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Clutton-Brock, T. H., Russell, A. F., Sharpe, L. L., Brotherton, P. N., McIlrath, G. M., White, S., et al. (2001). Effects of helpers on juvenile development and survival in meerkats. Science, 293(5539), 2446–2449.
Abstract: Although breeding success is known to increase with group size in several cooperative mammals, the mechanisms underlying these relationships are uncertain. We show that in wild groups of cooperative meerkats, Suricata suricatta, reductions in the ratio of helpers to pups depress the daily weight gain and growth of pups and the daily weight gain of helpers. Increases in the daily weight gain of pups are associated with heavier weights at independence and at 1 year of age, as well as with improved foraging success as juveniles and higher survival rates through the first year of life. These results suggest that the effects of helpers on the fitness of pups extend beyond weaning and that helpers may gain direct as well as indirect benefits by feeding pups.
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VanDierendonck, M. C., de Vries, H., & Schilder, M. B. H. (1995). An Analysis of Dominance, Its Behavioural Parameters and Possible Determinants in a Herd of Icelandic orses in Captivity. Netherl. J. Zool., 45(3-4), 362–385.
Abstract: Th e applicability of the concept of dominance was investigated in a captive herd of  Icelandic
horses and  ponies of diff erent breeds. Eight out of  behaviours possibly related
to dominance occurred frequently enough to be investigated in detail. For these eight agonistic
behaviours the coverage, the unidirectionality in the exchange, and the degree of
transitivity (Landau`s linearity index) were calculated. Four off ensive behaviours, together
with avoidance, were suitable for further analysis with regard to dominance. Th e patterns
of asymmetries with which these behaviours were exchanged were suffi ciently similar as to
justify the application of the dominance concept and to construct a (nearly) linear dominance
hierarchy. Th e rank order of the castrated stallions was completely linear, the hierarchy
of the mares was almost completely linear. Th e results suggest that off ensive and defensive
aggressive behaviours should be treated separately and that the concept of dominance
is applicable. However, ritualized formal dominance signals between adult horses appear to
be (almost) absent. Th e rank positions of the individuals were correlated with age and residency
in the herd but not with height. Middle ranking horses tended to be more frequently
in the close vicinity of another horse than high ranking or low ranking horses. Over and
above this correlation at the individual level, it was found that pairs of horses close in rank
to each other were more often also spatially close to each other. Being in oestrus did not infl
uence the dominance relationships between mares. For castrated stallions the rank positions
were correlated with the age at which they were castrated. Th is suggests that in male
horses experience prior to neutering infl uences the behaviour afterwards.
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Gammell, M. P., de Vries, H., Jennings, D. J., Carlin, C. M., & Hayden, T. J. (2003). David's score: a more appropriate dominance ranking method than Clutton-Brock et al.'s index. Anim. Behav., 66(3), 601–605.
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de Vries, H., Stevens, J. M. G., & Vervaecke, H. (2006). Measuring and testing the steepness of dominance hierarchies. Anim. Behav., 71(3), 585–592.
Abstract: In the analysis of social dominance in groups of animals, linearity has been used by many researchers as the main structural characteristic of a dominance hierarchy. In this paper we propose, alongside linearity, a quantitative measure for another property of a dominance hierarchy, namely its steepness. Steepness of a hierarchy is defined here as the absolute slope of the straight line fitted to the normalized David's scores (calculated on the basis of a dyadic dominance index corrected for chance) plotted against the subjects' ranks. This correction for chance is an improvement of an earlier proposal by de Vries (appendix 2 in de Vries, Animal Behaviour, 1998, 55, 827-843). In addition, we present a randomization procedure for determining the statistical significance of a hierarchy's steepness, which can be used to test the observed steepness against the steepness expected under the null hypothesis of random win chances for all pairs of individuals. Whereas linearity depends on the number of established binary dominance relationships and the degree of transitivity in these relationships, steepness measures the degree to which individuals differ from each other in winning dominance encounters. Linearity and steepness are complementary measures to characterize a dominance hierarchy.
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Poisbleau, M., Fritz, H., Valeix, M., Perroi, P. - Y., Dalloyau, S., & Lambrechts, M. M. (2006). Social dominance correlates and family status in wintering dark-bellied brent geese, Branta bernicla bernicla. Anim. Behav., 71(6), 1351–1358.
Abstract: In many gregarious species, including ducks and geese, being dominant provides more benefits than costs, because dominants have better access to resources essential for survival or reproduction. In geese, being in better body condition during migration towards the breeding grounds positively influences reproductive success. However, underlying proximate mechanisms linking prebreeding body condition on the wintering grounds to breeding success remain poorly understood. We investigated social dominance correlates and family status, in three consecutive winters, in a free-ranging, migrating, dark-bellied brent goose population. Families with juveniles dominated pairs, and pairs dominated singletons. Dominance rank did not increase with the number of juveniles per family. Males were dominant over females. Social dominance and reproductive status for a given winter were significantly correlated with body mass, body size and body condition during the previous winter, suggesting that body condition in winter also affects subsequent breeding success and hence also dominance. Levels of testosterone and triiodothyronine were not correlated with immediate or later dominance or reproductive status. We discuss the role of family status as a signal of social status in determining reproductive strategies.
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Poisbleau, M., Jenouvrier, S., & Fritz, H. (2006). Assessing the reliability of dominance scores for assigning individual ranks in a hierarchy. Anim. Behav., 72(4), 835–842.
Abstract: The dominance score (number of wins divided by the total number of interactions) is the most widely used procedure in field studies to rank individuals. Its reliability depends on the number of interactions on which it is calculated. However, most authors use it without any estimate of the associated error. We describe the precision associated with a dominance score estimate as a function of the number of interactions on which it is based, and hence provide a tool to plan field protocols and effort. The precision error decreases according to a power function with increasing number of interactions, but with more precision for extreme scores for any given number of interactions. We discuss the fact that the minimum number of interactions should be based on the precision associated with the 50% score, the least precise of all scores. We also emphasize the trade-off between recording effort and precision of the estimator, and give an example of our choice of 26 interactions for fieldwork on ducks and geese. When comparing individual ranks based on dominance scores with ranks given by the dominance matrix, we found a good correlation, with more mismatches around the middle of the hierarchy. This was consistent with the precision calculated with our model. We conclude that dominance score is a reliable tool, but conclusions must take into account the number of interactions on which the calculations are done. We also discuss the importance of initial assumptions and sources of bias in field studies.
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de VRIES, H. A. N. (1998). Finding a dominance order most consistent with a linear hierarchy: a new procedure and review. Anim. Behav., 55(4), 827–843.
Abstract: A procedure for ordering a set of individuals into a linear or near-linear dominance hierarchy is presented. Two criteria are used in a prioritized way in reorganizing the dominance matrix to find an order that is most consistent with a linear hierarchy: first, minimization of the numbers of inconsistencies and, second, minimization of the total strength of the inconsistencies. The linear ordering procedure, which involves an iterative algorithm based on a generalized swapping rule, is feasible for matrices of up to 80 individuals. The procedure can be applied to any dominance matrix, since it does not make any assumptions about the form of the probabilities of winning and losing. The only assumption is the existence of a linear or near-linear hierarchy which can be verified by means of a linearity test. A review of existing ranking methods is presented and these are compared with the proposed method.
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Klingel, H. (). Das Verhalten der Pferde (Equidae). Handb. o. Zool., 8(10), 1–68.
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Crowley, P. H., Provencher, L., Sloane, S., Dugatkin, L. A., Spohn, B., Rogers, L., et al. (1996). Evolving cooperation: the role of individual recognition. Biosystems, 37(1-2), 49–66.
Abstract: To evaluate the role of individual recognition in the evolution of cooperation, we formulated and analyzed a genetic algorithm model (EvCo) for playing the Iterated Prisoner's Dilemma (IPD) game. Strategies compete against each other during each generation, and successful strategies contribute more of their attributes to the next generation. Each strategy is encoded on a `chromosome' that plays the IPD, responding to the sequences of most recent responses by the interacting individuals (chromosomes). The analysis reported in this paper considered different memory capabilities (one to five previous interactions), pairing continuities (pairs of individuals remain together for about one, two, five, or 1000 consecutive interactions), and types of individual recognition (recognition capability was maximal, nil, or allowed to evolve between these limits). Analysis of the results focused on the frequency of mutual cooperation in pairwise interactions (a good indicator of overall success in the IPD) and on the extent to which previous responses by the focal individual and its partner were associated with the partner's identity (individual recognition). Results indicated that a fixed, substantial amount of individual recognition could maintain high levels of mutual cooperation even at low pairing continuities, and a significant but limited capability for individual recognition evolved under selection. Recognition generally increased mutual cooperation more when the recent responses of individuals other than the current partner were ignored. Titrating recognition memory under selection using a fitness cost suggested that memory of the partner's previous responses was more valuable than memory of the focal's previous responses. The dynamics produced to date by EvCo are a step toward understanding the evolution of social networks, for which additional benefits associated with group interactions must be incorporated.
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Naguib, M., Amrhein, V., & Kunc, H. P. (2004). Effects of territorial intrusions on eavesdropping neighbors: communication networks in nightingales. Behav. Ecol., 15(6), 1011–1015.
Abstract: Animal communication often occurs in communication networks in which multiple signalers and receivers are within signaling range of each other. In such networks, individuals can obtain information on the quality and motivation of territorial neighbors by eavesdropping on their signaling interactions. In songbirds, extracting information from interactions involving neighbors is thought to be an important factor in the evolution of strategies of territory defense. In a playback experiment with radio-tagged nightingales Luscinia megarhynchos we here demonstrate that territorial males use their familiar neighbors' performance in a vocal interaction with an unfamiliar intruder as a standard for their own response. Males were attracted by a vocal interaction between their neighbor and a simulated stranger and intruded into the neighbor's territory. The more intensely the neighbor had interacted with playback, the earlier the intrusions were made, indicating that males eavesdropped on the vocal contest involving a neighbor. However, males never intruded when we had simulated by a second playback that the intruder had retreated and sang outside the neighbor's territory. These results suggest that territorial males use their neighbors' singing behavior as an early warning system when territorial integrity is threatened. Simultaneous responses by neighboring males towards unfamiliar rivals are likely to be beneficial to the individuals in maintaining territorial integrity.
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