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White, A. M., Swaisgood, R. R., & Czekala, N. (2007). Ranging patterns in white rhinoceros, Ceratotherium simum simum: implications for mating strategies. Appl. Anim. Behav. Sci., 74(2), 349–356.
Abstract: How animals use space has important consequences for feeding ecology, social organization, mating strategies and conservation management. In white rhinoceros, female home ranges are much larger than male territories, suggesting that movement patterns are influenced by factors other than resource distribution. In this study we placed radiotransmitters on 15 female white rhinoceros, recording 1758 locations and collecting behavioural data during 1671 observation sessions, making this the largest data set of its kind in this species. We investigated how habitat variables and male territories influenced female movement and reproductive behaviour. Female home ranges were approximately 20 km2 and core areas were 5 km2, with male territories roughly the same size as female core areas. Female range size did not vary with season, but the pattern of space use did vary. Females used grassland habitat preferentially, utilizing these areas significantly more than expected based on availability. Findings relevant to the mating strategy include: (1) the amount of grassland in a male's territory predicted female use of the territory; (2) the time that a female spent in a male's territory was a significant predictor of reproductive activity with the male, indicating that females probably mate with the most familiar male; and (3) the temporal pattern of female space use suggests that females did not increase mate sampling behaviour nor did they become more choosy about which males they visited when reproductively active. These findings suggest that males may maximize reproductive success by defending areas containing more grassland habitat.
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Marneweck, C., Jürgens, A., & Shrader, A. M. (2017). Dung odours signal sex, age, territorial and oestrous state in white rhinos. Proc. R. Soc. Lond. B, 284(1846).
Abstract: Mammals commonly communicate olfactorily via urine. However, the extent to which they communicate via dung, another waste product, is unknown. Behavioural studies suggest that mammals can obtain information from dung odours but are unclear about the information transmitted. Moreover, an understanding of the volatile organic compounds (VOCs) released from dung is limited. To address this, we analysed the odours emitted from the dung of free-ranging white rhinos, and found that 2,3-dimethylundecane signalled an individual's sex, heptanal discriminated age class, nonane defined male territorial status and 2,6-dimethylundecane indicated female oestrous state. To validate these findings, we artificially reproduced key elements of the territorial and oestrous odour profiles (i.e. profiles likely to elicit behavioural responses from receivers). We then exposed free-ranging territorial males to these odours. In response, males elicited behaviours associated with the specific odours (e.g. territorial male (potential threat): reduced latency in assuming vigilance; oestrous female (potential mate): increased investigation). These results indicate that the VOCs identified from the dung of free-ranging individuals do transmit key information. Moreover, as white rhinos of all ages and sexes defecate communally, middens probably act as information centres. Furthermore, as many other mammals defecate communally, olfactory communication via dung odours is likely a widespread phenomenon.
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Brooks, C. J., & Harris, S. (2008). Directed movement and orientation across a large natural landscape by zebras, Equus burchelli antiquorum. Anim. Behav., 76(2), 277–285.
Abstract: We investigated how plains zebras moved across a large natural landscape by analysing the movement paths of nine zebra mares foraging out from spatially confined waterholes during the dry season in the Makgadikgadi Pans National Park, Botswana. Since it was essential to investigate directed movement over a range of spatial scales to determine the correct movement behaviour and strategy, we used Nams's scaling test for oriented movement. Zebras followed directed movement paths in the lower to medium spatial scales (10 m–3.7 km) and above their visual, and possibly olfactory, range. The spatial scale of directed movement suggests that zebras had a well-defined spatial awareness and cognitive ability. Seven zebras used directed movement paths, but the remaining two followed paths not significantly different to a correlated random walk (CRW). At large spatial scales (>3 km) no distinct movement pattern could be identified and paths could not be distinguished from a CRW. Foraging strategy affected the extent of directed movement: zebras with a confined dispersion of grazing patches around the central place directed their movements over a longer distance. Zebras may extend the distance at which they can direct their movement after improving their knowledge of the local environment.
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Murray, L. M. A., Byrne, K., & D’Eath, R. B. (2013). Pair-bonding and companion recognition in domestic donkeys, <em>Equus asinus</em>. Appl. Anim. Behav. Sci., 143(1), 67–74.
Abstract: Pair and social bonding has been documented in various taxa, where pair formations are often described as being driven by kinship or sexual motivation. However, pair-bonding between unrelated individuals where sexual motivation is not a factor is not well documented. Many social relationships and pair-bonds between members of a dyad are facilitated by each individual's ability to recognise their partner using cues which are characteristic of that particular individual. The aims of this study were i) to investigate the existence of pair-bonding in domestic donkeys and ii) to determine whether members of a dyad could recognise their companion during a Y-maze recognition test. Subjects were 55 unrelated donkeys (38 gelded males, 15 females) in seven groups of mixed or same sex, comprising 4?14 individuals. Spatial proximity (nearest-neighbour) was observed three times a day over a 22-day period. Using a simulation approach based on observed data to generate randomised nearest-neighbour matrices, the statistical significance of social relationships was estimated. Of these, 42 (79.2%) were involved in significantly (p<0.05) non-random nearest-neighbour relationships, most of which were reciprocal pair relationships. Based on the spatial data, 24 of the donkeys which had shown significant reciprocal nearest-neighbour preferences for one individual (companion) were then used in a Y-maze recognition test in which they were presented with a choice of their companion and either a familiar donkey from the same group or an unfamiliar donkey from a different group. Donkeys? spatial location in the Y-maze demonstrated a preference for their companion versus familiar (one sample Wilcoxon signed rank test, W=239, p=0.002) or unfamiliar donkeys (W=222, p=0.041). These results verify anecdotal evidence from donkey handlers that donkeys often form pair-bonds, and show that reciprocal social preference and recognition are the basis of these. Pair-bond formation and companionship among donkeys have potential implications for their management, husbandry and welfare.
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Schnabel, C. L., Babasyan, S., Freer, H., & Wagner, B. (2017). Quantification of equine immunoglobulin A in serum and secretions by a fluorescent bead-based assay. Veterinary Immunology and Immunopathology, 188, 12–20.
Abstract: Abstract Only few quantitative reports exist about the concentrations and induction of immunoglobulin A (IgA) in mucosal secretions of horses. Despite this, it is widely assumed that IgA is the predominant immunoglobulin on mucosal surfaces in the horse. Here, two new monoclonal antibodies (mAbs) against equine IgA, clones 84-1 and 161-1, were developed and characterized in detail. Both IgA mAbs specifically bound monomeric and dimeric equine IgA in different applications, such as Western blots and fluorescent bead-based assays. Cross-reactivity with other equine immunoglobulin isotypes was not observed. The new IgA mAb 84-1 was used in combination with the previously characterized anti-equine IgA mAb BVS2 for the development and validation of a fluorescent bead-based assay to quantify total IgA in equine serum and various secretions. The IgA assay's linear detection ranged from 64 pg/ml to 1000 ng/ml. For the quantification of IgA in serum or in secretions an IgA standard was purified from serum or nasal wash fluid (secretory IgA), respectively. The different standards were needed for accurate IgA quantification in the respective samples taking the different signal intensities of monomeric and dimeric IgA on the florescent bead-based assay into account. IgA was quantified by the bead-based assay established here in different equine samples of healthy adult individuals. In serum the median total IgA was 0.45 mg/ml for Thoroughbred horses (TB, n = 10) and 1.16 mg/ml in Icelandic horses (ICH, n = 12). In nasopharyngeal secretions of TB (n = 7) 0.13 mg/ml median total IgA was measured, and 0.25 mg/ml for ICH (n = 12). Saliva of ICH (n = 6) contained a median of 0.15 mg/ml, colostrum of Warmbloods (n = 8) a median of 1.89 mg/ml IgA. Compared to IgG1 and IgG4/7 quantified in the same samples, IgA appeared as the major immunoglobulin isotype in nasopharyngeal secretions and saliva while it is a minor isotype in serum and colostrum. The newly developed monoclonal antibodies against equine IgA and the resulting bead-based assay for quantification of total IgA can notably improve the evaluation of mucosal immunity in horses.
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Briard, L., Dorn, C., & Petit, O. (2015). Personality and Affinities Play a Key Role in the Organisation of Collective Movements in a Group of Domestic Horses. Ethology, 121(9), 888–902.
Abstract: Understanding how groups of individuals with different motives come to daily decisions about the exploitation of their environment is a key question in animal behaviour. While interindividual differences are often seen only as a threat to group cohesion, growing evidence shows that they may to some extent facilitate effective collective action. Recent studies suggest that personality differences influence how individuals are attracted to conspecifics and affect their behaviour as an initiator or a follower. However, most of the existing studies are limited to a few taxa, mainly social fish and arthropods. Horses are social herbivores that live in long-lasting groups and show identifiable personality differences between individuals. We studied a group of 38 individuals living in a 30-ha hilly pasture. Over 200 h, we sought to identify how far individual differences such as personality and affinity distribution affect the dynamic of their collective movements. First, we report that individuals distribute their relationships according to similar personality and hierarchical rank. This is the first study that demonstrates a positive assortment between unrelated individuals according to personality in a mammal species. Second, we measured individual propensity to initiate and found that bold individuals initiated more often than shy individuals. However, their success in terms of number of followers and joining duration did not depend on their individual characteristics. Moreover, joining process is influenced by social network, with preferred partners following each other and bolder individuals being located more often at the front of the movement. Our results illustrate the importance of taking into account interindividual behavioural differences in studies of social behaviours.
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McGreevy, P. (2012). Equine Behavior A Guide for Veterinarians and Equine Scientists.
Abstract: Chapter 1 – Introduction, Pages 1-36
Chapter 2 – Perception, Pages 37-54
Chapter 3 – Behavior and the brain, Pages 55-84, Caroline Hahn
Chapter 4 – Learning, Pages 85-118
Chapter 5 – Social behavior, Pages 119-150
Chapter 6 – Communication, Pages 151-163
Chapter 7 – Locomotory behavior, Pages 165-187
Chapter 8 – Ingestive behavior, Pages 189-215
Chapter 9 – Eliminative behavior, Pages 217-221
Chapter 10 – Body care, Pages 223-243
Chapter 11 – Behavior of the stallion, Pages 245-264
Chapter 12 – Behavior of the mare, Pages 265-290
Chapter 13 – Training, Pages 291-311, Andrew McLean, Paul McGreevy
Chapter 14 – Handling and transport, Pages 313-329
Chapter 15 – Miscellaneous unwelcome behaviors, their causes and resolution, Pages 331-345
Further reading, Page 347
Glossary, Pages 351-356
Index, Pages 357-369
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McDonnell, S. (1999). Understanding horse behavior. Your guide to horse health care and management. Lexington, KY 40544-4038: Blood-Horse Inc.
Abstract: The author has conducted much research on equine behaviour, and here presents her findings in a form suitable for owners of horses. Common behavioural problems are mentioned.
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Ward, A., & Webster, M. (2016). Sociality: The Behaviour of Group-Living Animals.
Abstract: Covers the aspects of social behaviour of animals in comprehensive form Provides a clear overview to up-to-date empirical and theoretical research on social animal behaviour
Discusses collective animal behaviour, social networks and animal personality in detail
The last decade has seen a surge of interest among biologists in a range of social animal phenomena, including collective behaviour and social networks. In ‘Animal Social Behaviour’, authors Ashley Ward and Michael Webster integrate the most up-to-date empirical and theoretical research to provide a new synthesis of the field, which is aimed at fellow researchers and postgraduate students on the topic. ​
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Wynne C. D. L. (2001). Animal Cognition: The Mental Lives of Animals. Palgrave.
Abstract: Covering a wide range of key topics, from reasoning and communication to sensation and complex problem-solving, this engagingly-written text presents a comprehensive survey of contemporary research on animal cognition. Written for anyone with an interest in animal cognition, but without a background in animal behaviour, it endeavours to explain what makes animals tick.
With numerous illustrations and including exciting recent studies from many little-studied species (such as the weakly electric African fish), this text is ideal for psychology students who are interested in how much of our human cognition is shared by other species, for students of biology who want to know how complex animal behaviour can get, and for all those with an interest in the animal mind.
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