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Jonart, L. M., Hill, G. E., & Badyaev, A. V. (2007). Fighting ability and motivation: determinants of dominance and contest strategies in females of a passerine bird. Anim. Behav., 74(6), 1675–1681.
Abstract: The communication of aggressive motivation or fighting ability has important fitness consequences for competing animals. Selection should favour rapid and honest communication between opponents to settle dominance relationships while avoiding prolonged and intense fighting. We investigated factors that influence fighting strategies and contest outcomes in female house finches, Carpodacus mexicanus, specifically focusing on the following questions. (1) What social contexts trigger an aggressive response? (2) Does body size and condition contribute to female fighting ability? (3) Do contextual factors, such as mate presence, nest status, nest proximity, and site experience contribute to fighting motivation? (4) Does contest intensity and duration increase as the differences in fighting ability between opponents decrease? (5) What is the relative contribution of fighting ability and aggressive motivation to the outcome of a contest? We found that aggression was triggered most frequently by female intrusions in the vicinity of nest and by extrapair female intrusions on an established pair. Female fighting and contest outcomes were strongly influenced by body condition and body size, and females were more motivated to initiate fights and won more contests when their mates were present. Females at the later breeding stages and those fighting closer to their nests were dominant and won more fights compared to females at earlier breeding stages or further from their nests. Females initiated a greater proportion of contests against opponents with similar local familiarity and breeding history. Escalated and prolonged contests, while rare, occurred exclusively between females of the most similar body size and condition. When differences in body condition between opponents are not easily perceived, contestants might escalate contests for more reliable assessments of relative fighting ability. Finally, body condition was not a strong determinant of contest outcome in the contexts with easily assessed differences in the resource value (e.g. mate presence), but without these motivational differences, body condition was the ultimate determinant of contest outcomes.
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Dreier, S., van Zweden, J. S., & D'Ettorre, P. (2007). Long-term memory of individual identity in ant queens. Biol Lett, 3(5), 459–462.
Abstract: Remembering individual identities is part of our own everyday social life. Surprisingly, this ability has recently been shown in two social insects. While paper wasps recognize each other individually through their facial markings, the ant, Pachycondyla villosa, uses chemical cues. In both species, individual recognition is adaptive since it facilitates the maintenance of stable dominance hierarchies among individuals, and thus reduces the cost of conflict within these small societies. Here, we investigated individual recognition in Pachycondyla ants by quantifying the level of aggression between pairs of familiar or unfamiliar queens over time. We show that unrelated founding queens of P. villosa and Pachycondyla inversa store information on the individual identity of other queens and can retrieve it from memory after 24h of separation. Thus, we have documented for the first time that long-term memory of individual identity is present and functional in ants. This novel finding represents an advance in our understanding of the mechanism determining the evolution of cooperation among unrelated individuals.
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Li, C., Jiang, Z., Tang, S., & Zeng, Y. (2007). Influence of enclosure size and animal density on fecal cortisol concentration and aggression in Pere David's deer stags. Gen Comp Endocrinol, 151(2), 202–209.
Abstract: We investigated the impact of enclosure size and animal density on behavior and adrenocortical secretion in Pere David's deer in Dafeng Nature Reserve, China. From February 15 to April 16 in 2004, we conducted two experiments. First, we studied maintenance behavior and conflict behavior of Pere David's deer stags in a large enclosure (200 ha) with low animal density (0.66 deer/ha) and a small display pen (0.75 ha) with high animal density (25.33 deer/ha). The maintenance behavior we recorded included standing, locomotion, foraging and rest. During the behavioral observations, we collected fresh voided fecal samples from the stags periodically, and analyzed the fecal cortisol concentrations in those samples using radioimmunoassay technique. Second, we monitored the fecal cortisol concentrations of one group of stags (12 deer lived in an enclosure of 100 ha) before and after transferred into a small pen (0.5 ha). We found that in the first experiment: (1) there were significant differences in standing and rest whereas no significant differences of locomotion and foraging between the free-ranging group and the display group; (2) frequency of conflict behavior in the display group was significantly higher than those in the free-ranging group; and (3) fecal cortisol concentration of the display group (326.17+/-16.98 ng/g dry feces) was significantly higher than that of the free-ranging group (268.98+/-15.21 ng/g dry feces). In the second experiment, there was no significant difference of the fecal cortisol concentrations among sampling days, but the mean fecal cortisol concentration of the day after transferring (337.46+/-17.88 ng/g dry feces) was significantly higher than that of the day before transferring (248.44+/-7.99 ng/g dry feces). Comparison with published findings, our results indicated that enclosure size and animal density affect not only behaviors, but also adrenocortical secretion in Pere David's deer. Small living space with high animal density may impose physiological stress to captive Pere David's deer. Moreover, long-term physiological stress and increase of conflict behavior may subsequently affect survival and reproduction of the deer.
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Ostner, J., Heistermann, M., & Schülke, O. (2008). Dominance, aggression and physiological stress in wild male Assamese macaques (Macaca assamensis). Hormones and Behavior, 54(5), 613–619.
Abstract: In group-living animals relative rank positions are often associated with differences in glucocorticoid output. During phases of social stability, when dominance positions are clear and unchallenged, subordinates often face higher costs in terms of social stress than dominant individuals. In this study we test this prediction and examine additional potential correlates of stress, such as reproductive season, age and amount of aggression received in wild, seasonally breeding Assamese macaques (Macaca assamensis). During a mating and a non-mating season we collected 394 h of focal observational data and 440 fecal samples of six adult and six large subadult males living in a multimale-multifemale group in their natural habitat in northeastern Thailand. The mating season was characterized by a general increase in aggressive behavior and glucocorticoid excretion across all males compared to the non-mating season. Among adult males, mating season glucocorticoid levels were significantly negatively related with dominance rank and positively with the amount of aggression received. Both relationships were non-significant among large subadult males. Thus, our results suggest that in adult Assamese macaques a high dominance position is not associated with high costs. Low costs of dominance might be induced by strong social bonds among top-ranking males, which exchange frequent affiliative interactions and serve as allies in coalitionary aggression against potentially rank-challenging subordinate males.
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Feuerstein, N., & Terkel, J. (2008). Interrelationships of dogs (Canis familiaris) and cats (Felis catus L.) living under the same roof. Appl. Anim. Behav. Sci., 113(1-3), 150–165.
Abstract: In the process of domestication, dogs (Canis familiaris) and cats (Felis catus) have undergone thousands of years of genetic changes that have adapted them to the human environment. Both species have acquired a global distribution and it has become quite common to find homes with the two living side by side. Nevertheless, there is widespread belief that interspecific communication between dogs and cats is problematic, stemming from their separate evolutionary development and different social structures. Consequently, many people considering possible adoption of both species are concerned about their ability to get along. Interrelationships of dogs and cats living together were studied here in an attempt to determine the main factors influencing the type of relationship likely to develop between the two species. Two approaches were used: (1) a questionnaire completed by owners of both dog(s) and cat(s), which provided a broad database of the animals' behaviors; and (2) observations carried out in participants' homes on their dog-cat interactions. Two separate ethograms for dogs and cats served for analyses of their body language. The findings revealed the following: Both species showed a similar ability to establish a relatively amicable relationship with the other species; the animals' gender had little influence on the nature of their interrelationship; and adoption of the cat prior to the dog appears to conduce to establishing an amicable relationship, as does their first encounter taking place at an early age (up to 6 months of age in cats and up to 1 year in dogs). The findings also suggest that the majority of these dogs and cats understood the particular body language displayed by one animal that has an opposite meaning for the other species; and that the earlier the age of first encounter between the two, the better this understanding. It can be concluded that exposure of both species at an early age to the presence of the other facilitates the learning of each other's body language, and the consequent establishment of an amicable relationship. A better understanding of the various factors that contribute to determining the two species' relationship should not only improve the quality of life of these pets, but also reassure and encourage more people to adopt both cat and dog.
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Hewitt, S. E., Macdonald, D. W., & Dugdale, H. L. (2009). Context-dependent linear dominance hierarchies in social groups of European badgers, Meles meles. Anim. Behav., 77(1), 161–169.
Abstract: A social hierarchy is generally assumed to exist in those mammalian societies in which the costs and benefits of group living are distributed unevenly among group members. We analysed infrared closed-circuit television footage, collected over 3 years in Wytham Woods, Oxfordshire, to test whether social groups of European badgers have dominance hierarchies. Analysis of directed aggression between dyads revealed linear dominance hierarchies in three social-group-years, but patterns within social groups were not consistent across years. Dominance hierarchies were significantly steeper than random in five out of six social-group-years. In those social-group-years where a linear hierarchy was determined, there was an effect of sex on dominance rank, with females gaining significantly higher rank than males in two social-group-years. Overall, rank was not related to age, nor did it appear to affect the likelihood of an individual being wounded, or an individual's breeding status. The latter resulted from nonorthogonality between sex and breeding status, as there were only two breeding males. Overall, hierarchies were primarily dominated by breeding females, and may occur when breeding competition arises. Relatedness, unreciprocated allogrooming and sequential allomarking were not consistently related to levels of directed aggression across social-group-years. We suggest that dominance structures within European badger groups may be context dependent, with future study required to complete our understanding of where, and when, they arise.
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Cozzi, A., Sighieri, C., Gazzano, A., Nicol, C. J., & Baragli, P. (2010). Post-conflict friendly reunion in a permanent group of horses (Equus caballus). Behav. Process., 85(2), 185–190.
Abstract: Gregarious animals living in permanent social groups experience intra-group competition. Conflicts over resources can escalate into costly aggression and, in some conditions, non-dispersive forms of conflict resolution may be favoured. Post-conflict friendly reunions, hence reconciliation, have been described in a variety of species. The aim of this study was to explore, for the first time, the occurrence of reconciliation in a group of domestic horses (Equus caballus) and learn more about strategies used to maintain group cohesion. The behaviour of seven horses living as permanent group in an enclosure for at least 2 years was observed by video for 108 h from June to August 2007. We used a Post-Conflict/Matched Control method to assess the existence of reconciliation and third-party affiliation. Behaviours recorded Post-Conflict, or during Matched Control periods, were classified as affiliative based on previous descriptions of visual communication patterns in horses. The proportion of attracted pairs over total post-conflict situations was significantly greater than the proportion of dispersed pairs, both during dyadic interactions (p < 0.001) and during triadic interactions (p = 0.002). The results of the present study show that both dyadic reconciliation and third-party post-conflict affiliative interactions form important social mechanisms for managing post-conflict situations in horses.
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Hartmann, E. (2010). Managing horses in groups to improve horse welfare and human safety. Ph.D. thesis, , .
Abstract: Managing horses in groups to improve horse welfare and human safety
:
reactions to mixing and separation
Hartmann, Elke (2010) Managing horses in groups to improve horse welfare and human safety . Doctoral diss. Dept. of Animal Environment and Health, SLU. Acta Universitatis agriculturae Sueciae vol. 2010:87.
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Abstract
The aim of this thesis was to investigate whether specific anecdotal concerns related to keeping horses in groups are supported by science and, if so, provide scientifically based recommendations that could be implemented in practice.
The aim of studies I and II was to identify methods for mixing unfamiliar horses that could minimise aggressive interactions and associated risk of injury. Results of study I revealed that pre-exposure of young horses in neighbouring boxes tended to lower contact-aggression (e.g. kicks, strikes) and biting behaviour in particular was reduced when the same pair of horses subsequently met in a paddock. This was not found when older horses were mixed (study II). Aggressive behaviour received by a new horse was not significantly different in meetings when it met one other horse compared to meeting two unfamiliar horses at the same time.
Removing a horse from a group of four in study III was generally unproblematic. Most horses approached the handler when she was catching the horse and while standing with it in the middle of the paddock. Thus, potential risk may be higher in situations when the handler remains relatively stationary, as other horses of the group have time to approach. Rank did not influence the number of horses following to the paddock gate and interactions between horses were rare.
Since horses naïve to social separation may be more difficult to handle away from the group, the objective in study IV was to investigate whether the initial presence of a companion horse would modify responses to separation. Results revealed no significant differences in heart rates and the number of training sessions required when the horses were subsequently trained in the absence of the partner compared to horses trained alone from the start.
In summary, results give little support for the original areas of concerns about mixing and separating horses. Risk of injury to both horses and humans should not be overestimated when handling horses in groups, but being aware of potential risk situations and being able to react accordingly is likely to increase horse welfare and human safety.
Faculty: Faculty of Veterinary Medicine and Animal Science
Keywords:
equine, behaviour, welfare, housing, mixing, aggression, injury, separation, habituation, learning
Agrovoc terms:
horses, behaviour, bites, injurious factors, animal learning, animal welfare, sweden
ISBN: 978-91-576-7532-3
Series.: Acta Universitatis agriculturae Sueciae
ISSN: 1652-6880
Volume: 2010:87
Papers/manuscripts:
I. Hartmann, E., Winther Christensen, J., Keeling, L.J. (2009). Social interactions of unfamiliar horses during paired encounters: Effect of pre-exposure on aggression level and so risk of injury. Applied Animal Behaviour Science 121, 214-221.
II. Hartmann, E., Rundgren, M., Keeling, L.J. (in press). Comparison of 3 methods for mixing unfamiliar horses (Equus caballus). Journal of Equine Veterinary Behavior: Clinical Applications and Research.
III. Hartmann, E., Søndergaard, E., Keeling, L.J. Identifying potential risk situations for humans when removing horses from groups. Manuscript.
IV. Hartmann, E., Christensen, J.W., Keeling, L.J. (in press). Training young horses to social separation: Effect of a companion horse on training efficiency. Equine Veterinary Journal.
Number of pages: 75
Year of publication: 2010
Language: eng
ID Code: 2396
Deposited By: Hartmann, Elke
Deposited On: 08 November 2010
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Hartmann, E., Keeling, L. J., & Rundgren, M. (2011). Comparison of 3 methods for mixing unfamiliar horses (Equus caballus). J Vet Behav Clin Appl Res, 6(1), 39–49.
Abstract: Horses are likely to exhibit aggression when meeting for the first time. Therefore, this study compared 3 methods for mixing horses to evaluate their effectiveness in reducing aggressive interactions: (1) mixing pairs of horses in a paddock (P, 10 minutes, 15 tests), (2) introducing 1 unfamiliar horse to a pair of familiar, resident horses in a paddock (PP, 10 minutes, 15 tests), (3) allowing limited physical contact between pairs of horses for a short period of pre-exposure in neighboring boxes (B, 5 minutes, 16 tests) before mixing them in a paddock (BP, 10 minutes 16 tests). A total of 16 Swedish Standardbred mares, aged 6-18 years (mean age ± SD: 11 ± 4.4), were included in the study. Half of the horses were familiar with each other (resident horses, n = 8), whereas the other half were bought in from a variety of sources (unfamiliar horses, n = 8). Social interactions, consisting of behaviors from the sender, the receiver, and the subsequent sender's response, were recorded continuously as frequencies. There were no differences in the frequencies of aggressive behaviors between the 3 mixing methods, including those aggressive behaviors in which physical contact had been attempted (kick, strike). Although resident horses were overall more aggressive (median number of aggressive behaviors per horse, 62; Q1, 36; Q3, 68.5) than unfamiliar horses (median per horse, 4; Q1, 2; Q3, 12.5) during all tests (U = 97, P = 0.003), none of the 62 tests needed to be terminated. Unfamiliar horses did not receive more aggression from resident horses in PP (mean per test ± SD: 5.1 ± 3.1) than in P (mean per test ± SD: 6.4 ± 4.9) (t = 0.63, P = 0.544). However, the behavior “attack” was more frequent in PP (median per test, 2; Q1, 0; Q3, 5) than in P (median per test, 0; Q1, 0; Q3, 1) (U = 282, P = 0.042), and “flee” was more frequent in PP (median per test, 6; Q1, 4; Q3, 8) than in P (median per test, 1; Q1, 0; Q3, 6) (U = 290, P = 0.018). Pre-exposure in boxes did not reduce aggression in BP (median per test, 7; Q1, 4.3; Q3, 11.8) as compared with P (median per test, 6; Q1, 2; Q3, 16) (U = 264, P = 0.767), but during pre-exposure in B tests, horses exchanged more nonaggressive (median per test, 2; Q1, 0.3; Q3, 4) than aggressive (median frequency of aggressive behavior, 0; Q1, 0; Q3, 1) (W = 71, P = 0.013) and mixed interactions (median per test, 0; Q1, 0; Q3, 1) (W = 92, P = 0.016) through the opening. Results suggest mixing an unfamiliar horse with 2 resident horses at the same time instead of one by one may be preferable. In this way, the total aggression received by the unfamiliar horse will potentially be less, even though aggressive interactions may be more intense.
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Flauger, B. (2011). The introduction of horses into new social groups with special regard to their stress level. Ph.D. thesis, , .
Abstract: Horses are a highly social species living in complex social systems which should require them to memorise and generalise social experiences and distinguish between familiar and unfamiliar conspecifics. In the main part of my thesis I concentrated on the specific conflict situation of a horse being introduced into a new social group, and investigated its behaviour and stress level. Horses were either introduced (1) immediately, (2) after an observation period, or (3) together with an integration horse after an observation period. Additionally, in the second part of my thesis I arranged several experiments to elaborate additional aspects which could affect the behaviour of horses during introductions. In this study I could describe a simplified method for measuring stress through the analysis of faecal GCMs in horses. An enzyme immunoassay (EIA) for 11-oxoaetiocholanolone using 11-oxoaetiocholanolone-17-CMO: BSA (3?,11-oxo-A EIA) as antigen showed high amounts of immunoreactive substances. The new assay increases the accuracy of the test and lowers the expenses per sample; also storing of samples at room temperature after collection is less critical. This is a big advantage both in the field of wildlife management of equids and in the field of equestrian sports (chapter 1). Comparing the different introduction techniques, the introduction with an integration horse led to significantly less total interactions and lower levels of aggression than the introduction of single horses, both immediately and after several days of observing the new group. Additionally, by observing the behaviour of the horses during everyday sociality I could develop a formula describing the interrelationship between expected aggression level and enclosure size per horse. The curve takes an exponential shape. Starting from a space allowance of 300 m2 and more per horse, the amount of aggressions per hour approaches zero. For the reduction of aggression levels and injury risks in socially kept horses I recommend an enclosure size of at least 300 m2 per horse (chapter 2). I further investigated the stress level of the introduced animals. Horses which were immediately introduced did not show elevated faecal GCMs. In contrast, horses which were introduced after an observation period had slightly elevated values 2 and 3 days after the introduction. For horses introduced together with an integration horse faecal GCMs were significantly above the baseline value on the day of introduction and 1 day after it. These differences between introduction techniques indicate that the introduction event itself is not as stressful as previously assumed. Rather standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses (chapter 3). In the commentary of chapter 4 several studies are discussed which failed to demonstrate social learning in horses. It is argued that they did not consider important aspects which could have an influence, such as the dominance status or the social background of the horses (chapter 4). In chapter 5 a social feeding situation was investigated. The social rank as well as the position of conspecifics affected the feeding strategy of horses. Domestic horses used social cognition and strategic decision making in order to decide where to feed. When possible they tended to return to the same, continuously supplied feeding site and switched to an ?avoidance tendency? in the presence of dominant horses or when another horse was already feeding there (chapter 5). One possibility to recognize group members is through olfactory recognition. In chapter 6 it is shown that horses are able to distinguish their own from their conspecifics? faeces. In addition, they paid most attention to the faeces of those group members from which they received the highest amount of aggressive behaviour (chapter 6). Horses show cognitive abilities because they are able to use humans as local enhancement cues when searching for food, independently of their body posture or gaze consistency when the persons face them. Moreover, they seem to orientate on the attention of familiar persons more than of unfamiliar persons (chapter 7). Altogether, the results of this thesis provide further support for the view that horses show good conflict resolution strategies. They are perfectly able to deal with the conflict situation of being introduced to new group members, and the introduction event itself is not as stressful as previously assumed. It is rather suggested that standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses. All additional experimental set-ups could demonstrate that horses are well capable of social cognition.
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