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Author Ginsberg, J. R; Rubenstein, D. I. doi  openurl
  Title Sperm competiton and variation in zebra mating behaviour Type Journal Article
  Year 1990 Publication Behavioral Ecology and Sociobiology Abbreviated Journal Behav. Ecol. Sociobiol.  
  Volume 26 Issue 6 Pages 427-434  
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  Abstract Data are presented on the breeding behavior of two zebra species to test whether intra- and interspecific variation in male reproductive behavior and physiology are correlated with differences in female promiscuity. In one species, plains zebra (Equus burchelli) females live in closed membership single male groups and mate monandrously. In the other species, the Grevy's zebra (E. grevyi) females live in groups whose membership is much more temporary. Typically, associations with individual males are brief and mating is polyandrous. However, some females – those having just given birth – reside with one male for long periods, mating monandrously. These differences in female mating behavior generate variability in the potential for sperm competition. We show that behavioral differences in male investment in reproductive activities correlate with the potential for sperm competition. When mating with promiscuous mares, Grevy's zebra stallions made a greater investment in reproductive behavior (calling, mounting, ejaculations) than did stallions of either species when mating with monandrous females. The evolution of large testes size in the Grevy's zebra, when compared to the congeneric plains zebra, horse, and mountain zebra, allows for this increased investment.  
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  Notes from Professor Hans Klingels Equine Reference List Approved yes  
  Call Number (up) Serial 1113  
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Author Rubenstein, D. I., openurl 
  Title Ecology and sociality in horses and zebras Type Book Chapter
  Year 1986 Publication Ecological Aspects of Social Evolution Abbreviated Journal Ecological Aspects of Social Evolution  
  Volume Issue Pages 282-302  
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  Publisher Princeton University Press Place of Publication Princeton, NJ. Editor Rubenstein, D. I. ; Wrangham, R. W.  
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  Notes from Professor Hans Klingels Equine Reference List Approved no  
  Call Number (up) Serial 1526  
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Author Rubenstein, D.I openurl 
  Title Life history and social organization in arid adapted ungulates Type Journal Article
  Year 1989 Publication Journal of Arid Environments Abbreviated Journal J. Arid. Environ.  
  Volume 17 Issue Pages 145-156  
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  Area Expedition Conference  
  Notes from Professor Hans Klingels Equine Reference List Approved yes  
  Call Number (up) Serial 1527  
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Author Rubenstein, D. I. isbn  openurl
  Title The ecology of female social behaviour in horses, zebras and asses Type Book Chapter
  Year 1994 Publication Animal Societies Abbreviated Journal Animal Societies  
  Volume Issue Pages 13-28  
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  Publisher Kyoto University Press Place of Publication Editor Jarman, P.J.;, Rositter, A.  
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  Series Volume Series Issue Edition  
  ISSN ISBN 4-87698-014-4 Medium  
  Area Expedition Conference  
  Notes Approved yes  
  Call Number (up) Serial 1528  
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Author Sundaresan, S.R.; Fischhoff, I.R.; Dushoff, J.; Rubenstein, D.I. doi  openurl
  Title Network metrics reveal differences in social organization between two fission-fusion species, Grevy's zebra and onager Type Journal Article
  Year 2007 Publication Oecologia Abbreviated Journal Oecologia  
  Volume 151 Issue 1 Pages 140-149  
  Keywords Analysis of Variance; Animals; Behavior, Animal/*physiology; *Environment; Equidae/*physiology; Female; India; Kenya; Male; *Models, Theoretical; *Social Behavior; *Social Environment  
  Abstract For species in which group membership frequently changes, it has been a challenge to characterize variation in individual interactions and social structure. Quantifying this variation is necessary to test hypotheses about ecological determinants of social patterns and to make predictions about how group dynamics affect the development of cooperative relationships and transmission processes. Network models have recently become popular for analyzing individual contacts within a population context. We use network metrics to compare populations of Grevy's zebra (Equus grevyi) and onagers (Equus hemionus khur). These closely related equids, previously described as having the same social system, inhabit environments differing in the distribution of food, water, and predators. Grevy's zebra and onagers are one example of many sets of coarsely similar fission-fusion species and populations, observed elsewhere in other ungulates, primates, and cetaceans. Our analysis of the population association networks reveals contrasts consistent with their distinctive environments. Grevy's zebra individuals are more selective in their association choices. Grevy's zebra form stable cliques, while onager associations are more fluid. We find evidence that females associate assortatively by reproductive state in Grevy's zebra but not in onagers. The current approach demonstrates the utility of network metrics for identifying fine-grained variation among individuals and populations in association patterns. From our analysis, we can make testable predictions about behavioral mechanisms underlying social structure and its effects on transmission processes.  
  Address Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ 08544, USA. ssundare@princeton.edu  
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  Language English Summary Language Original Title  
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  Series Volume Series Issue Edition  
  ISSN 0029-8549 ISBN Medium  
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  Notes PMID:16964497 Approved no  
  Call Number (up) Serial 1863  
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Author Nuñez, C.M.V.; Adelman, J.S.; Smith, J.; Gesquiere, L.R.; Rubenstein, D.I. url  doi
openurl 
  Title Linking social environment and stress physiology in feral mares (Equus caballus): Group transfers elevate fecal cortisol levels Type Journal Article
  Year 2014 Publication General and Comparative Endocrinology Abbreviated Journal  
  Volume 196 Issue Pages 26-33  
  Keywords Equus caballus; Fecal cortisol; Feral mare; Group transfer; Stress; Social instability  
  Abstract Abstract Feral horses (Equus caballus) have a complex social structure, the stability of which is important to their overall health. Behavioral and demographic research has shown that decreases in group (or band) stability reduce female fitness, but the potential effects on the physiological stress response have not been demonstrated. To fully understand how band stability affects group-member fitness, we need to understand not only behavioral and demographic, but also physiological consequences of decreases to that stability. We studied group changes in feral mares (an activity that induces instability, including both male and female aggression) on Shackleford Banks, NC. We found that mares in the midst of changing groups exhibit increased fecal cortisol levels. In addition, mares making more group transfers show higher levels of cortisol two weeks post-behavior. These results offer insights into how social instability is integrated into an animal’s physiological phenotype. In addition, our results have important implications for feral horse management. On Shackleford Banks, mares contracepted with porcine zona pellucida (PZP) make approximately 10 times as many group changes as do untreated mares. Such animals may therefore be at higher risk of chronic stress. These results support the growing consensus that links between behavior and physiological stress must be taken into account when managing for healthy, functional populations.  
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  Series Volume Series Issue Edition  
  ISSN 0016-6480 ISBN Medium  
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  Notes Approved no  
  Call Number (up) Equine Behaviour @ team @ Serial 5743  
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Author Rubenstein, D.I.; Hohmann, M.E doi  openurl
  Title Parasites and Social Behavior of Island Feral Horses Type Journal Article
  Year 1989 Publication Oikos Abbreviated Journal Oikos  
  Volume 3 Issue Pages 312-320  
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  Abstract The horses of Shackelford Banks, North Carolina, USA, are heavily parasitized by intestinal worms and harassed by dipterans, and although both types influence behavior only internal parasites affect bodily condition and the structuring of horse society. Thirteen species of internal parasites were identified, but only 4 of 13 groups contain them all and even within groups differences among individuals are large. Among individuals ova emissions vary ranging from 50 to 76,875 eggs per gram. The most important environmental factors influencing egg production are season and a group's location on the island, presumably because of salinity and soil differences and their effects on ova survival. Of the social and life history factors, age, and group size, but neither reproductive state nor dominance status are important. The fitness consequences of internal parasitism may be large since the number emitted is negatively correlated with next year's bodily condition. Biting fly burdens are also affected by a variety of environmental factors. In general, horses are covered with more flies on sunny days, when winds are moderately brisk, when occupying dunes, and around mid-day. In contrast to endoparasites, fly burden is affected by reproductive condition and dominance status and tends to decrease as groups increase in size. Since groups do not grow very large, nor do females attempt to bring groups together, the negative effects of endoparasites appear to overide those associated with ectoparasites. Consequently, endoparasites appear to exert a stronger influence on social structure, even though ectoparasites seem to play a stronger role in shaping details of behavior.  
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  Notes Approved no  
  Call Number (up) Equine Behaviour @ team @ Serial 2360  
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Author Fischhoff, I.R.; Sundaresan, S.R.; Cordingley, J.; Rubenstein, D.I. url  doi
openurl 
  Title Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions Type Journal Article
  Year 2007 Publication Behavioral Ecology Abbreviated Journal Behav. Ecol.  
  Volume 18 Issue 4 Pages 725-729  
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  Abstract Prey species must adapt their behavior to avoid predation. As a key prey item for lions (Panthera leo), plains zebras (Equus burchelli) were expected to respond to immediate threats posed by lions in their area. In addition, zebras were predicted to exhibit behavior tuned to reduce the potential for encounters with lions, by modifying their movement patterns in the times of day and habitats of greatest lion danger. We studied a population of approximately 600 plains zebra living in Ol Pejeta Conservancy, Kenya. We found that zebra abundance on or near a grassland patch was lower if lions had also been observed on that patch during the same day. Predation danger was highest in grassland habitat during the night, when lions were more active. Zebra sightings and global positioning system radio collar data indicated that zebras also reduced their use of grassland at night, instead using more woodland habitat. Zebras moved faster and took sharper turns in grassland at night. It is hypothesized that these more erratic movements assist zebras in avoiding detection or capture by lions.  
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  Notes 10.1093/beheco/arm036 Approved no  
  Call Number (up) Equine Behaviour @ team @ Serial 4360  
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Author Nuñez, C.M.V.; Adelman, J.S.; Mason, C.; Rubenstein, D.I. url  doi
openurl 
  Title Immunocontraception decreases group fidelity in a feral horse population during the non-breeding season Type Journal Article
  Year 2009 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.  
  Volume 117 Issue 1-2 Pages 74-83  
  Keywords Equus caballus; Horse; Harem; Stability; Immunocontraception; Porcine zona pellucidae (PZP); Behavior  
  Abstract The behavioral effects of the immunocontraceptive agent porcine zona pellucida (PZP) have not been adequately studied. Important managerial decisions for several species, including the wild horse (Equus caballus), have been based on this limited research. We studied 30 horses on Shackleford Banks, North Carolina, USA to determine the effects of PZP contraception on female fidelity to the harem male. We examined two classes of females: contracepts, recipients of the PZP vaccine (n = 22); and controls, females that have never received PZP (n = 8). We conducted the study during the non-breeding season from December 2005 to February 2006, totaling 102.2 h of observation. Contracepted mares changed groups more often than control mares (P = 0.04). Contracepts also visited more harem groups than did control mares (P = 0.02) and exhibited more reproductive interest (P = 0.05). For both contracepted and control females, the number of group changes (P = 0.01) and number of groups visited (P = 0.003) decreased with the proportion of years mares were pregnant. Our study shows that the application of PZP has significant consequences for the social behavior of Shackleford Banks horses. In gregarious species such as the horse, PZP application may disrupt social ties among individuals and inhibit normal social functioning at the population level.  
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  Series Volume Series Issue Edition  
  ISSN 0168-1591 ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number (up) Equine Behaviour @ team @ Serial 5095  
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Author Fischhoff, I.; Dushoff, J.; Sundaresan, S.; Cordingley, J.; Rubenstein, D. doi  openurl
  Title Reproductive status influences group size and persistence of bonds in male plains zebra (Equus burchelli) Type Journal Article
  Year 2009 Publication Behavioral Ecology and Sociobiology Abbreviated Journal Behav. Ecol. Sociobiol.  
  Volume 63 Issue 7 Pages 1035-1043-1043  
  Keywords Biomedical and Life Sciences  
  Abstract Animal groups arise from individuals’ choices about the number, characteristics, and identity of associates. Individuals make these choices to gain benefits from their associations. As the needs of an individual change with its phenotype, so too we expect the nature of its associations to vary. In this paper, we investigate how the social priorities of male plains zebra (Equus burchelli) depend on reproductive state. An adult male is either a bachelor, and lacking mating access, or a stallion defending a harem. Multiple harems and bachelor males aggregate in larger herds. Herds frequently split and merge, affording males opportunities to change associates. Over a 4-year period, we sampled the herd associations in a population of 500–700 zebras. To isolate the effects of reproductive state on male social behavior, we account for potential confounding factors: changes in population size, grouping tendencies, and sampling intensity. We develop a generally applicable permutation procedure, which allows us to test the null hypothesis that social behavior is independent of male status. Averaging over all individuals in the population, we find that a typical bachelor is found in herds containing significantly more adults, bachelors, and stallions than the herds of a typical stallion. Further, bachelors’ bonds with each other are more persistent over time than those among stallions. These results suggest that bachelors form cohesive cliques, in which we may expect cooperative behaviors to develop. Stallion–stallion associations are more diffuse, and less conducive to long-term cooperation.  
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  Publisher Springer Berlin / Heidelberg Place of Publication Editor  
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  Series Editor Series Title Abbreviated Series Title  
  Series Volume Series Issue Edition  
  ISSN 0340-5443 ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number (up) Equine Behaviour @ team @ Serial 5193  
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