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Stashak, T. S., & Wissdorf, H. (2007). Adams' Lahmheit bei Pferden. Hannover: Schaper M. & H.
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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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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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Taillon, J., & Côté, S. (2008). Are faecal hormone levels linked to winter progression, diet quality and social rank in young ungulates ? An experiment with white-tailed deer ( Odocoileus virginianus ) fawns. Behav. Ecol. Sociobiol., 62(10), 675–677.
Abstract: Abstract Hormones play a central role in the physiology and behaviour of animals. The recent development of noninvasive techniques has increased information on physical and social states of individuals through hormone measurements. The relationships among hormones, life history traits and behaviours are, however, still poorly known. For the first time, we evaluated natural winter glucocorticoid and testosterone levels in young ungulates in relation to winter progression, diet quality and social rank. Overwinter, levels of glucocorticoid and testosterone decreased, possibly due to the decline of fawns" body mass. The relationships between hormone levels and diet quality were surprising: Fawns fed the control diet presented higher glucocorticoid and lower testosterone levels then fawns fed the poor diet, suggesting that control fawns faced a higher nutritional stress than those on the poor diet. Similarly to other studies on social mammals, we found no relationship between faecal glucocorticoid levels and social rank, suggesting that social stress was similar for dominant and subordinate fawns during winter. Testosterone levels were not correlated to social rank as found previously in groups of individuals forming stable social hierarchies and maintaining stable dominance relationships. The simultaneous suppression of glucocorticoid and testosterone levels suggests for the first time that young ungulates present a hormonal strategy to prevent fast depletion of limited proteins and fat resources during winter.
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Krueger, K., Flauger, B., Farmer, K., & Hemelrijk, C. (2014). Movement initiation in groups of feral horses. Behav. Process., 103, 91–101.
Abstract: Abstract Herds of ungulates, flocks of birds, swarms of insects and schools of fish move in coordinated groups. Computer models show that only one or very few animals are needed to initiate and direct movement. To investigate initiation mechanisms further, we studied two ways in which movement can be initiated in feral horses: herding, and departure from the group. We examined traits affecting the likelihood of a horse initiating movement i.e. social rank, affiliative relationships, spatial position, and social network. We also investigated whether group members join a movement in dominance rank order. Our results show that whereas herding is exclusive to alpha males, any group member may initiate movement by departure. Social bonds, the number of animals interacted with, and the spatial position were not significantly associated with movement initiation. We did not find movement initiation by departure to be exclusive to any type of individual. Instead we find evidence for a limited form of distributed leadership, with higher ranking animals being followed more often.
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Krueger, K., Schneider, G., Flauger, B., & Heinze, J. (2015). Context-dependent third-party intervention in agonistic encounters of male Przewalski horses. Behav. Process., 121, 54–62.
Abstract: Abstract One mechanism to resolve conflict among group members is third party intervention, for which several functions, such as kin protection, alliance formation, and the promotion of group cohesion have been proposed. Still, empirical research on the function of intervention behaviour is rare. We studied 40 cases of intervention behaviour in a field study on 13 semi-wild bachelor horses (Equus ferus przewalskii) in (a) standard social situations, and (b) when new horses joined the group (i.e. introductions). Only interventions in agonistic encounters were analysed. Eight of 13 animals directed intervention behaviour toward threatening animal in agonistic encounters of group members. One stallion was particularly active. The stallions did not intervene to support former group mates or kin and interventions were not reciprocated. In introduction situations and in standard social situations, the interveners supported animals which were lower in rank, but targeted, threatening animals of comparable social rank. After introductions, stallions received more affiliative behaviour from animals they supported and thus appeared to intervene for alliance formation. In standard social situations, interveners did not receive more affiliative behaviour from animals they supported and may primarily have intervened to promote group cohesion and to reduce social disruption within the group.
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Heitor, F., & Vicente, L. (2010). Dominance relationships and patterns of aggression in a bachelor group of Sorraia horses (Equus caballus). J. Ethol., 28(1), 35–44.
Abstract: Abstract The influence of individual factors on dominance rank and the relationship between rank distance and patterns of aggression predicted by models of evolutionarily stable strategies (ESS) of animal conflict were investigated in a managed bachelor group of Sorraia horses, Equus caballus. The group was composed of four to six stallions 3- to 12-years-old during the study period. The dominance hierarchy was significantly linear and rank was not related to age, weight, height or aggressiveness. Frequency and intensity of agonistic interactions were low, but higher-ranking stallions did not receive lower aggressiveness than lower-ranking stallions. There was some evidence that dominance relationships were more contested among close-ranking stallions, as predicted. Agonistic-related interactions among close-ranking stallions served similar functions to those among distant-ranking stallions, but the latter interacted more frequently than expected for access to resting sites and/or resting partners. Therefore, we found some evidence that agonistic-related interactions among distant-ranking stallions play a larger role in providing access to valuable and defendable resources than those among close-ranking stallions. Nevertheless, the fact that space to escape from aggression was limited and breeding access was independent from dominance rank may have reduced the benefits relative to costs of aggression and therefore limited the occurrence of contests over dominance and resources.
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Vervaecke, H., Vries, H. D., & Elsacker, L. V. (1999). An Experimental Evaluation Of The Consistency Of Competitive Ability And Agonistic Dominance In Different Social Contexts In Captive Bonobos. Behaviour, 136(4), 423–442.
Abstract: Bonobos have been described as a relatively egalitarian and female dominant species. The exact nature and quality of their dominance relationships and the existence of female dominance are current topics of dispute. We investigated the consistency across social contexts, the stability in time, and the degree of expression of the competitive feeding ability and agonistic dominance in a captive group of bonobos. First, we examined whether the competitive feeding ranks and agonistic ranks differed in different dyadic contexts, triadic contexts and the whole group context. For some pairs of animals the dominance relationships with respect to competitive feeding altered with different group compositions. The agonistic dominance relationships changed accordingly. The competitive feeding ranks and agonistic ranks in the experiments correlated strongly with each other. The alpha position was occupied by a female, but not all females outranked all males. We suggest that females can profit from each others presence to gain inter-sexual dominance. Second, although the agonistic rank order in the whole group remained the same over at least five years, some dyadic competitive feeding ranks changed over time, resulting in a stronger female intersexual dominance. Third, the degree of expression of the behaviors used to quantify dyadic competitive and agonistic dominance was not high, in line with the popular 'egalitarian' epithet. Notwithstanding its low consistency across contexts, the dominance hierarchy in the whole group has a strong predictive value for other social relationships such as grooming. Given this strong effect of rank on other behaviours and given the strong dependency of rank on social context, the choice of the right party members may be a crucial factor in the fission-fusion processes of free-ranging bonobos.
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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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