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McDonnell, S. M., & Haviland, J. C. S. (1995). Agonistic ethogram of the equid bachelor band. Appl. Anim. Behav. Sci., 43(3), 147–188.
Abstract: An ethogram of agonistic and related behaviors among equid bachelor band members was developed. Several key English-language studies on equids were reviewed to derive a preliminary inventory of specific behaviors to be included in the ethogram. A bachelor band of domestic pony stallions pastured together was observed for approximately 50 daylight hours to obtain detailed descriptions of each behavior, enable photographic and video documentation of behaviors, and identify any behaviors to be added to the preliminary inventory. An initial draft of the ethogram was sent to 65 equine researchers for review. Twenty-eight critical reviews were received and their suggestions considered for the final draft. A total of 49 elemental behaviors including five distinct vocalizations was included in the ethogram. Three complex behavioral sequences were also included. Most of the behaviors catalogued from the direct observation of pastured pony stallions were also found in the equid literature. For many, references to these behaviors specifically among males or bachelor band members were not found. The results offer a practical tool for quantitative research and other studies of equid inter-male behavior as well as for teaching of equid behavior, and should facilitate progress toward development of a complete ethogram for the horse and other equids.
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Noë, R. (1992). Alliance formation among male hamadryas baboons: shopping for profitable partners. In A. H. Harcourt, & F. B. M. deWaal (Eds.), Coalitions and alliances in humans and other animals (pp. 284–321). Oxford: Oxford University Press.
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Feh, C. (1999). Alliances and reproductive success in Camargue stallions. Anim. Behav., 57(3), 705–713.
Abstract: A study of a herd of Camargue horsesEquus caballus, showed that while the majority of high-ranking stallions held single-male harems, some sons of low-ranking mares, being low ranking themselves, formed alliances that could last a lifetime. The two stallions were each other's closest associate and preferential grooming partner. Alliances were based on coalitions in which either both partners confronted an intruder synchronously or the dominant of the pair tended the female(s) while the subordinate simultaneously displayed towards the rival. Alliance partners were of similar age but were not more closely related to each other than to other stallions in the herd. Long-term paternity data revealed that subordinates sired close to a quarter of the foals born into the alliance group, and significantly more foals than low-ranking stallions in the herd adopting a `sneak'-mating strategy. The dominant appeared to benefit from the presence of his subordinate partner. Fights occurred all year round, and the subordinate stallion of each alliance pair fought outside competitors more than twice as often as the dominant. Forming short-term alliances before defending mares on their own may enhance long-term reproductive success for both partners. Other benefits to both partners include higher survivorship of their foals and increased access to proven reproductive mares. These results suggest that the relationship between alliance partners is based on mutualism, but several conditions for reciprocity seem to be fulfilled: the benefit to the dominant (assistance in fights), and the benefit to the subordinate (access to reproduction), are both costly to the other partner and delayed in time.
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Feh, C. (2001). Alliances between stallions are more than just multimale groups: reply to Linklater & Cameron (2000). Anim. Behav., 61, F27–F30.
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Weatherly, J. N., Arthur, E. I. L., & Tischart, L. M. (2003). Altering “motivational” variables alters induction produced by upcoming food-pellet reinforcement. Anim. Cogn., 6(1), 17–26.
Abstract: Previous research has demonstrated that rats will increase their rates of lever pressing for sucrose rewards in the first half of an experimental session when food pellets, rather than the same sucrose, continually serve as the reward in the second half of the session. This effect has been coined induction, and the present study investigated whether it could be altered by altering “motivational” variables. Experiment 1 manipulated subjects' motivation by altering, across conditions, their level of food deprivation. Predictably, the size of induction varied directly with level of deprivation. Experiments 2 and 3 manipulated subjects' motivation by feeding them food pellets and sucrose, respectively, prior to their responding in the experimental session. These pre-session feedings decreased the size of the observed induction in both experiments. The results from the present study indicate that the size of induction is correlated with subjects' motivation to respond for the available reinforcers. They are also consistent with the idea that operant processes underlie the effect. The notion that induction might encompass the concept of “anticipation” is also discussed.
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Lingle, S., Rendall, D., & Pellis, S. M. (2007). Altruism and recognition in the antipredator defence of deer: 1. Species and individual variation in fawn distress calls. Anim. Behav., 73(5), 897–905.
Abstract: Mule deer, Odocoileus hemionus, females actively defend fawns against predators, including nonoffspring conspecific fawns and heterospecific white-tailed deer, O. virginianus, fawns. We hypothesized that the defence of nonoffspring fawns was due to a recognition error. During a predator attack, females may have to decide whether to defend a fawn with imperfect information on its identity obtained from hearing only a few distress calls. We examined fawn distress calls to determine whether calls made by the two species and by different individuals within each species were acoustically distinctive. The mean and maximum fundamental frequencies of mule deer fawns were nearly double those of white-tailed deer fawns, with no overlap, enabling us to classify 100% of calls to the correct species using a single trait. A large proportion of calls was also assigned to the correct individual using a multivariate analysis (66% and 70% of mule deer and white-tailed deer fawns, respectively, chance = 6% and 10%); however, there was considerable statistical uncertainty in the probability of correct classification. We observed fawns approach conspecific females in an attempt to nurse; females probed most offspring fawns with their noses before accepting them, and always probed nonoffspring fawns before rejecting them, suggesting that close contact and olfactory information were needed to unequivocally distinguish nonoffspring from offspring fawns. Taken together, these results suggest that acoustic variation alone would probably be sufficient to permit rapid and reliable species discrimination, but it may not be sufficient for mothers to unequivocally distinguish their own fawn from conspecific fawns.
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Lingle, S., Rendall, D., Wilson, W. F., DeYoung, R. W., & Pellis, S. M. (2007). Altruism and recognition in the antipredator defence of deer: 2. Why mule deer help nonoffspring fawns. Anim. Behav., 73(5), 907–916.
Abstract: Both white-tailed deer, Odocoileus virginianus, and mule deer, O. hemionus, females defend fawns against coyotes, Canis latrans, but only mule deer defend nonoffspring conspecific and heterospecific fawns. During a predator attack, females may have to decide whether to defend a fawn while having imperfect information on its identity obtained from hearing a few distress calls. Although imperfect recognition can influence altruistic behaviour, few empirical studies have considered this point when testing functional explanations for altruism. We designed a series of playback experiments with fawn distress calls to test alternative hypotheses (by-product of parental care, kin selection, reciprocal altruism) for the mule deer's defence of nonoffspring, specifically allowing for the possibility that females mistake these fawns for their own. White-tailed deer females approached the speaker only when distress calls of white-tailed deer fawns were played and when their own fawn was hidden, suggesting that fawn defence was strictly a matter of parental care in this species. In contrast, mule deer females responded similarly and strongly, regardless of the caller's identity, the female's reproductive state (mother or nonmother) or the presence of their own offspring. The failure of mule deer females to adjust their responses to these conditions suggests that they do not defend nonoffspring because they mistake them for their own fawns. The lack of behavioural discrimination also suggests that kin selection, reciprocal altruism and defence of the offspring's area are unlikely to explain the mule deer's defence of nonoffspring. We identify causal and functional questions that still need to be addressed to understand why mule deer defend fawns so indiscriminately.
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Santi, A., Stanford, L., & Symons, J. (1998). An analysis of confusion errors in many-to-one matching with temporal and nontemporal samples. Anim. Cogn., 1(1), 37–46.
Abstract: In experiment 1, pigeons were trained to match temporal (2, 8, and 10 s of houselight) and location (feeder light, left key, center key illumination) samples to color comparison stimuli. Red choices were correct following the 2-s and feeder light, orange choices were correct following the 8-s and center key, and green choices were correct following the 10-s and left key. Samples that were harder to discriminate (8- vs 10-s, and left vs center key) were mapped onto comparisons that were easy to discriminate (orange vs green), while samples that were easier to discriminate (2- vs 8-s, and feeder light vs left key) were mapped onto comparisons that were hard to discriminate(red vs orange). The pattern of errors for temporal and location samples indicated that these samples were not represented by a common code even though they were associated with the same comparison stimuli. In experiment 2, the same pigeons were trained with visual samples in which samples that were hard to discriminate (triangle vs circle) were mapped onto comparisons that were easy to discriminate (orange vs green), while samples that were easy to discriminate(plus vs triangle) were mapped onto comparisons that were hard to discriminate (red vs orange). Following acquisition of the visual discrimination, the temporal samples were re-introduced and many-to-one training was continued. During delay testing, the pattern of errors for temporal and visual samples was equivalent and consistent with the hypothesis that visual samples were being coded in terms of the duration appropriate for the temporal sample with which it shared a common comparison response. Data from no-sample test sessions ruled out a simple response bias explanation of the data. The properties of common codes for temporal and nontemporal events can be somewhat flexible and more complicated than previously envisaged.
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Lonsdorf, E. V., Ross, S. R., Linick, S. A., Milstein, M. S., & Melber, T. N. (2009). An experimental, comparative investigation of tool use in chimpanzees and gorillas. Anim. Behav., 77(5), 1119–1126.
Abstract: Studies of ape tool use have been conducted in captivity since the early 1900s and in the wild since the 1960s. Chimpanzees are the most prolific tool users among the apes, and are known to use more tools than any other nonhuman animal. In contrast, reports of gorilla tool use are rare both in wild and captive settings. Studies of the processes involved in tool use learning have been limited in the wild by the lack of ability to control several unpredictable variables, and in captivity by tool use opportunities that are often presented in non-naturalistic contexts. We attempted to address both of these limitations by providing naïve subjects with a naturalistic tool use device (built to simulate a termite mound) while housed in a more natural social setting to approximate how learning would occur in the wild. Both gorillas and chimpanzees participated in the experiment to allow comparative analyses of acquisition of tool behaviour and the factors that may affect acquisition. Both species showed low frequencies of interaction with the mound in the baseline condition, before baiting with a food reward. Once baited, chimpanzees both attempted and succeeded to extract the reward more quickly than did gorillas. The number of social group members at the mound was significantly higher for chimpanzees than for gorillas and may have affected skill acquisition. We advocate that comparative approaches to skill acquisition and learning are valuable, but that researchers need to be cognizant of species differences in social structure that may affect results.
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de Vries, H. (1995). An improved test of linearity in dominance hierarchies containing unknown or tied relationships. Anim. Behav., 50(5), 1375–1389.
Abstract: Appleby (1983, Anim. Behav., 31, 600-608) described a statistical test, based on the work of Kendall (1962, Rank Correlation Methods), for the significance of linearity in dominance hierarchies. He suggested that unknown relationships should be assigned the value 1/2 and that subsequently the same test procedure can be used. In this paper it is shown that incorrect results are obtained by this method whenever there are unknown relationships. Values of the linearity index are systematically too low. P-values can be too high (underestimating the significance) or too low (overestimating), and seem to differ by not much more than a factor two (respectively a half) from the correct P-value. An improved method is developed for testing linearity in a set of dominance relationships containing unknown relationships. Furthermore, it is argued that, if one admits the possibility of tied dominance relationships, which should indeed be assigned the value 1/2, Landau's linearity index is to be preferred to Kendall's index. A randomization test is developed for assessing the significance of linearity or non-linearity in a set of dominance relationships containing unknown or tied relationships. The test statistic employed in this testing procedure is based on Landau's linearity index, but takes the unknown and tied relationships into account.
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