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Stashak, T. S., & Wissdorf, H. (2007). Adams' Lahmheit bei Pferden. Hannover: Schaper M. & H.
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Obergfell, J. (2012). Einf lus s v on St ruktur e l ement en auf da s Li eg ev e rha l t en v on Pf e rden in Gruppenha l tung unt e r Be rücks i cht igung de s Ag g r e s s i ons v e rha l t ens. Ph.D. thesis, , Karlsruhe.
Abstract: Durch die vorliegende Studie wurde der Einfluss von Strukturelementen auf das Liege- und Aggressionsverhalten von Pferden in Gruppenhaltung untersucht. Die Strukturelemente sollten Rückzugsmöglichkeiten bieten, sowie in ihrem Bereich zur Aufhebung der Individualdistanz führen und dadurch die Fläche relativ vergrößern. Für die Versuche stellte das Haupt- und Landesgestüt Marbach drei unabhängige Pferdegruppen mit verschiedener Herdengröße zur Verfügung, die jeweils in Einraum- Innenlaufställen gehalten wurden. Die Datenerfassung fand zwischen 23 und sieben Uhr statt. Insgesamt wurden 366 Stunden Videomaterial ausgewertet. Beim Ruheverhalten wurden mit Hilfe des event-sampling- Verfahrens die Parameter Gesamtliegedauer, Dauer in Seitenlage, Dauer der Einzelphasen in
Seitenlage und Abliegehäufigkeit pro Nacht erfasst. Es wurden Versuchsphasen ohne
Strukturelemente und mit hängenden Planen als Strukturierung durchgeführt. Im ersten Stall
kamen außerdem über einander gestapelte Strohballen zum Einsatz. Diese Art der
Strukturierung stellte sich jedoch als nicht praktikabel heraus und führte im Vergleich zu den Planen zu einer signifikanten Verschlechterung der Gesamtliegedauer. In Stall 1 konnte man eine tendenzielle Verbesserung der Parameter Gesamtliegedauer und Gesamtdauer in Seitenlage durch das Anbringen von Planen sehen. In Stall 3 dagegen verschlechterte sich das Ruheverhalten in den Versuchsphasen mit Planen gegenüber den Versuchsphasen ohne Struktur. Die Werte der Gesamtdauer in Seitenlage nahmen signifikant ab. In Stall 2 zeigten sich keine Unterschiede in den verschiedenen Versuchsphasen. Möglicherweise ist die Wirkung der Strukturelemente auf das Liegeverhalten abhängig von der Flächengröße. Stall 1 hatte bezogen auf die Leitlinien des BMELV die größte und Stall 3 die kleinste Fläche. Bei den anderen Parametern des Liegeverhaltens gab es keine signifikanten Unterschiede in den verschiedenen Versuchsphasen. Mit Hilfe des time-sampling-Verfahrens wurde die Anzahl gleichzeitig liegender Pferde und gleichzeitig liegender Pferde in Seitenlage bestimmt. Auch hier zeigten sich keine Unterschiede in den verschiedenen Versuchsphasen. Bei der Gegenüberstellung der Werte der Gesamtliegedauer und der Gesamtdauer in Seitenlage mit dem Alter der Pferde (Stall 1 und Stall 3) und mit dem Integrationszeitpunkt (Stall 1) konnte kein Zusammenhang festgestellt werden. Beim Aggressionsverhalten wurden mit Hilfe des
event-sampling-Verfahrens in den Ställen 2 und 3 verschiedene Arten von Aggressionen
erfasst, die dann in die drei Intensitätsgrade Low-Level-, Mid-Level- und High-Level-
Aggressionen unterteilt wurden. Neben der Anzahl wurde die Dauer der verschiedenen
Aggressionen bewertet. Insgesamt konnte eine positive Wirkung der Planen auf das
Aggressionsverhalten beobachtet werden. Die Gesamtanzahl an Aggressionen nahm in beiden
Ställen tendenziell in den Versuchsphasen mit Planen ab. In Stall 3 konnte, wenn man die
Aggressionen stundenweise betrachtet, ein signifikanter Unterschied festgestellt werden.
Auch der Hinterhandschlag und die Aggressionen, welche das Ruheverhalten stören,
verringerten sich tendenziell nach dem Anbringen von Strukturelementen. In beiden Ställen
nahm die relative Häufigkeit von Mid-Level-Aggressionen nach dem Anbringen von Planen
zu. Dagegen konnte bei den High-Level-Aggressionen und in Stall 3 bei den Low-Level-
Aggressionen eine relative Abnahme beobachtet werden. Sowohl die Anzahl als auch die
Dauer der Mid-Level-Aggressionen verringerten sich in Stall 3 stundenweise betrachtet
signifikant in den Versuchsphasen mit Planen. In Stall 2 war bei den High-Level-
Aggressionen sowohl bei der Dauer als auch bei der Anzahl eine signifikante Abnahme zu
sehen. Wenn man das Aggressionsverhalten in Bezug zu der Fläche in den zwei Ställen
betrachtet, schien diese vor allem einen Einfluss auf die High-Level-Aggressionen zu
nehmen. Mit Hilfe des Rangindex der Pferde im Stall 3 wurde eine Rangordnung aufgestellt.
Zwischen dem Platz der Pferde in der Rangordnung und den Parametern des Ruheverhaltens
(Gesamtliegedauer, Gesamtdauer in Seitenlage) sowie dem Alter der Pferde konnte kein
Zusammenhang festgestellt werden. Zwischen dem Rangindex und der Gesamtanzahl an
Aggressionen bestand dagegen ein hoch signifikanter Zusammenhang. Im Rahmen dieser
Studie ist das Anbringen von Strukturelementen in Bezug auf das Aggressionsverhalten in
Einraum-Innenlaufställen von Pferden zu empfehlen. Die Wirkung auf das Liegeverhalten der
Pferde sollte in weiteren Studien untersucht werden.
[In this study the influence of structural elements on lying and aggression behavior in horses
kept in groups was exanimated. Structural elements should offer the possibility of retreat.
Furthermore, within the respective area, the individual distance should be limited, thus
increasing the overall space relatively. For the studies the “Haupt- und Landesgestüt
Marbach” provided three non-related groups of different sized horses, each kept in separate
“Einraum-Innenlaufställen”. Data collection took place between 11 pm and 7 am. Altogether,
366 hours of video material were analyzed. Using the event-sampling-method, the following
parameters within the horses´ resting behavior were determined: The total lying period, the
total lying period in lateral position, the periods of the single lying in lateral position and the
frequency of lying down. Test phases were carried out without structural elements and with
hanging canvases as structural elements. In the first stable there were also three bales of straw
piled up on top of each other. This kind of structuring emerged as impractical and in
comparison to the canvas, led to a significant deterioration to the total period of lying. After
the canvases were fixed in the first stable, a slight improvement with the parameters of total
lying period and total lying period in the lateral position could be seen. In contrast, the resting
behavior in stable 3 worsened in the test phases with the canvases in comparison to the test phases without structural elements. The values for the total lying period in the lateral position
decreased significantly. In stable 2, there were no differences in the different test phases.
Possibly the effect of the structural elements is dependent of the space. Stable 1 had the
largest space and stable 3 the least space referred to the guideline of the BMELV. The other
parameters of the lying behavior showed no significant differences in the different test phases.
The number of horses lying at the same time, and the number of horses lying at the same time
in the lateral position were detected with the aid of the time-sampling-method. There were
also no differences between the different test phases. If the values of the total lying period and
the total lying period in lateral position were compared to the age of the horses (stable 1 and
stable 3) and to the point of integration (stable 1), there was no correlation.
Different kinds of aggression were detected by the means of the event-sampling-method in
stable 2 and 3 and were subdivided into the three levels of intensity: Low-level-, mid-level
and high-level-aggressions. Both the number and the duration of the different aggressions
were evaluated. All in all a positive influence of the canvas on the aggression behavior could
be observed. In both stables, the total number of aggressions decreased in the test phases with
canvases. In stable 3 a significant difference could be detected, if the aggressions were
observed per hour. Also hind limb kicks and aggressions disturbing the resting behavior,
decreased by trend, after fixing the canvases in the stable. In both stables the relative
frequency of mid-level-aggressions increased after the canvases were mounted. In contrast
there was a relative decrease at the high-level-aggressions and in stable 3 at the low-levelaggressions.
The number and the duration of the mid-level-aggressions in stable 3 decreased,
when considered by the hour in the test phases with canvases. In stable 2 a significant
decrease in duration and number of high-level-aggressions was observed. When examining
aggression behavior in reference to space in the two stables, high-level-aggressions seemed to
be mostly influenced. In stable 3 a rank order was established with the aid of the rank index of
the horses. There was no correlation between the horses´ rank and the parameters of the
resting behavior (the total lying period, the total lying period in lateral position) and between
horses´ rank and age. In contrast, there was a high significant correlation between the rank
index and the total number of aggressions. In context to this study, structural elements in
“Einraum-Innenlaufställen” (BMELV 2009) seem to have a positive influence on the
aggression behavior and thus seem to be recommendable. The influence on the lying behavior
should be investigated in further studies.]
Keywords: Pferde, Ruheverhalten, Liegeverhalten, Aggressionsverhalten, Strukturelemente, Rangordnung [horses, rest behavior, lying behavior, aggression behavior, structural elements, rank order]
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Schneider, G., & Krueger, K. (2012). Third-party interventions keep social partners from exchanging affiliative interactions with others. Anim. Behav., 83(2), 377–387.
Abstract: Third-party interventions are defined as the interruption of dyadic interactions by third animals through direct physical contact, interposing or threats. Previous studies focused on the analysis of interventions against agonistic encounters. However, there have been no evaluations of interventions against affiliative behaviours, particularly in relation to the intervening animal�s social relationships and its social and spatial position. Horses, Equus caballus, are an interesting model species, as interventions against affiliative interactions occur more frequently than against agonistic interactions. In this study, 64 feral horses displayed 67 interventions in affiliative interactions and eight interventions in agonistic interactions within the observation period. We analysed the interventions in affiliative encounters, and found that it was mainly higher-ranking females that intervened in the affiliative interactions of group mates in the stable horse harems. The intervening animals took an active part in affiliative and agonistic encounters within the group, but did not occupy particular social roles or spatial positions. They intervened in affiliative interactions in which group mates with which they had social bonds interacted with other members of the group. They targeted the nonbonded animal and approached the one with which they were socially bonded. We suggest some species use third-party interventions in affiliative interactions to prevent competition for preferred social interaction partners from escalating into more costly agonistic encounters.
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Neumann, C., Duboscq, J., Dubuc, C., Ginting, A., Irwan, A. M., Agil, M., et al. (2011). Assessing dominance hierarchies: validation and advantages of progressive evaluation with Elo-rating. Animal Behaviour, 82(4), 911–921.
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Bang, A., Deshpande, S., Sumana, A., & Gadagkar, R. (2010). Choosing an appropriate index to construct dominance hierarchies in animal societies: a comparison of three indices. Animal Behaviour, 79(3), 631–636.
Abstract: A plethora of indices have been proposed and used to construct dominance hierarchies in a variety of vertebrate and invertebrate societies, although the rationale for choosing a particular index for a particular species is seldom explained. In this study, we analysed and compared three such indices, viz Clutton-Brock et al.'s index (CBI), originally developed for red deer, Cervus elaphus, David's score (DS) originally proposed by the statistician H. A. David and the frequency-based index of dominance (FDI) developed and routinely used by our group for the primitively eusocial wasps Ropalidia marginata and Ropalidia cyathiformis. Dominance ranks attributed by all three indices were strongly and positively correlated for both natural data sets from the wasp colonies and for artificial data sets generated for the purpose. However, the indices differed in their ability to yield unique (untied) ranks in the natural data sets. This appears to be caused by the presence of noninteracting individuals and reversals in the direction of dominance in some of the pairs in the natural data sets. This was confirmed by creating additional artificial data sets with noninteracting individuals and with reversals. Based on the criterion of yielding the largest proportion of unique ranks, we found that FDI is best suited for societies such as the wasps belonging to Ropalidia, DS is best suited for societies with reversals and CBI remains a suitable index for societies such as red deer in which multiple interactions are uncommon.
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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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Smith, J. E., Kolowski, J. M., Graham, K. E., Dawes, S. E., & Holekamp, K. E. (2008). Social and ecological determinants of fission-fusion dynamics in the spotted hyaena. Anim. Behav., 76(3), 619–636.
Abstract: Theory predicts that individuals living in fission-fusion societies, in which group members frequently change subgroups, should modify grouping patterns in response to varying social and environmental conditions. Spotted hyaenas, Crocuta crocuta, are long-lived carnivores that reside in permanent social groups called clans. Clans are complex, fission-fusion societies in which individual members travel, rest and forage in subgroups that frequently change composition. We studied two clans in Kenya to provide the first detailed description of fission-fusion dynamics in this species. Because social and ecological circumstances can influence the cohesiveness of animal societies, we evaluated the extent to which specific circumstances promote the formation of subgroups of various sizes. We found that cooperative defence of shared resources during interclan competition and protection from lions were cohesive forces that promoted formation of large subgroups. We also tested hypotheses suggesting factors limiting subgroup size. Mothers with small cubs avoided conspecifics, thereby reducing infanticide risk. Victims of aggression either reconciled fights or separated from former opponents to reduce the immediate costs of escalated aggression in the absence of food. As predicted by the ecological constraints hypothesis, hyaenas adjusted their grouping patterns over both short and long time scales in response to feeding competition. Crocuta were most gregarious during periods of abundant prey, joined clanmates at ephemeral kills in numbers that correlated with the energetic value of the prey and gained the most energy when foraging alone because cooperative hunting attracted numerous competitors. Overall, our findings indicate that resource limitation constrains grouping in this species.
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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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Heitor, F., & Vicente, L. (2008). Maternal care and foal social relationships in a herd of Sorraia horses: Influence of maternal rank and experience. Appl. Anim. Behav. Sci., 113(1-3), 189–205.
Abstract: The influence of maternal rank and experience on patterns of maternal care and social relationships of foals were investigated in a managed herd of Sorraia horses, Equus caballus. Social interactions and spatial relationships of 13 foals (seven females and six males) born to seven mares were examined from birth to 10 months of life, within the three major periods of foal development. Conflict over suckling between dam and foal was not generally affected by rank and experience, but higher-ranking mothers allowed more suckling during late lactation than lower-ranking mothers. Foals of higher-ranking mares spent more time in proximity to the mother during socialization. Maternal rank and experience did not significantly affect maternal protectiveness, foal independence from the mother or the development of affiliative relationships between foals and group members. Foals of higher-ranking mares received lower frequencies of aggression from other horses only in the first month of life. Dominance relationships among foals depended mainly on aggressiveness and were not associated with maternal rank. The large variability in maternal behaviour, the absence of a significant association between maternal rank and body condition at parturition and the stable social environment within this herd may partly account for the reported results.
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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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