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Larose, C., Richard-Yris, M. - A., Hausberger, M., & Rogers, L. J. (2006). Laterality of horses associated with emotionality in novel situations. Laterality, 11(4), 355–367.
Abstract: We have established that lateral biases are characteristic of visual behaviour in 65 horses. Two breeds, Trotters and French Saddlebreds aged 2 to 3, were tested on a novel object test. The main finding was a significant correlation between emotionality index and the eye preferred to view the novel stimulus: the higher the emotionality, the more likely that the horse looked with its left eye. The less emotive French Saddlebreds, however, tended to glance at the object using the right eye, a tendency that was not found in the Trotters, although the emotive index was the same for both breeds. The youngest French Saddlebreds did not show this trend. These results are discussed in relation to the different training practices for the breeds and broader findings on lateralisation in different species.
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Henderson, A. J. Z. (2007). Don't fence me in: managing psychological well being for elite performance horses. J. Appl. Anim. Welf. Sci., 10(4), 309–329.
Abstract: This article posits that stereotypical behavior patterns and the overall psychological well being of today's performance horse could be substantially enhanced with care that acknowledges the relationship between domesticated horses and their forerunners. Feral horses typically roam in stable, social groups over large grazing territories, spending 16-20 hr per day foraging on mid- to poor-quality roughage. In contrast, today's elite show horses live in relatively small stalls, eat a limited-but rich-diet at specific feedings, and typically live in social isolation. Although the horse has been domesticated for more than 6000 years, there has been no selection for an equid who no longer requires an outlet for these natural behaviors. Using equine stereotypies as a welfare indicator, this researcher proposes that the psychological well being of today's performance horse is compromised. Furthermore, the article illustrates how minimal management changes can enhance horses' well being while still remaining compatible with the requirements of the sport-horse industry. The article discusses conclusions in terms of Fraser, Weary, Pajor, and Milligan's “integrative welfare model” (1997).
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Johnstone, R. A. (2001). Eavesdropping and animal conflict. Proc. Natl. Acad. Sci. U.S.A., 98(16), 9177–9180.
Abstract: Fights between pairs of animals frequently take place within a wider social context. The displays exchanged during conflict, and the outcome of an encounter, are often detectable by individuals who are not immediately involved. In at least some species, such bystanders are known to eavesdrop on contests between others, and to modify their behavior toward the contestants in response to the observed interaction. Here, I extend Maynard Smith's well known model of animal aggression, the Hawk-Dove game, to incorporate the possibility of eavesdroppers. I show that some eavesdropping is favored whenever the cost of losing an escalated fight exceeds the value of the contested resource, and that its equilibrium frequency is greatest when costs are relatively high. Eavesdropping reduces the risk of escalated conflict relative to that expected by chance, given the level of aggression in the population. However, it also promotes increased aggression, because it enhances the value of victory. The net result is that escalated conflicts are predicted to occur more frequently when eavesdropping is possible.
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Lee, R. D. (2003). Rethinking the evolutionary theory of aging: transfers, not births, shape senescence in social species. Proc Natl Acad Sci U S A, 100(16), 9637–9642.
Abstract: The classic evolutionary theory of aging explains why mortality rises with age: as individuals grow older, less lifetime fertility remains, so continued survival contributes less to reproductive fitness. However, successful reproduction often involves intergenerational transfers as well as fertility. In the formal theory offered here, age-specific selective pressure on mortality depends on a weighted average of remaining fertility (the classic effect) and remaining intergenerational transfers to be made to others. For species at the optimal quantity-investment tradeoff for offspring, only the transfer effect shapes mortality, explaining postreproductive survival and why juvenile mortality declines with age. It also explains the evolution of lower fertility, longer life, and increased investments in offspring.
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Jansen, T., Forster, P., Levine, M. A., Oelke, H., Hurles, M., Renfrew, C., et al. (2002). Mitochondrial DNA and the origins of the domestic horse. Proc. Natl. Acad. Sci. U.S.A., 99(16), 10905–10910.
Abstract: The place and date of the domestication of the horse has long been a matter for debate among archaeologists. To determine whether horses were domesticated from one or several ancestral horse populations, we sequenced the mitochondrial D-loop for 318 horses from 25 oriental and European breeds, including American mustangs. Adding these sequences to previously published data, the total comes to 652, the largest currently available database. From these sequences, a phylogenetic network was constructed that showed that most of the 93 different mitochondrial (mt)DNA types grouped into 17 distinct phylogenetic clusters. Several of the clusters correspond to breeds and/or geographic areas, notably cluster A2, which is specific to Przewalski's horses, cluster C1, which is distinctive for northern European ponies, and cluster D1, which is well represented in Iberian and northwest African breeds. A consideration of the horse mtDNA mutation rate together with the archaeological timeframe for domestication requires at least 77 successfully breeding mares recruited from the wild. The extensive genetic diversity of these 77 ancestral mares leads us to conclude that several distinct horse populations were involved in the domestication of the horse.
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Mettke-Hofmann, C., & Gwinner, E. (2003). Long-term memory for a life on the move. Proc. Natl. Acad. Sci. U.S.A., 100(10), 5863–5866.
Abstract: Evidence is accumulating that cognitive abilities are shaped by the specific ecological conditions to which animals are exposed. Long-distance migratory birds may provide a striking example of this. Field observations have shown that, at least in some species, a substantial proportion of individuals return to the same breeding, wintering, and stopover sites in successive years. This observation suggests that migrants have evolved special cognitive abilities that enable them to accomplish these feats. Here we show that memory of a particular feeding site persisted for at least 12 months in a long-distance migrant, whereas a closely related nonmigrant could remember such a site for only 2 weeks. Thus, it seems that the migratory lifestyle has influenced the learning and memorizing capacities of migratory birds. These results build a bridge between field observations suggesting special memorization feats of migratory birds and previous neuroanatomical results from the same two species indicating an increase in relative hippocampal size from the first to the second year of life in the migrant but not in the nonmigrant.
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Chase, I. D., Tovey, C., Spangler-Martin, D., & Manfredonia, M. (2002). Individual differences versus social dynamics in the formation of animal dominance hierarchies. Proc. Natl. Acad. Sci. U.S.A., 99(8), 5744–5749.
Abstract: Linear hierarchies, the classical pecking-order structures, are formed readily in both nature and the laboratory in a great range of species including humans. However, the probability of getting linear structures by chance alone is quite low. In this paper we investigate the two hypotheses that are proposed most often to explain linear hierarchies: they are predetermined by differences in the attributes of animals, or they are produced by the dynamics of social interaction, i.e., they are self-organizing. We evaluate these hypotheses using cichlid fish as model animals, and although differences in attributes play a significant part, we find that social interaction is necessary for high proportions of groups with linear hierarchies. Our results suggest that dominance hierarchy formation is a much richer and more complex phenomenon than previously thought, and we explore the implications of these results for evolutionary biology, the social sciences, and the use of animal models in understanding human social organization.
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Reader, S. M., & Laland, K. N. (2002). Social intelligence, innovation, and enhanced brain size in primates. Proc. Natl. Acad. Sci. U.S.A., 99(7), 4436–4441.
Abstract: Despite considerable current interest in the evolution of intelligence, the intuitively appealing notion that brain volume and “intelligence” are linked remains untested. Here, we use ecologically relevant measures of cognitive ability, the reported incidence of behavioral innovation, social learning, and tool use, to show that brain size and cognitive capacity are indeed correlated. A comparative analysis of 533 instances of innovation, 445 observations of social learning, and 607 episodes of tool use established that social learning, innovation, and tool use frequencies are positively correlated with species' relative and absolute “executive” brain volumes, after controlling for phylogeny and research effort. Moreover, innovation and social learning frequencies covary across species, in conflict with the view that there is an evolutionary tradeoff between reliance on individual experience and social cues. These findings provide an empirical link between behavioral innovation, social learning capacities, and brain size in mammals. The ability to learn from others, invent new behaviors, and use tools may have played pivotal roles in primate brain evolution.
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Plotnik, J. M., de Waal, F. B. M., & Reiss, D. (2006). Self-recognition in an Asian elephant. Proc. Natl. Acad. Sci. U.S.A., 103(45), 17053–17057.
Abstract: Considered an indicator of self-awareness, mirror self-recognition (MSR) has long seemed limited to humans and apes. In both phylogeny and human ontogeny, MSR is thought to correlate with higher forms of empathy and altruistic behavior. Apart from humans and apes, dolphins and elephants are also known for such capacities. After the recent discovery of MSR in dolphins (Tursiops truncatus), elephants thus were the next logical candidate species. We exposed three Asian elephants (Elephas maximus) to a large mirror to investigate their responses. Animals that possess MSR typically progress through four stages of behavior when facing a mirror: (i) social responses, (ii) physical inspection (e.g., looking behind the mirror), (iii) repetitive mirror-testing behavior, and (iv) realization of seeing themselves. Visible marks and invisible sham-marks were applied to the elephants' heads to test whether they would pass the litmus “mark test” for MSR in which an individual spontaneously uses a mirror to touch an otherwise imperceptible mark on its own body. Here, we report a successful MSR elephant study and report striking parallels in the progression of responses to mirrors among apes, dolphins, and elephants. These parallels suggest convergent cognitive evolution most likely related to complex sociality and cooperation.
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Scheffer, M., & van Nes, E. H. (2006). Self-organized similarity, the evolutionary emergence of groups of similar species. Proc. Natl. Acad. Sci. U.S.A., 103(16), 6230–6235.
Abstract: Ecologists have long been puzzled by the fact that there are so many similar species in nature. Here we show that self-organized clusters of look-a-likes may emerge spontaneously from coevolution of competitors. The explanation is that there are two alternative ways to survive together: being sufficiently different or being sufficiently similar. Using a model based on classical competition theory, we demonstrate a tendency for evolutionary emergence of regularly spaced lumps of similar species along a niche axis. Indeed, such lumpy patterns are commonly observed in size distributions of organisms ranging from algae, zooplankton, and beetles to birds and mammals, and could not be well explained by earlier theory. Our results suggest that these patterns may represent self-constructed niches emerging from competitive interactions. A corollary of our findings is that, whereas in species-poor communities sympatric speciation and invasion of open niches is possible, species-saturated communities may be characterized by convergent evolution and invasion by look-a-likes.
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