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Moehlman, P. D. (1998). Behavioral patterns and communication in feral asses (Equus africanus). Appl. Anim. Behav. Sci., 60(2-3), 125–169.
Abstract: The behavior of feral populations of the African wild ass (Equus africanus) were studied in the Northern Panamint Range of Death Valley National Monument for 20 months from 1970 to 1973 [Moehlman, P.D., 1974. Behavior and ecology of feral asses (Equus asinus). PhD dissertation, University of Wisconsin, Madison, 251 pp.; Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Natl. Geogr. Soc. Res. Reports, 1970: 405-411]. Maintenance behavior is described and behavior sequences that were used in social interactions are quantified by sex and age class. Agonistic, sexual, and greeting behavior patterns are described and analyzed in conjunction with the responses they elicited. Mutual grooming mainly occurred between adult males, and between females and their offspring. Five types of vocalizations were distinguished: brays, grunts, growls, snorts, and whuffles. A second population was studied for 1 month on Ossabaw Island, GA (Moehlman, 1979). This population had more permanent social groups and had a higher rate of mutual grooming and foal social play.
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Khalil, A. M., & Kaseda, Y. (1998). Early experience affects developmental behaviour and timing of harem formation in Misaki horses. Appl. Anim. Behav. Sci., 59(4), 253–263.
Abstract: A study was made of the behavior of young male Misaki feral horses in the developmental stage, by observing nine of them once a week from January 1988 to December 1996. The relationship between behavior before separation and in the developmental stage was also investigated. This stage begins just after young males separate from their natal band or mothers, and it continues until they start to form harems. The duration of the developmental stage in the study ranged from 0.6 to 3.9 years, depending on the age of the young males at the time of separation. Young males associated with three types of social groups at the beginning of the developmental stage, according to their social groups before separation. These were bachelor groups, harem groups and wandering female groups. During this period, males joined the three groups, mixed sex groups and sometimes were solitary. It was considered that these associations provided a good opportunity for males to acquire different behavioral patterns and experiences before they entered the next stage. Depending on the groups with which they associated, young males that spent more time with bachelor groups had the longest average developmental stage. They associated with harem groups more often during the breeding season and more frequently with other groups or were solitary during the non-breeding season. This may be a transition period because by the end of this stage all males had spent time in solitude before forming their own harem bands.
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Dugatkin, L. A., & Mesterton-Gibbons, M. (1996). Cooperation among unrelated individuals: reciprocal altruism, by-product mutualism and group selection in fishes. Biosystems, 37(1-2), 19–30.
Abstract: Cooperation among unrelated individuals can evolve not only via reciprocal altruism but also via trait-group selection or by-product mutualism (or some combination of all three categories). Therefore the (iterated) prisoner's dilemma is an insufficient paradigm for studying the evolution of cooperation. We replace this game by the cooperator's dilemma, which is more versatile because it enables all three categories of cooperative behavior to be examined within the framework of a single theory. Controlled studies of cooperation among fish provide examples of each category of cooperation. Specifically, we describe reciprocal altruism among simultaneous hermaphrodites that swap egg parcels, group-selected cooperation among fish that inspect dangerous predators and by-product mutualism in the cooperative foraging of coral-reef fish.
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Foster, T. M., Temple, W., Cameron, B., & Poling, A. (1997). Demand curves for food in hens: Similarity under fixed-ratio and progressive-ratio schedules. Behav. Process., 39(2), 177–185.
Abstract: Demand curves were generated for five domestic hens under progressive-ratio 5 schedules of food delivery and under fixed-ratio schedules of food delivery that began at fixed-ratio 5 and were incremented by 5 each session. All sessions ended after 10 consecutive minutes without a response. Although response rates at a given ratio were higher under the progressive-ratio schedule, all hens completed higher ratios under the fixed-ratio schedule. Similar, but not identical, demand curves were generated under progressive-ratio and fixed-ratio schedules. Under both schedules, consumption (reinforcers earned) decreased as cost (ratio size) increased. Data generally were well described by an equation in which elasticity of demand is constant, although an equation in which elasticity could vary accounted for slightly more of the variance.
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Robinson, T. A., Foster, T. M., Temple, W., & Poling, A. (1995). Performance of domestic hens under progressive-ratio schedules of food delivery. Behav. Process., 34(3), 233–239.
Abstract: Domestic hens were exposed to progressive-ratio 2 and progressive-ratio 10 schedules of food delivery with different initial ratios (2, 10, 20, 30, and 40). Breaking points, defined as the largest ratios completed before responding ceased for 600 consecutive seconds, were recorded under all conditions. In general, breaking points were higher under the PR 10 schedule than under the PR 2 schedule, and the value of the initial ratio did not systematically affect the breaking point. The former finding suggests that relative satiation affected breaking points in the present study, but the latter finding suggests that the primary determinant was the `price' of the reinforcer, defined in terms of the number of responses required to produce it. Breaking points were similar under conditions where initial ratios changed from session to session and under more conventional conditions, where initial ratios remained unchanged over several sessions.
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Visser, E. K., van Reenen, C. G., van der Werf, J. T. N., Schilder, M. B. H., Knaap, J. H., Barneveld, A., et al. (2002). Heart rate and heart rate variability during a novel object test and a handling test in young horses. Physiol. Behav., 76(2), 289–296.
Abstract: Forty-one Dutch Warmblood immature horses were used in a study to quantify temperamental traits on the basis of heart rate (HR) and heart rate variability (HRV) measures. Half of the horses received additional training from the age of 5 months onwards; the other half did not. Horses were tested at 9, 10, 21 and 22 months of age in a novel object and a handling test. During the tests, mean HR and two heart variability indices, e.g. standard deviation of beat-to-beat intervals (SDRR) and root mean square of successive beat-to-beat differences (rMSSD), were calculated and expressed as response values to baseline measures. In both tests, horses showed at all ages a significant increase in mean HR and decrease in HRV measures, which suggests a marked shift of the balance of the autonomic nervous system towards a sympathetic dominance. In the novel object test, this shift was more pronounced in horses that had not been trained. Furthermore, statistical analysis showed that the increase in mean HR could not be entirely explained by the physical activity. The additional increase in HR, the nonmotor HR, was more pronounced in the untrained horses compared to the trained. Hence, it is suggested that this nonmotor HR might be due to the level of emotionality. HR variables showed consistency between years, as well as within the second year. These tests bring about a HR response in horses, part of which may indicate a higher level of emotionality; and horses show individual consistency of these HR variables over ages. Therefore, it is concluded that mean HR and HRV measures used with these tests quantify certain aspects of a horse's temperament.
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Shultz, S., & Dunbar, R. I. M. (2006). Both social and ecological factors predict ungulate brain size. Proc Biol Sci, 273(1583), 207–215.
Abstract: Among mammals, the members of some Orders have relatively large brains. Alternative explanations for this have emphasized either social or ecological selection pressures favouring greater information-processing capacities, including large group size, greater foraging efficiency, higher innovation rates, better invasion success and complex problem solving. However, the focal taxa for these analyses (primates, carnivores and birds) often show both varied ecological competence and social complexity. Here, we focus on the specific relationship between social complexity and brain size in ungulates, a group with relatively simple patterns of resource use, but extremely varied social behaviours. The statistical approach we used, phylogenetic generalized least squares, showed that relative brain size was independently associated with sociality and social complexity as well as with habitat use, while relative neocortex size is associated with social but not ecological factors. A simple index of sociality was a better predictor of both total brain and neocortex size than group size, which may indicate that the cognitive demands of sociality depend on the nature of social relationships as well as the total number of individuals in a group.
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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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Mech L.D. (2000). Leadership in Wolf, Canis lupus, Packs. Can Field Nat, 114(2), 259–263.
Abstract: I examine leadership in Wolf (Canis lupus) packs based on published observations and data gathered during summers from 1986 to 1998 studying a free-ranging pack of Wolves on Ellesmere Island that were habituated to my presence. The breeding male tended to initiate activities associated with foraging and travel, and the breeding female to initiate, and predominate in, pup care and protection. However, there was considerable overlap and interaction during these activities such that leadership could be considered a joint function. In packs with multiple breeders, quantitative information about leadership is needed.
Keywords: Wolf, Canis lupus, leadership, behavior, foraging, movements, pup care, provisioning, sociality, reproduction, breeding, Northwest Territories.
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Reboreda, J. C., Clayton, N. S., & Kacelnik, A. (1996). Species and sex differences in hippocampus size in parasitic and non-parasitic cowbirds. Neuroreport, 7(2), 505–508.
Abstract: To test the hypothesis that selection for spatial abilities which require birds to locate and to return accurately to host nests has produced an enlarged hippocampus in brood parasites, three species of cowbird were compared. In shiny cowbirds, females search for host nests without the assistance of the male; in screaming cowbirds, males and females inspect hosts' nests together; in bay-winged cowbirds, neither sex searches because this species is not a brood parasite. As predicted, the two parasitic species had a relatively larger hippocampus than the non-parasitic species. There were no sex differences in relative hippocampus size in screaming or bay-winged cowbirds, but female shiny cowbirds had a larger hippocampus than the male.
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