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Bartos, L., Bartosova, J., Pluhacek, J., & Sindelarova, J. (2012). Promiscuous behaviour disrupts pregnancy block in domestic horse (Equus caballus) mares: A counterstrategy against possible male infanticide. In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: It is a common practice that the domestic horse mare is removed from her home environment and transported elsewhere for mating. After conceiving she is returned back into her home environment and social group, containing often familiar stallions and geldings. If we presume that the behavioural adaptation for infanticide relevant for wild, or free-ranging horses has not be lost in domestication (and it is clear that it remains at least within feral populations), then we may expect that, unless prevented by fencing or other management measures, the dominant males in the home social group may subsequently attempt infanticide. Foetal loss is a common phenomenon in domestic horses, being usually substantially higher than that in other domestic ungulates reaching up to 40%. One adaptive response to infanticide risk in polygynous populations is the Bruce effect. Pregnant females terminate their pregnancies when exposed to a dominant strange male. Our previous studies on plain zebra suggested the possibility of the Bruce effect also in equids. In this study we therefore tested the presumption that the Bruce effect could explain foetal loss in domestic horses. Based on reproduction records from horse breeders, we have recently published that bringing a pregnant mare which had been mated away from home into a vicinity of a familiar male who was not the father of her foetus increased probability of pregnancy disruption (Behav Ecol Sociobiol DOI: 10.1007/s00265-011-1166-6). These mares aborted in 31% of cases, while none of those mated within the home stable aborted. Repeated sexual activity either by a stallion or dominant gelding from the normal home group was observed shortly after the mare came from away-mating. Pregnant mares isolated from home males by a fence were even seen soliciting them over the fence. Therefore, there is probably some other mechanism than the Bruce effect leading to pregnancy block in the mare. We speculate that, once returned to the home “herd”, and introduced to familiar males, mares were more likely to terminate their pregnancy to save energy and avoid likely future infanticidal loss of their progeny by dominant male(s) of the home social group. Additional data has now showed that if a mare was mated away from home and was brought into an environment containing mares only, she was less likely to abort than a mare returning to an environment containing familiar male or males. This further supports our above hypothesis. Supported by AWIN, EU FP7 project No. 266213.
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Bartosova, J., Komarkova, M., Dubcova, J., Bartos, L., & Pluhacek J. (2012). Nursing behaviour in pregnant domestic mares (Equus caballus): Can they cope with dual maternal investment? In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Among mammals, lactation is the most energy demanding part of parental care and so parentoffspring conflict should arise over milk provided by the mother. Mother and offspring should disagree over the length and amount of the milk provision. We focused on effect of pregnancy on suckling behaviour variables as indicators of mother-offspring conflict in domestic horses. We presumed shorter suckling bouts and higher rates of rejected and/or terminated suckling in pregnant mares compared to non-pregnant ones. Increasing conflict over amount of maternal investment between mother and her young are to be expected because of her parallel investment into a nursed foal and a foetus. Eight groups of loose housed lactating mares with foals of Kladruby horse were studied at the National Stud Kladruby nad Labem (Czech Republic) from deliveries to abrupt weaning (at the age of 127 to 210 days). We recorded 10 848 suckling solicitations of 79 mare-foal pairs, from which 10 607 resulted in a suckling bout. In 41 cases a nursing mare became pregnant during lactation. We found no significant effect of pregnancy either on probability of the mother rejecting suckling solicitation of her foal or probability that she terminated a suckling bout. However the overall effect of mother’s pregnancy on suckling bout duration was not significant, there were considerable differences in pregnant and non-pregnant mares according to who terminated a suckling bout, whether the mother or the foal (F(1, 9776) = 12.1, P < 0.001). In case it was the mother then the suckling bout was longer if she was pregnant (65.36 ± 1.25 s) than barren (60.55 ± 1.36 s). We found no impact of pregnancy on duration of suckling bouts terminated by the foal. Further, nursing a foal during the first two trimesters of pregnancy had no negative impact on birth weight of the foetus. In conclusion, we found not higher, but a lower mother-offspring conflict in pregnant than in non-pregnant lactating mares while expecting just the opposite. We suggest that pregnant mares compensate their nursed foals during intensive stages of lactation through a relaxed mother-offspring conflict for later decrease in investment due to increasing demands of the foetus and/or for the shorter period of milk supply. Our results (partly published in Bartosova et al. 2011, PLoS ONE 6(8): e22068) are of high importance in horse breeding. One of the main arguments for early weaning of the foals is regeneration of their pregnant mothers before upcoming delivery. Here we present evidence that a pregnant mare “counts” with her dual maternal investment and “employs” evolutionary mechanisms enabling her to rear a vital foetus. From this point of view there is no objective reason for stressful weaning of her nursed offspring practised in conventional breeding. Supported by AWIN, EU FP7 project No. 266213.
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Hartmann, E., Søndergaard, E., & Keeling, L. J. Keeping horses in groups: A review. Appl. Anim. Behav. Sci., .
Abstract: Although husbandry conditions for horses have improved over the last decades, many horses are still kept singly with limited or no physical contact to other horses. This is surprising, given the fact that keeping horses in groups is recognised best to fulfil their physical and behavioural needs, especially their need for social contact with conspecifics, as well as to have a beneficial effect on horse–human interactions during training. Group housing of farm animals is widely applied in practice. As a consequence, scientists have investigated numerous aspects of group housing to help further improve animal welfare and human–animal interactions under these conditions. However, compared to this literature available in farm animals, and the plentiful studies conducted of feral horse populations, there is much less done when it comes to the management of horses kept in groups in the domestic environment. In particular, limited scientific information is available into the effect of group size and group composition on behaviour and methods of introducing new horses into established groups, even though problems related to social integration are repeatedly taken as arguments against keeping horses in groups. This review, therefore, aims to provide an overview of the current scientific knowledge regarding keeping horses in groups. Furthermore, it aims to give insight into whether or not some of the concerns related to keeping horses in groups are justified and to review scientifically based solutions that could be useful in practice to improve horse welfare and human safety.
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König von Borstel, U., Pasing, S., & Gauly, M. (2011). Towards a more objective assessment of equine personality using behavioural and physiological observations from performance test training. Appl. Anim. Behav. Sci., 135(4), 277–285.
Abstract: Current definitions of horse personality traits are rather vague, lacking clear, universally accepted guidelines for evaluation in performance tests. Therefore, the aim of the present study was to screen behavioural and physiological measurements taken during riding for potential links with scores the same horses received in the official stallion performance test for rideability and personality traits. Behaviour, heart rate (HR) and HR variability from thirty-six stallions participating in a performance test were recorded repeatedly during their performance test training. Using the coefficient of determination, regression analysis revealed that about 1/3 of variation (ranging between r = 0.26 (“constitution” (i.e. fitness, health)) and r = 0.46 (rideability)) in the personality trait scores could be explained by selecting the three most influential behaviour patterns per trait. These behaviour patterns included stumbling (with all traits except character), head-tossing (temperament, rideability), tail-swishing (willingness to work), involuntary change in gait (character) and the rider's use of her/his hands (constitution, rideability), voice (temperament) or whip (constitution). Subsequent mixed model analysis revealed a significant (P < 0.05) influence of the behaviour pattern “horse-induced change in gait” on character (-0.98 ± 0.31 scores per additional occurrence of change in gaits), of head-tossing (-0.25 ± 0.08 scores) and rider's use of voice (-0.51 ± 0.25; P = 0.0594) on temperament, and of stumbling on each of the following: willingness to work (-2.5 ± 1.2), constitution (-2.5 ± 1.2 scores; P = 0.0516) and rideability scores (-3.3 ± 1.4). In addition, constitution scores tended (P = 0.0889) to increase with higher low frequency/high frequency heart rate variation ratios (LF/HF), indicating a shift towards sympathetic dominance and thus a higher stress load in horses with higher scores for constitution. Rideability scores from the training phase were also significantly influenced by head-tossing (-0.5 ± 0.1), and in addition rideability scores from the final test were influenced by the training rider, ranging between average estimated rideability scores of 6.8 ± 0.4 for one training rider and 8.36 ± 0.3 scores for another training rider. Horses ridden with their nose-line predominantly behind the vertical received higher scores for rideability (8.3 ± 0.3) than horses ridden with their nose-line at the vertical (7.7 ± 0.2). These findings indicate that either judges perceive horses to have a better rideability when they readily offer a more extreme poll flexion, or that riders make use of horses’ better rideability by imposing a more extreme poll flexion. Several of the above described associations, but also of the non-existing links (e.g. no association between shying or heart rate and temperament) between behaviour patterns and scores for personality traits are rather surprising, warranting further investigation regarding the underlying causes of these relationships. Some of these behaviour patterns should be considered when redesigning the current guidelines for evaluation of personality traits during breeding horse performance tests, ultimately leading to improved genetic selection for equine personality traits. However, ethical implication of defining aversive behaviour such as head-tossing as an indicator of, for example, poor temperament, should not be neglected when devising new guidelines: such aversive behaviour may in fact be an indication of inadequate training techniques rather than poor horse personality.
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Menke, C., Waiblinger, S., Foelsch, D. W., & Wiepkema, P. R. (1999). Social Behaviour and Injuries of Horned Cows in Loose Housing Systems. Anim Welfare, 8(3), 243–258.
Abstract: The relationship between social behaviour and skin injuries (caused by horns) of loose housed horned cows was investigated on 35 dairy farms. While the frequencies of two agonistic behaviour elements (push and chase away) were positively correlated with the occurrence of skin injuries, the frequencies of butting and homing were not. Butting appears to have an ambivalent motivation, in that its occurrence is correlated positively both with agonistic behaviour and with social licking. Horning showed a positive correlation with social licking only. Four groups of husbandry conditions that may be associated with the occurrence of social behaviour and of injuries were distinguished: i) herd management, with variables including problem solving management by the farmer, integration of new cows, and dealing with periparturient and oestrus cows; ii) human-animal relationship, with variables including ability to identify individual cows, frequency of brushing the cows, number of milkers, and frequency of personnel changes; iii) animal characteristics, with the variable of herd size; and iv) stable characteristics, with the variable of space per cow (m2). The relevance of the husbandry variables investigated here had been confirmed in a previous stepwise regression analysis (Menke 1996). The variables for herd management and human-animal relationship conditions correlated in a consistent way with the occurrence of agonistic behaviour and/or of injuries, while most of them also correlated in the opposite direction with the occurrence of social licking. Herd size correlated positively with agonistic behaviour, but negatively with social licking. Space per cow correlated negatively with agonistic behaviour and injuries. In more than 70 per cent of the herds investigated, the levels of agonistic behaviour and of skin injuries were low, implying that horned dairy cows can be kept with less risk than is often assumed. We argue that such risks strongly depend on management factors that can be improved.
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Weng, R. C., Edwards, S. A., & English, P. R. (1998). Behaviour, social interactions and lesion scores of group-housed sows in relation to floor space allowance. Appl. Anim. Behav. Sci., 59(4), 307–316.
Abstract: The space allowance appropriate for sows in group housing remains scientifically undefined, since the social space requirement of a group of animals and the factors which affect this are unknown. Eight established groups of six pregnant, multiparous sows were used in a replicated Latin Square design of experiment, with 7 day periods, to compare four pen sizes providing 2.0, 2.4, 3.6 or 4.8 m2/sow. For the last 48 h of each 7 day period, a continuous video recording was made to determine general behaviour and all social interactions. Time spent rooting increased progressively with increasing space allowance, whereas time spent sitting and standing inactive were both progressively reduced. The total frequency of social interactions and aggressive behaviour both increased with decreasing space allowance. The Attack:Retreat ratio was significantly higher, and the Avoidance Index significantly lower, in the smallest pen. All body regions had the highest count of lesions after sows had been in the smallest pen, with damage levels being reduced as pen area increased. Analysis of body lesion scores, combining incidence and severity, gave the same treatment effects. In conclusion, the results indicated that a minimum space of between 2.4 and 3.6 m2/sow was necessary in the conditions of this experiment to promote good welfare. This result cannot be generalised to situations of different group size, group stability or feeding method.
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Garamszegi, L. Z., Møller, A. P., & Erritzøe, J. (2002). Coevolving avian eye size and brain size in relation to prey capture and nocturnality. Proc Roy Soc Lond B Biol Sci, 269(1494), 961–967.
Abstract: Behavioural adaptation to ecological conditions can lead to brain size evolution. Structures involved in behavioural visual information processing are expected to coevolve with enlargement of the brain. Because birds are mainly vision–oriented animals, we tested the predictions that adaptation to different foraging constraints can result in eye size evolution, and that species with large eyes have evolved large brains to cope with the increased amount of visual input. Using a comparative approach, we investigated the relationship between eye size and brain size, and the effect of prey capture technique and nocturnality on these traits. After controlling for allometric effects, there was a significant, positive correlation between relative brain size and relative eye size. Variation in relative eye and brain size were significantly and positively related to prey capture technique and nocturnality when a potentially confounding variable, aquatic feeding, was controlled statistically in multiple regression of independent linear contrasts. Applying a less robust, brunching approach, these patterns also emerged, with the exception that relative brain size did not vary with prey capture technique. Our findings suggest that relative eye size and brain size have coevolved in birds in response to nocturnal activity and, at least partly, to capture of mobile prey.
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Søndergaard, E., Jensen, M. B., & Nicol, C. J. (2011). Motivation for social contact in horses measured by operant conditioning. Appl. Anim. Behav. Sci., 132(3-4), 131–137.
Abstract: Although horses are social animals they are often housed individually with limited social contact to other horses and this may compromise their welfare. The present study included eight young female horses and investigated the strength of motivation for access to full social contact, head contact and muzzle contact, respectively, to a familiar companion horse. Horses were housed individually next to their companion horse and separations between pens prevented physical contact. During daily test sessions horses were brought to a test area where they could access an arena allowing social contact. Arena access during 3 min was given after completion of a predetermined number of responses on a panel. Fixed ratios (FR) of 8, 16, 24, 32 and 40 responses per arena access were applied in a random order, one per daily test session, within each test week (Monday to Friday), and the number of rewards per daily test session was recorded. All horses could access all three types of social contact in a cross-over design, and an empty arena was used as control. Motivational strength was assessed using elasticity of demand functions, which were estimated based on the number of rewards earned and FR. Elasticities of demand for the three types of social contact were low (-0.20), and not significantly different, although increasing FR still resulted in a decrease in rewards obtained for all three types of social contact (P < 0.001). Across FR-levels horses earned more rewards for social contact than for an empty arena, as shown by much higher intercept values (2.51 vs. 0.99; P < 0.001). However, the elasticity of demand for infrequent access to an empty arena (-0.08) was lower than for social contact (P < 0.01) and not significantly different from zero (P = 0.07). Horses performed more social behaviour the lesser the restriction on social contact (full > head > muzzle). However, the finding that horses showed a similar and high motivation for all three types of social contact suggests that they are valued equally highly in a situation where the alternative is no social contact.
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Normando, S., Meers, L., Samuels, W. E., Faustini, M., & Ödberg, F. O. (2011). Variables affecting the prevalence of behavioural problems in horses. Can riding style and other management factors be significant? Appl. Anim. Behav. Sci., 133(3-4), 186–198.
Abstract: The effects of riding style and various management factors on the prevalence of stereotypies and other behavioural problems among 346 mixed-breed saddle horses (phase 1) and 101 Arabian horses (phase 2) were analysed through a questionnaire answered by owners. In phase 1, the questionnaire data were partially validated through 20-min observations of 81 (23.3%) of the cases. Results indicate that horses primarily ridden in the English style were reported to be significantly more likely to display stereotypies (p < 0.001), problems when transported (p = 0.001), multiple behavioural problems (p < 0.001), and to have more restrictive stabling (p < 0.001) than horses ridden with other styles. When only Arabian horses were assessed in phase 2, however, there was no significant difference in behavioural problems between the Arabian horses ridden English style versus other riding styles. However Arabian horses were housed less restrictively than horses in phase 1 and English riding style and restrictive stabling tended to exacerbate each other's association with stereotypies. Management-related effects were found when, e.g., horses housed in restrictive stabling were more frequently reported to show locomotion stereotypies (p = 0.02) and those denied ad libitum hay displayed stereotypic wood-chewing behaviour (p = 0.02). To aid diagnosing and prioritizing interventions and care, the most predictive subsets of factors were computed for the various problem behaviours. E.g., among saddle horses, a statistical model comprised of the main riding style, duration of access to a paddock, and horse's age predicted whether a horse was reported to display any behavioural problem 62% of the time. This study supports the effects of management and handling on the prevalence of behavioural problems, and helps prioritize the relative importance of broad management categories on equine welfare. In particular, it underscores the importance of riding style on the well-being of saddle horses.
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Weissing, F. J. (2011). Animal behaviour: Born leaders. Nature, 474(7351), 288–289.
Abstract: Social animals face a dilemma. To reap the benefits of group living, they have to stay together. However, individuals differ in their preferences as to where to go and what to do next. If all individuals follow their own preferences, group coherence is undermined, resulting in an outcome that is unfavourable for everyone. Neglecting one's own preferences and following a leader is one way to resolve this coordination problem. But what attributes make an individual a 'leader'? A modelling study by Johnstone and Manica1 illuminates this question.
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