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Cameron, E. Z.,, Linklater, W. L.,, Stafford, K. J.,, & Minot, E. O.,. (2003). Social grouping and maternal behaviour in feral horses (Equus caballus): the influence of males on maternal protectiveness. Behav. Ecol. Sociobiol., 53(2), 92–101.
Abstract: The risk of infant injury or mortality influences maternal behaviour, particularly protectiveness. Mares are found in bands with a single stallion or bands with more than one stallion in which paternity is less certain. We investigated maternal behaviour in relation to band type. Mares in bands with more than one stallion were more protective of their foals, particularly when stallions and foals approached one another. The rate of aggression between the stallion and foal was a significant predictor of maternal protectiveness, and mare protectiveness was significantly correlated with reduced reproductive success in the subsequent year. Mares that changed band types with a foal at foot, or had their band type experimentally altered, were more protective of their foal in multi-stallion bands than they were in single-stallion bands. Equids are unusual amongst ungulates in that infanticide and feticide have been reported. Both occur where paternity has been uncertain, and equid social structure is similar to other species in which infanticide has been reported. Stallions benefit from infanticide as the mare has greater reproductive success in the subsequent year. Stallion aggression is a significant modifier of mare behaviour and maternal effort, probably due to the risk of infanticide.
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Dugatkin, L., & Alfieri, M. (1991). Tit-For-Tat in guppies (Poecilia reticulata): the relative nature of cooperation and defection during predator inspection. Evol. Ecol., 5(3), 300–309.
Abstract: Summary The introduction of game-theoretical thinking into evolutionary biology has laid the groundwork for a heuristic view of animal behaviour in which individuals employ “strategies” – rules that instruct them how to behave in a given circumstance to maximize relative fitness. Axelrod and Hamilton (1981) found that a strategy called Tit-For-Tat (TFT) is one robust cooperative solution to the iterated Prisoner's Dilemma game. There exists, however, little empirical evidence that animals employ TFT. Predator inspection in fish provides one ecological context in which to examine the use of the TFT strategy.
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Rubenstein, D. I., & Hack, M. A. (1992). Horse signals: The sounds and scents of fury. Evol. Ecol., 6(3), 254–260.
Abstract: During contests animals typically exchange information about fighting ability. Among feral horses these signals involve olfactory or acoustical elements and each type can effectively terminate contests before physical contact becomes necessary. Dung transplant experiments show that for stallions, irrespective of rank, olfactory signals such as dung sniffing encode information about familiarity suggesting that such signals can be used as signatures. As such they can provide indirect information about fighting ability as long as opponents associate identity with past performance. Play-back experiments, however, show that vocalizations, such as squeals, directly provide information about status regardless of stallion familiarity. Sonographs reveal that squeals of dominants are longer than those of subordinates and that only those of dominants have at their onset high-frequency components.
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Houston, A. I., & McNamara, J. M. (1988). Fighting for food: a dynamic version of the Hawk-Dove game. Evol. Ecol., 2(1), 51–64.
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Berger, J. (1977). Organizational systems and dominance in feral horses in the Grand Canyon. Behav. Ecol. Sociobiol., 2(2), 131–146.
Abstract: 1. Several aspects of the behavioral ecology of feral horses (Equus caballus) were studied in Grand Canyon, Arizona, USA. Most bands contained three to five horses that included one stallion and his harem. Males that did not obtain a harem remained solitary. Throughout the study bands remained stable in composition.
2. Home ranges for all bands decreased in size in successive warm months, probably due to increased ambient temperature and drought. This resulted in greater utilization of spring areas that led to increased interband confrontation and agonistic display.
3. Territoriality was not observed in individual horses or bands, but bands hierarchial in both inter- and intraband structures. Interband stallion dominance was reinforced through posturing and fighting. Intraband hierarchies, as determined by dominance coefficients, were independent of individual size in three of four bands.
4. Indexes of nervousness (NER), calculated while horses were drinking, showed that stallions were less nervous than mares. A low NER was correlated with individuals leading toward drinking areas, whereas a high NER existed in individuals initiating flight although no single horse acted consistently as a leader.
5. Diurnal activity patterns were correlated with ambient temperatures.
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Dugatkin, L. A., & Alfieri, M. (1991). Guppies and the TIT FOR TAT strategy: preference based on past interaction. Behav. Ecol. Sociobiol., 28(4), 243–246.
Abstract: The evolution of cooperation requires either (a) nonrandom interactions, such that cooperators preferentially interact with other cooperators, or (b) conditional behaviors, such that individuals act cooperatively primarily towards other cooperators. Although these conditions can be met without assuming sophisticated animal cognition, they are more likely to be met if animals can remember individuals with whom they have interacted, associate past interactions with these individuals, and base future behavior on this information. Here we show that guppies (Poecilia reticulata), in the context of predator inspection behavior, can identify and remember (for at least 4 h) the “more cooperative” among two conspecifics and subsequently choose to be near these individuals in future encounters.
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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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Noë, R., & Hammerstein, P. (1994). Biological markets: supply and demand determine the effect of partner choice in cooperation, mutualism and mating. Behav. Ecol. Sociobiol., 35(1), 1–11.
Abstract: The formation of collaborating pairs by individuals belonging to two different classes occurs in the contexts of reproduction and intea-specific cooperation as well as of inter-specific mutualism. There is potential for partner choice and for competition for access to preferred partners in all three contexts. These selective forces have long been recognised as important in sexual selection, but their impact is not yet appreciated in cooperative and mutualistic systems. The formation of partnerships between members of different classes has much in common with the conclusion of trade agreements in human markets with two classes of traders, like producers and consumers, or employers and employees. Similar game-theoretical models can be used to predict the behaviour of rational traders in human markets and the evolutionarily stable strategies used in biological markets. We present a formal model in which the influence of the market mechanism on selection is made explicit. We restrict ourselves to biological markets in which: (1) Individuals do not compete over access to partners in an agonistic manner, but rather by outcompeting each other in those aspects that are preferred by the choosing party. (2) The commodity the partner has to offer cannot be obtained by the use of force, but requires the consent of the partner. These two restrictions ensure a dominant role for partner choice in the formation of partnerships. In a biological market model the decision to cooperate is based on the comparison between the offers of several potential partners, rather than on the behaviour of a single potential partner, as is implicitly assumed in currently accepted models of cooperation. In our example the members of one class A offer a commodity of fixed value in exchange for a commodity of variable value supplied by the other class, B. We show that when the B-class outnumbers the A-class sufficiently and the cost for the A-class to sample the offers of the B-class are low, the choosiness of the A-class will lead to selection for the supply of high value commodities by the B-class (Fig. 3a). Under the same market conditions, but with a high sampling cost this may still be the evolutionariy stable outcome, but another pair of strategies proves to be stable too: relaxed choosiness of class A coupled with low value commodities supplied by class B (Fig. 3b). We give a number of examples of mating, cooperative and mutualistic markets that resemble the low sampling cost situation depicted in Fig. 3a.
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Dugatkin, L. A. (1991). Dynamics of the TIT FOR TAT strategy during predator inspection in the guppy (Poecilia reticulata). Behav. Ecol. Sociobiol., 29(2), 127–132.
Abstract: One well-known solution to the iterated Prisoner's Dilemma is the TIT FOR TAT strategy. This strategy has three “characteristics” associated with it. TIT FOR TAT is nice (cooperates on the first move of a game), retaliatory (plays defect against an individual that defected on the prior move), and forgiving (cooperates with an individual which has defected in the past but cooperates in the present). Predator inspection behavior in guppies (Poecilia reticulata) was examined in order to determine whether guppies displayed these three characteristics. Results indicate that while it can be quite difficult to translate the abstract concepts of niceness, retaliation, and forgiveness into measurable behaviors, the data support the hypothesis that guppies display the three characteristics associated with the TIT FOR TAT strategy.
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Kirkpatrick, J. F., & Turner, J. W. (1991). Changes in herd stallions among feral horse bands and the absence of forced copulation and induced abortion. Behav. Ecol. Sociobiol., 29(3), 217–219.
Abstract: Forced copulation and induced abortion were investigated in a herd of feral horses inhabiting a coastal barrier island. Eight mares were diagnosed pregnant in August and October 1989 by means of urinary and fecal steroid metabolites, prior to documented changes in herd stallions. These mares were observed for harassment and forced copulation by the new stallions and for the presence of foals during the spring and summer of 1990. No incidents of harassment or attempts at forced copulation were witnessed and seven of the eight mares produced foals in 1990. These data indicate that forced copulation and induced abortion are not common events among all feral horse herds and suggest reinvestigation of this hypothesized phenomenon.
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