, we review how changes in vocal output are not solely due to changes in arousal levels and their effect on the vocal apparatus; extrinsic factors like the vocalizations from other conspecifics also have an important influence. With regard to vocal learning, we describe not only how infant marmoset vocalizations undergo dramatic acoustic changes during development that are not wholly explained by physical growth, but also how, as in humans, contingent vocal responses from parents influence the rate of vocal development. We argue that the similarities in the vocal systems of marmoset monkeys and humans may be due to their shared cooperative breeding strategy and prosociality.
  • Birdsong learning, avian cognition and the evolution of language
    • Abstract: Publication date: Available online 23 February 2019Source: Animal BehaviourAuthor(s): William A. Searcy, Stephen NowickiThe most language-like aspect of the song of songbirds is its development: as with human speech, birdsong develops through vocal production learning, in which individuals modify the structure of their vocalizations in response to experience with the vocalizations of others. As is true of speech development, birdsong learning qualifies as a cognitive ability, as it involves the acquisition, storage and processing of information obtained from the environment. Accordingly, if cognitive abilities are in general positively associated in songbirds, as has been argued for humans and other mammals, then song learning ability should be positively associated with other cognitive abilities, and learned attributes of song should serve as an indicator of domain-general cognition. A review of studies in which songbirds have been subjected to batteries of cognitive tasks finds, however, that different cognitive measures are not consistently positively associated. Moreover, learned attributes of birdsong do not show consistent positive associations with other cognitive measures. These results argue that, rather than being a component of domain-general cognition, song learning is an autonomous cognitive module. Birdsong learning shows other characteristics of modularization in that it is domain specific, is based on a localized and highly structured neural system and exhibits a level of innate specification. Whether language learning in humans is similarly modularized has been much debated. Despite a possible difference in modularization, much can be learned about the evolution of human language learning from studying birdsong learning. Aspects of birdsong learning that are especially relevant include vocal interaction learning, pragmatics and the initial selective benefits and neural underpinnings of vocal production learning.
  • Syntactic rules in avian vocal sequences as a window into the evolution
           of compositionality
    • Abstract: Publication date: Available online 14 February 2019Source: Animal BehaviourAuthor(s): Toshitaka N. Suzuki, Michael Griesser, David WheatcroftUnderstanding the origins and evolution of language remains a deep challenge, because its complexity and expressive power are unparalleled in the animal world. One of the key features of language is that the meaning of an expression is determined both by the meanings of its constituent parts and the syntactic rules used to combine them; known as the principle of compositionality. Although compositionality has been considered unique to language, recent field studies suggest that compositionality may have also evolved in vocal combinations in nonhuman animals. Here, we discuss how compositionality can be explored in animal communication systems and review recent evidence that birds use an ordering rule to generate compositional expressions composed of meaningful calls. Also, we suggest that birdsongs, particularly when incorporating calls, may represent unrecognized examples of compositionality in animal communication. Finally, we outline future research directions to uncover the development, neural mechanisms and evolution of compositionality.
  • Rules, rhythm and grouping: auditory pattern perception by birds
    • Abstract: Publication date: Available online 18 December 2018Source: Animal BehaviourAuthor(s): Carel ten Cate, Michelle SpieringsBoth language and music are universal and characteristic for humans. The evolution of the cognitive abilities underlying language and music are widely debated. A core question is whether these abilities find their origins in a modification or extension of general cognitive abilities for processing auditory input also present in other species. If so, comparative studies of nonhuman animals should reveal similarities in processing abilities. In this paper, we review some examples of such studies. We focus on whether birds (in particular zebra finches, Taeniopygia guttata, and budgerigars, Melopsittacus undulates) can detect structural patterns based on relational rather than on physical similarities among auditory stimuli – an essential ability for processing and producing language and music. We briefly discuss why birds are suitable model species. Next, we discuss three domains of pattern detection: the ability to (1) detect ‘grammatical rules’ underlying sound strings; (2) perceive regular rhythms and (3) spontaneously group separate sounds into a larger pattern. In all of these there is evidence that birds show some ability to detect relational patterns. However, there is also variation between species: while budgerigars show relational rule learning, zebra finches attend to local physical similarities between sound strings used for training and testing. For rhythm detection, zebra finches and budgerigars show no clear differences. However, a broader comparison indicates that here too differences are present in the extent to which different bird species attend to relational patterns or to local features. Finally, spontaneous grouping of sounds was shown in zebra finches. The clear variation among bird species in their perceptual and cognitive abilities, in combination with their accessibility for experimental studies, provides opportunities to study the variation in auditory processing mechanisms and how these evolved. This may also provide hypotheses for the evolution of these abilities in humans.
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