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Author Wasilewski A.
Title 'Friendship' in ungulates? – Sociopositive relationships between non-related herd members of the same species Type Manuscript
Year 2003 Publication Abbreviated Journal
Volume Issue Pages
Keywords
Abstract http://www.staff.uni-marburg.de/~z-phylog/wisstaff/wasi-sum.htm

Summary

* The quantitative study presented here demonstrates and verifies for ungulates the existence of social bonds, which in humans are termed 'friendships'. Whereas 'friendships' in non-human primates have become increasingly accepted over the past 30 years, their existence in non-primates remained as yet to be proved.

* The investigation was conducted on ungulates, because like primates the majority of their species are highly social; at the same time they are phylogenetically distant to the primates. To date, research into the social behaviour of ungulates in general, and of domestic species in particular, focuses predominantly on agonistic (socionegative) components, such as conflicts and resulting dominance hierarchies; sociopositive behaviours have been largely neglected.

As a consequence, previous evidence in favour of 'friendships' among ungulates remained largely anecdotal. Studies explicitly investigating sociopositive relationships among herd members are rare and typically confound bonds based on 'friendships' with reproductive relationships (kinship and sexually motivated bonds).

* Of each of the two large ungulate orders two species were investigated: horses and donkeys (Perissodactyla, Equidae) and sheep and cattle (Artiodactyla, Bovidae). The comparative, individual-based (and hence highly discriminative) longitudinal approach addressed all three levels of social organisation: associations, interactions and bond patterns within social units. Due to its systematic, quantitative, individual-based and comparatively broad design, this work may be considered to some degree a 'pioneering study'. As a consequence, a substantial amount of time and effort was required, particularly with respect to developing methods for the collection and analysis of data.

* Two reasons led to conducting this study on domestic ungulates: For farm animals information about who is related to whom is usually readily available, facilitating discrimination between friendship and kinship. Furthermore, successful proof of 'friendships' among ungulates would imply practical consequences for animal husbandry, since a social environment accommodating the animals' psychosocial needs is an essential prerequisite for their well-being (ethical reasons for animal protection) and ultimately results in higher productivity and hence economic efficiency (anthropocentric reasons for animal protection).

* Field research was conducted in England. Cattle and sheep were studied under commercial husbandry conditions, whereas appropriately large herds of horses and donkeys could only be found at a sanctuary.

* Two to three herds of each of the four species, namely cattle, sheep, donkeys and horses, comprising 11 to 60 members (average herd size approx. 25) were studied in two consecutive years (1996: 36 weeks, 1997: 28 weeks). The herds of cattle mainly consisted of subadult females, whereas those of the other species were predominantly composed of adults. Equids were kept in mixed-sex herds of mares and geldings (castrated males); the bachelor flocks of sheep consisted of 'intact' males (rams). The data presented here are based on 234 individuals and approximately 1500 hours of observation.

* Since even human social psychology provides neither a consistent definition nor a universally applicable terminology, it was necessary to establish a generally valid, precise definition of 'friendship'. Care was taken to exclude characteristics containing a bias as to the function of friendships (criteria were limited to those regarding contents and form). The definition contains all relevant criteria including those necessary for discrimination against alternative sociopositive relationships.



Friendship is defined as a voluntary, reciprocal, non-reproductive bond between individuals. It is primarily dyadic and carries a subjective value to both participants. Friendship is characterised by positive affect ('affection') and is expressed in a consistent inter-individual preference.

This definition is applicable to social relationships among both human and non-human animals, as well as interspecific relationships.

* Two types of parameters were used as indicators of 'friendship': frequency of spatial proximity (nearest neighbours; consistent neighbour preferences as a reflection of associations) and frequencies of different spontaneous as well as experimentally induced sociopositive interactions. For different reasons, of the four kinds of interactions (social grooming s.l., resting in physical contact, sharing limited feed, documenting behaviours s.l.) only grooming and feed sharing proved reliable.

* Aspects of the animals' spontaneous behaviour were recorded for all herds and in both years, whereas husbandry conditions demanded that complementary experiments be restricted to a few selected herds only. Data were collected using different Sampling and Recording Techniques, which were usually combined for maximum efficiency (e.g. Scan Sampling with nearest neighbour data, Ad libitum Sampling and Continuous Recording with sociopositive interactions).

* Data were subjected to an extensive statistical analysis. In addition to commonly used uni- and bivariate methods (Binomial and Chi-Square Goodness-of-Fit Tests, bivariate Correlations and Mantel Tests) a number of multivariate statistics were applied (Multidimensional Scaling, hierarchical Cluster Analysis, partial Correlations and multiple Mantel-Tests). Dyadic data are best represented as matrices; matrix data, however, pose the difficulty of partial interdependence, requiring Randomisation Tests (such as the one developed by Mantel) for their analysis. The complexity and amount of neighbour data required the development and optimisation of a 'customised' evaluation software (NENESYS = NEarest NEighbour SYstematic Standardisation programme) in close interdisciplinary cooperation with computer scientists and mathematicians.

* Seven different aspects of 'ungulate friendships' were addressed. The presented results differ in extent and depth.

1. Demonstration and verification of friendships in ungulates

2. Quantification of inter-individual preferences

3. Situation-specific aspects of these preferences

4. Dynamics and duration of friendships

5. Asymmetry within the relationships

6. Conditions enhancing friendships

7. Functions of animal friendships

* All four species expressed distinct inter-individual ('personal') preferences both for certain neighbours and for specific partners for sociopositive interactions. Endeavouring to present conservative and hence reliable assessments, only those neighbour preferences exceeding a threshold criterion were considered as associations. As the set-up of the study ensured that these associations met the criteria outlined in the definition of 'friendship' by excluding alternative bases for bonds (such as kinship and sexual motivation = reproductive relationships), the first explicit attempt at verifying the existence of friendship can be considered successful for all four species of ungulates examined. As a consequence, the concept of 'friendship' needs to be extended beyond the order of primates.

* Due to methodological reasons, the frequencies of interactions are only semi-quantitative and hence can be compared within herds only.

Frequencies of spatial proximity, on the other hand, provide a fully quantitative indicator, enabling direct comparisons across species, between different herds of the same species and between individuals. Due to the 'spatial limitations' of this thesis, it mainly focuses on inter-species comparisons.

With the exception of donkeys, the species could be arranged in sequence of increasing values of three parameters characterising their associative structure: maximum strength of association, maximum group size and proportion of herd members associated in distinct groups. Cattle showed the lowest values in all three parameters, sheep were intermediate, and horses reached the highest values. In these three species the strength of their associations was positively related to group size and the proportion of individuals associated in distinct groups. The donkeys did not fit this pattern: On the one hand, their association pattern comprised the lowest proportion of associated members and the smallest groups (pairs); on the other hand, these pairs exhibited very strong associations.

* Comparing inter-individual preferences across up to four different situations clear interspecific differences became evident. Some of these were greater between species of the same order than between orders.

The comparison of structures of association during grazing to those during resting, revealed two distinct patterns: Herds of horses and sheep flocks divided themselves into separate, usually mutually exclusive and independent groups and their associations corresponded greatly in both situations. Donkeys and cattle, on the other hand, formed diffuse, incomplete associative networks. The proportion of associations corresponding in both situations was far lower than in horses and sheep.

In all four species frequencies of social grooming s.l. (sheep are generally believed not to engage in social grooming, but see below) were compared to proximity frequencies during grazing and resting; and additionally to frequencies of feed sharing in the equid species.

Whereas horses restricted their sociopositive interactions entirely (grooming) or mostly (feed sharing) to members of their own group, cattle showed very little if any correspondence between partner preferences when grazing, resting and grooming. Donkeys spread their preferences widely across herd members, 'typically' approaching individuals for grooming with whom they were not associated during either grazing or resting. Considering their patterns of feed sharing, two different types of behaviour became evident: Pairs of donkeys that were closely associated during both grazing and resting, preferentially shared feed between them, whereas the remaining herd members did not discriminate between associates and non-associates when sharing feed. For sheep the findings of this study substantially increased the likelihood that unidirectional head rubbing and reciprocal cheek-to-cheek contact serve to strengthen 'personal' bonds and reduce tension – two functions generally attributed to social grooming. Both behaviours occurred mostly, but not exclusively, within groups of associated animals; these associations corresponded largely during grazing and resting.

* Results indicate that different types of social interactions and associations during different activities, respectively, may be particularly suited to initiate, consolidate and intensify friendships and occur particularly frequently during different phases of friendship formation and development. In horses, for instance, social grooming appears to some extent to be associated with the initiation of bonds, whereas resting in close proximity seems to strengthen newly formed relationships.

The longitudinal character of the study permitted to trace existing bonds over a maximum of 18 months. The duration of bonds in horses and sheep was comparable to findings of other authors for herds whose social networks resulted from a mixture of bonds based on both friendship and kinship; cattle relationships lasted shorter than described in the literature, while those in donkeys turned out to be approximately twice as long as had been documented to date.

* Aspects of asymmetry within dyadic relationships become particularly evident in grooming behaviours in bovids, which are typically performed unilaterally (social licking in cattle and head rubbing in sheep). Both behaviours can occur either spontaneously or as a consequence of solicitation.

* Addressing the question of factors potentially encouraging friendships, relationships between the degree of similarity with respect to some physical characteristics and nearest neighbour frequencies were found to be statistically significant. Cattle of similar age associated more closely, and horned rams preferred other individuals with horns as neighbours. The widely accepted preference of horses for associates of similar coat colour (and height) was not verified.

It remains to be decided whether (in addition to this preference for similar individuals) a preference based on complementary characteristics does exist.

* Following a critical review of the methodology employed, the benefits ungulates derive from friendships and possible applications for animal husbandry are discussed. Very little convincing evidence of a (direct) concrete, practical benefit was found. A psychological benefit in the form of social / emotional support, on the opposite, became highly probable.

Anecdotal evidence indicates, that situations causing a state of anxiety, may favour the initiation and intensification of friendships. Moreover, the study was able to statistically prove connection between (roughly) simultaneous introduction to a new herd (novel social environment) and persisting associations (as an indicator of friendships) in horses and sheep. These findings suggest that ungulate friendships provide a feeling of security. Social / emotional support among friends is predominantly given in a passive manner, e.g. in the form of – both literally and metaphorically – standing by a friend during conflicts with a third party, followed by social grooming with the friend. Social grooming has been shown by other authors to reduce physiological symptoms of stress. In addition, external signs of relaxation indicate a decrease in psychological tension. Hence, psychological support enhances the animal's physical health (and ultimately increases its biological fitness), providing an indirect practical benefit.

* Since all larger species of farm animals are highly social ungulates, the demonstration and verification of friendships implies consequences for husbandry practice. Intensive husbandry conditions are characterised by high animal densities, resulting in increased social stress, and grouping of animals of similar ages, which interferes with or prevents the formation and maintenance of bonds between kin. As friendships develop preferentially between individuals of similar age, they can be accommodated with relatively little effort when deciding about regrouping. The study presented here provides explicit recommendations to interested stockpersons on how to expeditiously and efficiently identify existing friendships and on which aspects to take into account when regrouping established herds or introducing new members.

* The different aspects of the study and their complex and often interwoven interrelations are condensed into a 'Regelkreismodell'. This model facilitates the generation of precise hypotheses for future research. Further detailed investigation into a variety of questions raised by this study seems in many ways promising and rewarding.

* The successful demonstration and verification of the existence of friendship in ungulates requires an extension of the 'concept of friendship' beyond the order of primates and encourages more detailed research into the bonding behaviour of other social vertebrates (e.g. elephants, dolphins, carnivores and also parrots).

* This extension demands a reinterpretation of some behaviours which so far have been unsatisfactorily or controversially classified, such as social head or horn rubbing in sheep, which has been mentioned occasionally, but received very little attention in the literature. Due to its occurrence during aggressive interactions, some authors consider horn or head rubbing to be agonistic. Others suppose the preorbital gland to play a significant role in this type of interaction and interpret head or horn rubbing as a 'respectful gesture' (adopting the dominant sheep's individual scent).

The study presents the first description of cheek-to-cheek contact in sheep. These cheek contacts are unequivocally sociopositive in nature. Since they are frequently associated with head rubbing, a third interpretation of head rubbing behaviour is offered by classifying both behaviours within the context of social grooming s.l.. More detailed examinations of these behaviours will be conducted as part of a research project into the evolution of the bovidae.
Address
Corporate Author Thesis Ph.D. thesis
Publisher University of Marburg Place of Publication Marburg Editor
Language German Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
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Notes Approved no
Call Number refbase @ user @ Serial 824
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Author Rybarczyk, P.; Rushen, J.; de Passille, A.M.
Title Recognition of people by dairy calves using colour of clothing Type Journal Article
Year 2003 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.
Volume 81 Issue 4 Pages 307-319
Keywords Cattle handling; Discrimination; Human-animal relationships; Colour cues; Y-maze
Abstract We examined whether very young dairy calves are able to discriminate between two people, and whether they use the colour of clothing or other indices to do so. During the familiarisation phase, one person (the familiar rewarder), who always wore the same colour clothes, gave milk, spoke gently and patted the calves in their individual pen for 6 days each week. During the test phase, the calf had to make a choice in an Y-maze placed in front of the gate of its stall. When the calves chose the familiar rewarder, they received 200 ml of milk as reinforcement. When they made the incorrect choice, they received nothing and were returned to their stall. On each test day, the calves made eight choices. The criterion of success was that the calf made at least six correct choices in eight trials on each of two consecutive test days (P<0.021 by the binomial law). The first experiment was carried out with fourteen 1-week-old male and female Holstein calves to see if calves could approach a person, who changed position in the maze, in order to obtain a feed reward. The familiar rewarder wore the same clothes as during the period of familiarisation and was in one arm of the Y-maze. The other arm was empty and the position of the familiar rewarder in the maze was randomised. Eleven of the 14 calves reached the criterion for success, after only three tests. The second experiment, carried out with five 2-week-old calves, examined whether the calves can differentiate the familiar rewarder (wearing the same clothing as during the period of familiarisation) from another person (the non-rewarder) wearing clothes of a different colour. The criterion of success was reached by all five calves. The third experiment was carried out with seven 2-week-old calves. It examined whether the calves can differentiate the familiar rewarder and the non-rewarder, when the two people are wearing clothes of the same colour (i.e. the same colour worn by the familiar rewarder during the phase of familiarisation). None of the calves were able to reach the criterion of success within a limited number of four test days. Often, calves would always choose the same arm of the maze. The fourth experiment was carried out on six 1-month-old calves. It was similar to experiment 3 with the difference that the familiar rewarder and the non-rewarder both wore the same colour clothes, but which were not the same colour as worn during the phase of familiarisation. Only one calf achieved the criterion of success within two test days. Results demonstrated that colour cues help very young calves to discriminate between two people, when these people wear different colour clothing. Some calves may be able to use other indicators than the colour of clothing. The Y-maze method is an promising way of examining calves' abilities to recognise people.
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Call Number refbase @ user @ Serial 848
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Author Chase, I.D.; Tovey, C.; Murch, P.
Title Two's Company, Three's a Crowd: Differences in Dominance Relationships in Isolated Versus Socially Embedded Pairs of Fish Type Journal Article
Year 2003 Publication Behaviour Abbreviated Journal Behaviour
Volume 140 Issue 10 Pages 1193-1217
Keywords
Abstract We performed experiments with cichlid fish to test whether several basic aspects of dominance were the same in isolated pairs as in pairs within a social group of three or four. We found that the social context, whether a pair was isolated or within a group, strongly affected the basic properties of dominance relationships. In particular, the stability of relationships over time, the replication of relationships in successive meetings, and the extent of the loser effect were all significantly less in socially embedded pairs than in isolated pairs. We found no significant winner effect in either isolated or socially embedded pairs. These findings call into question many current approaches to dominance that do not consider social context as an important factor in dominance behavior. These findings also cast serious doubt on the validity of empirical and theoretical approaches based on dyadic interactions. Among these approaches are game theoretic models for the evolution of aggressive behavior, experimental designs evaluating how asymmetries in attributes influence the outcome of dominance
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Call Number refbase @ user @ Serial 857
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Author Huebener, E. (ed)
Title Fortschritt des Wissens zu Grundsitz und Schenkelhilfen und die “Entdeckung” der Bewegungen des Pferderückens und des Pferderumpfes Type Manuscript
Year 2003 Publication www.reiten-lesen-denken.de Abbreviated Journal
Volume Issue Pages 6
Keywords Grundsitz – Stehsitz – Unterschenkel-Vorstreck-Sitz – Balancesitz – Erkenntnisse seit Erfindung des Buchdrucks – vom 16. bis 20.Jh.
Abstract Tabellarische, chronologische übersicht zur Entwicklung des Grundsitzes durch fünf Jahrhunderte, wobei “Grundsitz” als Oberbegriff für verschiedene Formen des wirklich sitzenden “Hoch zu Roß” dient. Fehlentwicklungen (anstatt von Fortschritten, nur in Ausnahmefällen erwähnt)sind fett kursiv als solche gekennzeichnet.
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Corporate Author Thesis
Publisher Place of Publication Editor Huebener, E.
Language Deutsch Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
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Notes Approved no
Call Number Reiten Lesen Denken @ eberhardhuebener @ Serial 1762
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Author Huebener, E.
Title Basis Seat and Leg Aids – Evolution of Knowledge Type Manuscript
Year 2003 Publication www.reiten-lesen-denken.de Abbreviated Journal
Volume Issue Pages 6
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Abstract
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Language Deutsch Summary Language Original Title
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Call Number Reiten Lesen Denken @ eberhardhuebener @ Serial 1763
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Author Altmann, S.A.; Altmann, J.
Title The transformation of behaviour field studies Type Journal Article
Year 2003 Publication Animal Behaviour. Abbreviated Journal Anim. Behav.
Volume 65 Issue 3 Pages 413-423
Keywords
Abstract As areas of science mature, they pass through three, broadly overlapping stages of development, characterized respectively by description, explanation and synthesis. Field research on animal behaviour is making the transition from an area with a preponderance of purely descriptive studies to one that also includes the development and testing of verifiable hypotheses about the structure, causes and consequences of behaviour. We survey several reasons for this transformation of behaviour field studies and some of the major trends that characterize it, including: (1) patterns discerned in our cumulative knowledge of natural history; (2) increased support for behaviour field studies; (3) interfaces with related areas of science; (4) the development of observational sampling methods and other aspects of data sampling and analysis; (5) the development of models of behaviour's adaptive functions and life-history consequences; (6) long-term field sites that make possible complete life histories, increased attention to individual differences and intergenerational studies of behaviour; and (7) the development of techniques for remote tracking of animals and for noninvasive, hands-off sampling of a range of behavioural, physiological, genetic and environmental phenomena. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.
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Notes Approved no
Call Number Serial 1800
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Author Momozawa, Y.; Ono, T.; Sato, F.; Kikusui, T.; Takeuchi, Y.; Mori, Y.; Kusunose, R.
Title Assessment of equine temperament by a questionnaire survey to caretakers and evaluation of its reliability by simultaneous behavior test Type Journal Article
Year 2003 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.
Volume 84 Issue 2 Pages 127-138
Keywords Anxiety; Heart rate; Horse; Questionnaire; Temperament
Abstract We carried out a questionnaire survey of the caretakers, using 86 riding horses kept in the Equestrian Park, Tokyo (Japan Racing Association). The questionnaire survey used a 5-point scale and a 3-point scale to assess several caretakers' impressions of each horse's temperament, on the basis of the norm and the horse's tendencies in ordinary care and daily training. Factor analysis of the temperament scores obtained with the 5-point scale questionnaire revealed three mutually independent factors that we named “anxiety”, “novelty seeking” and “understanding”. In order to verify the reliability of this questionnaire survey, a balloon reactivity test was conducted using the same horses. Each horse was introduced into an unfamiliar indoor arena (7 mx12.5 mx3 m) in the center of which two balloons slowly revolved. The horses' responses were assessed by recording changes in their behavior and heart rate (HR) during the 5 min experimental period. By comparing the questionnaire survey and the balloon reactivity test, it was found that the horses evaluated as highly anxious by the caretakers tended to show greater HR increases and defecate more often during exposure to the balloon stimuli than did the other horses. Additionally, the horses assessed by caretakers to have problems with ordinary care and/or training showed greater increases of HR and frequency of defecation in the balloon reactivity test, and the horses assessed as having `a long adaptation time to unfamiliar objects' were found to be unwilling to touch the balloons. Thus, the horses' behavior during the balloon reactivity test was highly consistent with their temperament as determined by the questionnaire. These results suggest that the questionnaire survey would be an effective means to assess equine temperamental traits, especially those related to anxiety.
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Language Summary Language Original Title
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Call Number refbase @ user @ Serial 1984
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Author Visser, E.K.; van Reenen, C.G.; Schilder, M.B.H.; Barneveld, A.; Blokhuis, H.J.
Title Learning performances in young horses using two different learning tests Type Journal Article
Year 2003 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.
Volume 80 Issue Pages 311-326
Keywords Horse; Personality; Learning performance; Consistency; Emotionality
Abstract To achieve optimal performance in equine sports as well as in leisure not only the physical abilities of the horse should be considered, but also the horse's personality. Besides temperamental aspects, like emotionality, or the horse's reactivity towards humans in handling situations, the learning ability of the horse is another relevant personality trait. To study whether differences in learning performance are consistent over time and whether individual learning performance differs between learning tests or is affected by emotionality, 39 young horses (Dutch Warmblood) were tested repeatedly in two learning tests. An aversive stimulus (AS) was used in one learning test (the avoidance learning test) and a reward was used in the other learning test (the reward learning test). During both learning tests behaviour as well as heart rate were measured. Each test was executed four times, twice when horses were 1 year of age, and twice when they were 2 years of age. Half of the horses received additional physical training from 6 months onwards. In both tests horses could be classified as either performers, i.e. completing the daily session, or as non-performers, i.e. returning to the home environment without having completed the daily session. There were some indications that emotionality might have caused non-performing behaviour, but these indications are not convincing enough to exclude other causes. Furthermore, there seem to be no simple relationships between measures of heart rate, behavioural responses putatively related to emotionality and learning performance. Horses revealed consistent individual learning performances within years in both tests, and in the avoidance learning test also between years. There was no significant correlation between learning performances in the avoidance learning test and the learning performances in the reward learning test. It is concluded that individual learning abilities are consistent over a short time interval for an avoidance learning test and a reward learning test and over a longer time for the avoidance learning test. Furthermore, results indicate that some horses perform better when they have to learn to avoid an aversive stimulus while others perform better when they are rewarded after a correct response. It is suggested that these differences may be relevant to design optimal individual training programmes and methods.
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Call Number refbase @ user @ Serial 2009
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Author Leca, J.-B.; Gunst, N.; Thierry, B.; Petit, O.
Title Distributed leadership in semifree-ranging white-faced capuchin monkeys Type Journal Article
Year 2003 Publication Animal Behaviour. Abbreviated Journal Anim. Behav.
Volume 66 Issue 6 Pages 1045-1052
Keywords
Abstract We investigated the initiation of group movements in white-faced capuchin monkeys, Cebus capucinus, with the aim of determining whether a single individual with high dominance status consistently leads movements or whether leadership is distributed between group members. The group studied was reared in semifree-ranging conditions. A multivariate analysis followed by univariate analyses demonstrated that leadership was not concentrated on a single individual in this species. All individuals could initiate a collective movement. Nearly half of group members regularly succeeded in recruiting at least three followers. Although both sexes had similar rates of start attempts, females succeeded more frequently than males. We found no significant effect of the dominance status on the percentage of successful attempts. The use of a slow speed, looking back towards the other group members, or trills by the initiator heightened the likelihood of success in group movement initiation. An initiator starting from a core position in a clumped group was more successful than one starting from an edge position in a clumped group or from a dispersed group. Furthermore, the probability of successful start attempts was higher when the group remained stationary for a long period. Leadership in white-faced capuchins appears to be distributed between group members rather than exclusively concentrated on high-ranking individuals. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
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Call Number Serial 2028
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Author Reale, D.; Festa-Bianchet, M.
Title Predator-induced natural selection on temperament in bighorn ewes Type Journal Article
Year 2003 Publication Animal Behaviour. Abbreviated Journal Anim. Behav.
Volume 65 Issue 3 Pages 463-470
Keywords
Abstract Temperament traits in animals may have important fitness consequences, but have received little attention from ecologists or evolutionary biologists. A few studies have linked variation in temperament with fitness, but none has measured selection on temperament traits. We estimated the strength of selection on female boldness and docility on bighorn sheep ewes, Ovis canadensis. The Ram Mountain population experienced little predation pressure during the first 25 years of study, then 2 years (1997 and 1998) of frequent predation by cougars, Puma concolor, during which adult ewe mortality almost tripled over the long-term average, to 27% a year. During years of high predation, we found moderate selection favouring bold ewes, and age-specific selection on docility. Old ewes appeared more vulnerable to predation than young ewes. In contrast, no evidence of selection on temperament traits was observed during 2 years of low predation (1996 and 1999). These results suggest predator-induced selection favouring bold and nondocile ewes. Leadership was highly correlated with age and may increase the risk of predator encounter. Leadership alone, however, could not explain the higher vulnerability of old ewes to predation. Cougar predation on bighorn sheep occurs sporadically and unpredictably, probably because individual cougars often are prey specialists. Cougar predation may have limited microevolutionary effects on temperament in bighorn sheep, because it mostly affects ewes near the end of their reproductive life span and because of potential countervailing selection on boldness and docility. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.
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Language Summary Language Original Title
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Notes Approved no
Call Number Serial 2029
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