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Whishaw, I. Q., Sacrey, L. - A. R., & Gorny, B. (2009). Hind limb stepping over obstacles in the horse guided by place-object memory. Behav. Brain. Res., 198(2), 372–379.
Abstract: An animal that has stepped over an obstacle with its forelimbs uses a memory of the obstacle to guide the hind limbs so that they also clear the obstacle, even in situations in which long pauses are introduced between forelimb and hind limb stepping. To further clarify the features of hind limb obstacle clearance memory, the present study examined hind limb obstacle clearance in the horse. A rider guided horses over obstacles and paused the horse over obstacles in tests that examined the relationship between forelimb and hind limb stepping, with the following results. First, the horses displayed memory for an obstacle as measured by hind limb lifting over the obstacle for durations lasting as long as 15Â min. The response was not dependent upon ongoing visualization of the obstacle, as limb lifting was unaffected by visual occlusion with blinders, a blindfold, or by removing the obstacle during the pause. Second, previous experience of stepping over an obstacle led to pause-related hind limb lifting at the object's previous location even on trials for which there was no obstacle and so no preceding forelimb lifting. Third, whereas a horse would lift its hind limbs to clear two successively presented obstacles, replacing an obstacle before the horse after the forelimbs had cleared the obstacle prevented subsequent hind limb lifting at the obstacle's previous location. Taken together the results show that hind limb obstacle clearance is guided by a place-object memory. The results are discussed in relation to the differential sensory and memonic control of forelimb and hind limb stepping with the suggestion that place-object memory can guide hind stepping as well as overshadow working memory from front leg stepping.
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Proops, L., McComb, K., & Reby, D. (2009). Cross-modal individual recognition in domestic horses (Equus caballus). Proc. Natl. Acad. Sci. U.S.A., 106(3), 947–951.
Abstract: Individual recognition is considered a complex process and, although it is believed to be widespread across animal taxa, the cognitive mechanisms underlying this ability are poorly understood. An essential feature of individual recognition in humans is that it is cross-modal, allowing the matching of current sensory cues to identity with stored information about that specific individual from other modalities. Here, we use a cross-modal expectancy violation paradigm to provide a clear and systematic demonstration of cross-modal individual recognition in a nonhuman animal: the domestic horse. Subjects watched a herd member being led past them before the individual went of view, and a call from that or a different associate was played from a loudspeaker positioned close to the point of disappearance. When horses were shown one associate and then the call of a different associate was played, they responded more quickly and looked significantly longer in the direction of the call than when the call matched the herd member just seen, an indication that the incongruent combination violated their expectations. Thus, horses appear to possess a cross-modal representation of known individuals containing unique auditory and visual/olfactory information. Our paradigm could provide a powerful way to study individual recognition across a wide range of species.
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Cameron, E. Z., Setsaas, T. H., & Linklater, W. L. (2009). Social bonds between unrelated females increase reproductive success in feral horses. Proc. Natl. Acad. Sci. U.S.A., 106(33), 13850–13853.
Abstract: In many mammals, females form close social bonds with members of their group, usually between kin. Studies of social bonds and their fitness benefits have not been investigated outside primates, and are confounded by the relatedness between individuals in primate groups. Bonds may arise from kin selection and inclusive fitness rather than through direct benefits of association. However, female equids live in long-term social groups with unrelated members. We present 4 years of behavioral data, which demonstrate that social integration between unrelated females increases both foal birth rates and survival, independent of maternal habitat quality, social group type, dominance status, and age. Also, we show that such social integration reduces harassment by males. Consequently, social integration has strong direct fitness consequences between nonrelatives, suggesting that social bonds can evolve based on these direct benefits alone. Our results support recent studies highlighting the importance of direct benefits in maintaining cooperative behavior, while controlling for the confounding influence of kinship.
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Finarelli, J. A., & Flynn, J. J. (2009). Brain-size evolution and sociality in Carnivora. Proc. Natl. Acad. Sci. U.S.A., 106(23), 9345–9349.
Abstract: Increased encephalization, or larger brain volume relative to body mass, is a repeated theme in vertebrate evolution. Here we present an extensive sampling of relative brain sizes in fossil and extant taxa in the mammalian order Carnivora (cats, dogs, bears, weasels, and their relatives). By using Akaike Information Criterion model selection and endocranial volume and body mass data for 289 species (including 125 fossil taxa), we document clade-specific evolutionary transformations in encephalization allometries. These evolutionary transformations include multiple independent encephalization increases and decreases in addition to a remarkably static basal Carnivora allometry that characterizes much of the suborder Feliformia and some taxa in the suborder Caniformia across much of their evolutionary history, emphasizing that complex processes shaped the modern distribution of encephalization across Carnivora. This analysis also permits critical evaluation of the social brain hypothesis (SBH), which predicts a close association between sociality and increased encephalization. Previous analyses based on living species alone appeared to support the SBH with respect to Carnivora, but those results are entirely dependent on data from modern Canidae (dogs). Incorporation of fossil data further reveals that no association exists between sociality and encephalization across Carnivora and that support for sociality as a causal agent of encephalization increase disappears for this clade.
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Liker, A., & Bókony, V. (2009). Larger groups are more successful in innovative problem solving in house sparrows. Proc Natl Acad Sci USA, 106(19), 7893–7898.
Abstract: Group living offers well-known benefits to animals, such as better predator avoidance and increased foraging success. An important additional, but so far neglected, advantage is that groups may cope more effectively with unfamiliar situations through faster innovations of new solutions by some group members. We tested this hypothesis experimentally by presenting a new foraging task of opening a familiar feeder in an unfamiliar way to house sparrows in small and large groups (2 versus 6 birds). Group size had strong effects on problem solving: sparrows performed 4 times more and 11 times faster openings in large than in small groups, and all members of large groups profited by getting food sooner (7 times on average). Independently from group size, urban groups were more successful than rural groups. The disproportionately higher success in large groups was not a mere consequence of higher number of attempts, but was also related to a higher effectiveness of problem solving (3 times higher proportion of successful birds). The analyses of the birds' behavior suggest that the latter was not explained by either reduced investment in antipredator vigilance or reduced neophobia in large groups. Instead, larger groups may contain more diverse individuals with different skills and experiences, which may increase the chance of solving the task by some group members. Increased success in problem solving may promote group living in animals and may help them to adapt quickly to new situations in rapidly-changing environments.
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Gácsi, M., Gyoöri, B., Virányi, Z., Kubinyi, E., Range, F., Belényi, B., et al. (2009). Explaining Dog Wolf Differences in Utilizing Human Pointing Gestures: Selection for Synergistic Shifts in the Development of Some Social Skills. PLoS ONE, 4(8), e6584.
Abstract: <sec> <title>Background</title> <p>The comparison of human related communication skills of socialized canids may help to understand the evolution and the epigenesis of gesture comprehension in humans. To reconcile previously contradicting views on the origin of dogs' outstanding performance in utilizing human gestures, we suggest that dog-wolf differences should be studied in a more complex way.</p> </sec><sec> <title>Methodology/Principal Findings</title> <p>We present data both on the performance and the behaviour of dogs and wolves of different ages in a two-way object choice test. Characteristic behavioural differences showed that for wolves it took longer to establish eye contact with the pointing experimenter, they struggled more with the handler, and pups also bit her more before focusing on the human's signal. The performance of similarly hand-reared 8-week-old dogs and wolves did not differ in utilizing the simpler proximal momentary pointing. However, when tested with the distal momentary pointing, 4-month-old pet dogs outperformed the same aged hand reared wolves. Thus early and intensive socialisation does not diminish differences between young dogs and wolves in behaviour and performance. Socialised adult wolves performed similarly well as dogs in this task without pretraining. The success of adult wolves was accompanied with increased willingness to cooperate.</p> </sec><sec> <title>Conclusion/Significance</title> <p>Thus, we provide evidence for the first time that socialised adult wolves are as successful in relying on distal momentary pointing as adult pet dogs. However, the delayed emergence of utilising human distal momentary pointing in wolves shows that these wild canines react to a lesser degree to intensive socialisation in contrast to dogs, which are able to control agonistic behaviours and inhibition of actions in a food related task early in development. We suggest a “synergistic” hypothesis, claiming that positive feedback processes (both evolutionary and epigenetic) have increased the readiness of dogs to attend to humans, providing the basis for dog-human communication.</p> </sec>
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Gácsi, M., McGreevy, P., Kara, E., & Miklósi, Á. (2009). Effects of selection for cooperation and attention in dogs. Behav Brain Funct, 5, 31.
Abstract: ABSTRACT: BACKGROUND: It has been suggested that the functional similarities in the socio-cognitive behaviour of dogs and humans emerged as a consequence of comparable environmental selection pressures. Here we use a novel approach to account for the facilitating effect of domestication in dogs and reveal that selection for two factors under genetic influence (visual cooperation and focused attention) may have led independently to increased comprehension of human communicational cues. METHOD: In Study 1, we observed the performance of three groups of dogs in utilizing the human pointing gesture in a two-way object choice test. We compared breeds selected to work while visually separated from human partners (N = 30, 21 breeds, clustered as independent worker group), with those selected to work in close cooperation and continuous visual contact with human partners (N = 30, 22 breeds, clustered as cooperative worker group), and with a group of mongrels (N = 30).Secondly, it has been reported that, in dogs, selective breeding to produce an abnormal shortening of the skull is associated with a more pronounced area centralis (location of greatest visual acuity). In Study 2, breeds with high cephalic index and more frontally placed eyes (brachycephalic breeds, N = 25, 14 breeds) were compared with breeds with low cephalic index and laterally placed eyes (dolichocephalic breeds, N = 25, 14 breeds). RESULTS: In Study 1, cooperative workers were significantly more successful in utilizing the human pointing gesture than both the independent workers and the mongrels.In study 2, we found that brachycephalic dogs performed significantly better than dolichocephalic breeds. DISCUSSION: After controlling for environmental factors, we have provided evidence that at least two independent phenotypic traits with certain genetic variability affect the ability of dogs to rely on human visual cues. This finding should caution researchers against making simple generalizations about the effects of domestication and on dog-wolf differences in the utilization of human visual signals.
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Langergraber, K., Mitani, J., & Vigilant, L. (2009). Kinship and social bonds in female chimpanzees (Pan troglodytes). Am. J. Primatol., 71(10), 840–851.
Abstract: A large body of theoretical and empirical research suggests that kinship influences the development and maintenance of social bonds among group-living female mammals, and that human females may be unusual in the extent to which individuals form differentiated social relationships with nonrelatives. Here we combine behavioral observations of party association, spatial proximity, grooming, and space use with extensive molecular genetic analyses to determine whether female chimpanzees form strong social bonds with unrelated individuals of the same sex. We compare our results with those obtained from male chimpanzees who live in the same community and have been shown to form strong social bonds with each other. We demonstrate that party association is as good a predictor of spatial proximity and grooming in females as it is in males, that the highest party association indices are consistently found between female dyads, that the sexes do not differ in the long-term stability of their party association patterns, and that these results cannot be explained as a by-product of the tendency of females to selectively range in particular areas of the territory. We also show that close kin (i.e. mother-daughter and sibling dyads) are very rare, indicating that the vast majority of female dyads that form strong social bonds are not closely related. Additional analyses reveal that “subgroups” of females, consisting of individuals who frequently associate with one another in similar areas of the territory, do not consist of relatives. This suggests that a passive form of kin-biased dispersal, involving the differential migration of females from neighboring communities into subgroups, was also unlikely to be occurring. These results show that, as in males, kinship plays a limited role in structuring the intrasexual social relationships of female chimpanzees.
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Gosden, T. P., & Svensson, E. I. (2009). Density-Dependent Male Mating Harassment, Female Resistance, and Male Mimicry. Am Nat, 173(6), 709–721.
Abstract: Abstract:
Genetic variation in female resistance and tolerance to male mating harassment can affect the outcome of sexually antagonistic mating interactions. We investigated female mating rates and male mating harassment in natural populations of a damselfly (Ischnura elegans). This damselfly species has a heritable sex‐limited polymorphism in females, where one of the morphs is a male mimic (androchrome females). The three female morphs differ in mating rates, and these differences are stable across populations and years. However, the degree of premating resistance toward male mating attempts varied across generations and populations. Male mating harassment of the female morphs changed in a density‐dependent fashion, suggesting that male mate preferences are plastic and vary with the different morph densities. We quantified morph differences in male mating harassment and female fecundity, using path analysis and structural equation modeling. We found variation between the morphs in the fitness consequences of mating, with the fecundity of one of the nonmimetic morphs declining with increasing male mating harassment. However, androchrome females had lower overall fecundity, presumably reflecting a cost of male mimicry. Density‐dependent male mating harassment on the morphs and fecundity costs of male mimicry are thus likely to contribute to the maintenance of this female polymorphism.
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Basile, M., Boivin, S., Boutin, A., Blois-Heulin, C., Hausberger, M., & Lemasson, A. (2009). Socially dependent auditory laterality in domestic horses (Equus caballus). Anim. Cogn., 12, 611–619.
Abstract: Laterality is now known to be an ubiquitous phenomenon among the vertebrates. Particularly, laterality of auditory processing has been demonstrated in a variety of species, especially songbirds and primates. Such a hemispheric specialization has been shown to depend on factors such as sound structure, species specificity and types of stimuli. Much less is known on the possible influence of social familiarity although a few studies suggest such an influence. Here we tested the influence of the degree of familiarity on the laterality of the auditory response in the domestic horse. This species is known for its social system and shows visible reactions to sounds, with one or two ears moving towards a sound source. By comparing such responses to the playback of different conspecific whinnies (group member, neighbor and stranger), we could demonstrate a clear left hemisphere (LH) preference for familiar neighbor calls while no preference was found for group member and stranger calls. Yet, we found an opposite pattern of ear side preference for neighbor versus stranger calls. These results are, to our knowledge, the first to demonstrate auditory laterality in an ungulate species. They open further lines of thought on the influence of the social “value” of calls and the listener's arousal on auditory processing and laterality.
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