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Nuñez, C. M. V., Adelman, J. S., Smith, J., Gesquiere, L. R., & Rubenstein, D. I. (2014). Linking social environment and stress physiology in feral mares (Equus caballus): Group transfers elevate fecal cortisol levels. General and Comparative Endocrinology, 196, 26–33.
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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Nuñez, C. M. V., Adelman, J. S., Mason, C., & Rubenstein, D. I. (2009). Immunocontraception decreases group fidelity in a feral horse population during the non-breeding season. Appl. Anim. Behav. Sci., 117(1-2), 74–83.
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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Gueron, S., Levin, S. A., & Rubenstein, D. I. (1996). The Dynamics of Herds: From Individuals to Aggregations. J. Theor. Biol., 182(1), 85–98.
Abstract: The dynamic behavior of small herds is investigated by means of simulations of two-dimensional discrete-stochastic models. An individual-based approach is used to relate collective behavior to individual decisions. In our model, the motion of an individual in a herd is assumed to be the combined result of both density-independent and density-dependent decisions, in the latter case based on the influence of surrounding neighbors; assumed decision rules are hierarchical, balancing short range repulsion against long-range attraction. The probability of fragmentation of the model herd depends on parameter values. We explore the variety and characteristics of spatial patterns that develop during migration, for herds that are homogeneous and heterogeneous regarding intrinsic walking speeds. Group integrity can be maintained even in mixed populations, but fragmentation results for these more easily than for a homogeneous herd. Observations of natural populations suggest that animals move away from individuals that intrude too closely into their environment, but are attracted to individuals at a distance. Between these extremes, there appears to be a neutral zone, within which other individuals engender no response. We explore the importance of this neutral zone, and offer evolutionary interpretations. In particular, the neutral zone, if not too large, permits the individual to remain in contact with the herd, while reducing the frequency with which acceleration or deceleration must be undertaken. This offers obvious energetic benefits.
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Ginsberg, J. R., & Rubenstein, D. I. (1990). Sperm competiton and variation in zebra mating behaviour. Behav. Ecol. Sociobiol., 26(6), 427–434.
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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Fischhoff, I. R., Sundaresan, S. R., Cordingley, J., & Rubenstein, D. I. (2007). Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions. Behav. Ecol., 18(4), 725–729.
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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Fischhoff, I. R., Sundaresan, S. R., Cordingley, J., Larkin, H. M., Sellier, M. - J., & Rubenstein, D. I. (2007). Social relationships and reproductive state influence leadership roles in movements of plains zebra, Equus burchellii. Anim. Behav., 73(5), 825–831.
Abstract: In animal groups, collective movements emerge from individual interactions. Biologists seek to identify how characteristics of actors in these groups, and their relationships, influence the decision-making process. We distinguished two basic factors determining leadership in group choices: identity and state. We hypothesized that identity is more important to leadership in groups with stable relationships, which permit the development of habitual roles. In groups with fluid membership, particular individuals or subgroups are less likely to emerge as consistent leaders. Instead, we predicted that movement initiation in unstable groups depends on individual state at the time of the decision. We characterized how identity and reproductive state influenced leadership patterns in the movements of plains zebra. As in many other mammals, lactation in this species significantly alters water and energy needs. We investigated leadership in tightly knit harems and loosely bonded herds of multiple harems. Harem females tended to have habitual roles in the initiation of harem movement. In herds, however, we found no consistent leaders among harems. At both levels of social organization, lactation was a key determinant of leadership. In harems, lactating females were more likely to initiate movement than nonlactating females. In turn, harems containing lactating females were more likely to lead herd movements. Thus, we conclude that social relationships and reproductive state together shape the interactions that produce group behaviours. One benefit to lactating females of leading herd movements is preferential access to scarce water. Thus, leadership roles in group decisions may have fitness consequences.
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Fischhoff, I., Dushoff, J., Sundaresan, S., Cordingley, J., & Rubenstein, D. (2009). Reproductive status influences group size and persistence of bonds in male plains zebra (Equus burchelli). Behav. Ecol. Sociobiol., 63(7), 1035-1043.
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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