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VanDierendonck, M. C., de Vries, H., Schilder, M.B.H. (1995). An Analysis of Dominance, Its Behavioural Parameters and Possible Determinants in a Herd of Icelandic horses in captivity. Netherl. J. Zool., 45(3-4), 362–385.
Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
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Wolter, R., & Krueger, K. (2015). The analysis of social bonds in feral horses. In Proceedings of the 3. International Equine Science Meeting.
Abstract: In many social mammals, individuals preferentially affiliate with a small subset of available partners instead of distributing their social behaviors equally among all group members. The resulting social bonds have been investigated in several mammalian taxa, especially in primate societies, but also in other taxa such as birds, dolphins, rodents and ungulates. In feral horses, a great number of studies on social bonds can be found, but with a huge variety between methods for the analysis. There seems to be a lack of a clear and common definition of social bonds in horses and of comparable analyses. For example, there are irregularities between the studies regarding the research designs, the selection of recording methods and the interpretation of the measurements. Mutual grooming is used most often for the analysis of social relationships in many species. As mutual grooming is rare in horses, especially measurements of spatial proximity are commonly used for the analysis of social bonds in addition to other behavioral patterns. However, the combination of mutual grooming and nearest neighborhood analyses for the analysis of social bond is debatable, as in contrast to mutual grooming, which must occur deliberately by both grooming partners, the spatial distribution can be influenced by one partner alone, which may even force the other horse to keep a certain distance or to stay in close proximity.
In this study, we investigated the comparability of mutual grooming and nearest neighborhood data for social bond analyses in feral horses. Therefore, we observed five groups of semi-wild living Przewalski’s horses and six groups of feral horses.
We analysed the horses’ social ranks by applying an Average Dominance Index, we recorded the distances between the animals and observed the number of mutual grooming events as well as friendly approaches.
Our results show that there was only a weak correlation between the frequency of staying in nearest neighborhood and mutual grooming in all observed horse groups. In contrast to this, the correlation between the number of friendly approaches and mutual grooming events was higher in most groups.
Hierarchies did not affect social bonds, as mutual grooming was similarly induced by higher and lower ranking animals and the social rank did not affect the choice of the grooming partner. Similarly, likelihoods of staying in the neighborhood of particular animals were not affected by the animals’ social rank.
The grooming frequencies differed between the different horse groups and between the individual horses living in the particular groups. They seem to be effected by individual predisposition.
Altogether we suggest that the ratio of mutual grooming seems to be a better indicator for social bonds in feral horses than the frequency of staying in the nearest neighborhood. Mutual grooming occurs deliberately and is bidirectional, whereas nearest neighborhoods could be enforced and unidirectional. For the calculation of social bonds in horses, we consider it to be more reliable to combine the frequency of mutual grooming with the frequency of friendly approaches.
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Creel, S. (2001). Social dominance and stress hormones. Trends. Ecol. Evol, 16(9), 491–497.
Abstract: In most cooperatively breeding birds and mammals, reproductive rates are lower for social subordinates than for dominants, and it is common for reproduction in subordinates to be completely suppressed. Early research conducted in captivity showed that losing fights can increase glucocorticoid (GC) secretion, a general response to stress. Because GCs can suppress reproduction, it has been widely argued that chronic stress might underlie reproductive suppression of social subordinates in cooperative breeders. Contradicting this hypothesis, recent studies of cooperative breeders in the wild show that dominant individuals have elevated GCs more often than do subordinates. The findings that elevated GCs can be a consequence of subordination or a cost of dominance complicate the conventional view of social stress, with broad ramifications for the evolution of dominance and reproductive suppression.
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Hemelrijk, C. K., Wantia, J., & Gygax, L. (2005). The construction of dominance order: comparing performance of five methods using an individual-based model. Behaviour, 142(8), 1043–1064.
Abstract: In studies of animal behaviour investigators correlate dominance with all kinds of behavioural
variables, such as reproductive success and foraging success. Many methods are used to
produce a dominance hierarchy from a matrix reflecting the frequency of winning dominance
interactions. These different methods produce different hierarchies. However, it is difficult to
decide which ranking method is best. In this paper, we offer a new procedure for this decision:
we use an individual-based model, called DomWorld, as a test-environment. We choose this
model, because it provides access to both the internal dominance values of artificial agents
(which reflects their fighting power) and the matrix of winning and losing among them and,
in addition, because its behavioural rules are biologically inspired and its group-level patterns
resemble those of real primates. We compare statistically the dominance hierarchy based on
the internal dominance values of the artificial agents with the dominance hierarchy produced
by ranking individuals by (a) their total frequency of winning, (b) their average dominance
index, (c) a refined dominance index, the David`s score, (d) the number of subordinates each
individual has and (e) a ranking method based on maximizing the linear order of the hierarchy.
Because dominance hierarchies may differ depending on group size, type of society, and the
interval of study, we compare these ranking methods for these conditions.We study complete
samples as well as samples randomly chosen to resemble the limitations of observing real
animals. It appears that two methods of medium complexity (the average dominance index
and David`s score) lead to hierarchical orders that come closest to the hierarchy based on
internal dominance values of the agents. We advocate usage of the average dominance index,
because of its computational simplicity.
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Flauger, B., & Krueger, K. (2012). Social feeding decisions in horses (Equus caballus). In Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Like many other herbivores equids feed on rather evenly distributed resources. Especially in ruminants several studies have proved the influence of social organisations, rank, sex and the depletion of feeding sites on the feeding behaviour of individuals. However, it is not yet understood whether social aspects affect horses´ feeding decisions. Horses roam on vast habitats with constantly changing vegetation. In non-competitive situations domestic horses tend to return to the same feeding site until it is overgrazed. Whereas, for competition over limited food the social status of the individuals appears to be important. Curiosity about the influence of social rank and different social feeding conditions on the horses´ feeding decisions between two buckets, equally filled with high-quality surplus food, led us to create the test situation described here. The observer horses were alternately tested with a dominant and a subordinate demonstrator placed in one of three different positions. We conclude that domestic horses use cognitive strategic decision making in order to decide where to feed in a social feeding situation. When possible they tend to return to the same, continuously supplied feeding site and switch to an “avoidance tendency” when another horse is already feeding from it or in the presence of a dominant horse. Thus the position and the social rank of conspecifics affect the feeding strategy of horses.
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Krueger, K., & Flauger, B. (2008). Social feeding decisions in horses (Equus caballus). Behav. Process., 78(1), 76–83.
Abstract: Like many other herbivores, in a natural environment equids feed on rather evenly distributed resources. However, the vegetation in their vast habitats constantly changes. If food is plentiful only little competition occurs over food, and in non-competitive situations domestic horses tend to return to the same feeding site until it is overgrazed. In contrast, they compete over limited food for which the social status of the individuals appears to be important. Especially in ruminants several studies have proved an influence of social organisations, rank, sex and the depletion of feeding sites on the feeding behaviour of individuals. However, it is not yet understood whether and how social aspects affect horses“ feeding decisions. Curiosity about the influence of social rank on the horses” feeding decisions between two, equally with high-quality surplus food-filled buckets placed in different social feeding conditions, led us to create the test below. The observer horses were alternately tested with a dominant and a subordinate demonstrator placed in one of the three different positions. We conclude that domestic horses use social cognition and strategic decision making in order to decide where to feed in a social feeding situation. When possible they tend to return to the same, continuously supplied feeding site and switch to an “avoidance tendency” in the presence of dominant horses or when another horse is already feeding there. Thus, the social rank and the position of conspecifics affect the feeding strategy of horses.
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Keil, N. M., Sambraus, H.H. (1998). “Intervenors” in agonistic interactions amongst domesticated goats. Z. Säugetierk., 63(5), 266–272.
Abstract: Social behaviour was observed in individually marked goats in two herds. The goats from one herd (n = 98) were horned, those of the other herd (n = 83) were polled. By recording agonistic interactions within the herds, a dominance index was determined for each animal. In both herds, intervention took place. Intervention is defined as one animal pushing in between two fighters, and thus ending the fight. More cases of intervention took place per individual animal amongst the horned goats than amongst the polled ones. Goats which intervened in fights on several occasions usually had a high dominance index. Members of the herd which were observed intervening only once had an average dominance index in both herds of almost 0.5. In some cases, goats very low in the rank order intervened a fight. Only rarely did the intervenors have a lower dominance index than the two fighters. In 103 cases, the direct dominance relationship between a fighting animal and the intervenor was known. In 95 cases (92.2%), the intervenor was dominant to the herd member in this fight and in just eight cases (7.8%), it was subordinate. It could not be determined what advantage the intervenor gained from its activity. It is possible that, at least in certain cases, a particularly relationship existed between the intervenor and one of the fighters.
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VanDierendonck, M. C., de Vries, H., & Schilder, M. B. H. (1995). An Analysis of Dominance, Its Behavioural Parameters and Possible Determinants in a Herd of Icelandic orses in Captivity. Netherl. J. Zool., 45(3-4), 362–385.
Abstract: Th e applicability of the concept of dominance was investigated in a captive herd of  Icelandic
horses and  ponies of diff erent breeds. Eight out of  behaviours possibly related
to dominance occurred frequently enough to be investigated in detail. For these eight agonistic
behaviours the coverage, the unidirectionality in the exchange, and the degree of
transitivity (Landau`s linearity index) were calculated. Four off ensive behaviours, together
with avoidance, were suitable for further analysis with regard to dominance. Th e patterns
of asymmetries with which these behaviours were exchanged were suffi ciently similar as to
justify the application of the dominance concept and to construct a (nearly) linear dominance
hierarchy. Th e rank order of the castrated stallions was completely linear, the hierarchy
of the mares was almost completely linear. Th e results suggest that off ensive and defensive
aggressive behaviours should be treated separately and that the concept of dominance
is applicable. However, ritualized formal dominance signals between adult horses appear to
be (almost) absent. Th e rank positions of the individuals were correlated with age and residency
in the herd but not with height. Middle ranking horses tended to be more frequently
in the close vicinity of another horse than high ranking or low ranking horses. Over and
above this correlation at the individual level, it was found that pairs of horses close in rank
to each other were more often also spatially close to each other. Being in oestrus did not infl
uence the dominance relationships between mares. For castrated stallions the rank positions
were correlated with the age at which they were castrated. Th is suggests that in male
horses experience prior to neutering infl uences the behaviour afterwards.
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Christensen, J. W., Ahrendt, L. P., Lintrup, R., Gaillard, C., Palme, R., & Malmkvist, J. (2012). Does learning performance in horses relate to fearfulness, baseline stress hormone, and social rank? In Applied Animal Behaviour Science (Vol. 140, pp. 44–52).
Abstract: The ability of horses to learn and remember new tasks is fundamentally important for their use by humans. Fearfulness may, however, interfere with learning, because stimuli in the environment can overshadow signals from the rider or handler. In addition, prolonged high levels of stress hormones can affect neurons within the hippocampus; a brain region central to learning and memory. In a series of experiments, we aimed to investigate the link between performance in two learning tests, the baseline level of stress hormones, measured as faecal cortisol metabolites (FCM), fearfulness, and social rank. Twenty-five geldings (2 or 3 years old) pastured in one group were included in the study. The learning tests were performed by professional trainers and included a number of predefined stages during which the horses were gradually trained to perform exercises, using either negative (NR) or positive reinforcement (PR). Each of the learning tests lasted 3 days; 7min/horse/day. The NR test was repeated in a novel environment. Performance, measured as final stage in the training programme, and heart rate (HR) were recorded. Faeces were collected on four separate days where the horses had been undisturbed at pasture for 48h. Social rank was determined through observations of social interactions during feeding. The fear test was a novel object test during which behaviour and HR were recorded. Performance in the NR and PR learning tests did not correlate. In the NR test, there was a significant, negative correlation between performance and HR in the novel environment (rS=-0.66, P<0.001, i.e. nervous horses had reduced performance), whereas there was no such correlation in the home environment (both NR and PR). Behavioural reactions in the fear test correlated significantly with performance in the NR test in the novel environment (e.g. object alertness and final stage: rS=-0.43, P=0.04), suggesting that performance under unfamiliar, stressful conditions may be predicted by behavioural responses in a fear test. There was a negative correlation between social rank and baseline stress hormones (rS=-0.43, P=0.04), i.e. high rank corresponded to low FCM concentrations, whereas neither rank nor FCM correlated with fearfulness or learning performance. We conclude that performance under stressful conditions is affected by activation of the sympathetic nervous system during training and related to behavioural responses in a standardised fear test. Learning performance in the home environment, however, appears unrelated to fearfulness, social rank and baseline FCM levels.
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Amici, F., Widdig, A., Lehmann, J., & Majolo, B. (2019). A meta-analysis of interindividual differences in innovation. Anim. Behav., 155, 257–268.
Abstract: The ability to innovate and the social transmission of innovations have played a central role in human evolution. However, innovation is also crucial for other animals, by allowing them to cope with novel socioecological challenges. Although innovation plays such a central role in animals' lives, we still do not know the conditions required for innovative behaviour to emerge. Here, we focused on interindividual differences in innovation by (1) extensively reviewing existing literature on innovative behaviour in animals and (2) quantitatively testing the different evolutionary hypotheses that have been proposed to explain interindividual variation in innovation propensity during foraging tasks. We ran a series of phylogenetically controlled mixed-effects meta-regression models to determine which hypotheses (if any) are supported by currently available empirical studies. Our analyses show that innovation is more common in individuals that are older and belong to the larger sex, but also in more neophilic and/or explorative individuals. Moreover, these effects change depending on the study setting (i.e. wild versus captive). Our results provide no clear support to the excess of energy or the bad competitor hypotheses and suggest that study setting and interindividual differences in traits related to personality are also important predictors of innovation.
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