In many animal groups, certain individuals consistently appear at the forefront of coordinated movements [1-4]. How such leaders emerge is poorly understood [5, 6]. Here, we show that in pairs of sticklebacks, Gasterosteus aculeatus, leadership arises from individual differences in the way that fish respond to their partner's movements. Having first established that individuals differed in their propensity to leave cover in order to look for food, we randomly paired fish of varying boldness, and we used a Markov Chain model to infer the individual rules underlying their joint behavior. Both fish in a pair responded to each other's movements-each was more likely to leave cover if the other was already out and to return if the other had already returned. However, we found that bolder individuals displayed greater initiative and were less responsive to their partners, whereas shyer individuals displayed less initiative but followed their partners more faithfully; they also, as followers, elicited greater leadership tendencies in their bold partners. We conclude that leadership in this case is reinforced by positive social feedback.

Central to our understanding of social group formation and maintenance is the question of how withingroup conflict resolution is achieved in the face of asymmetrical competition over resources and reproduction. A crucial yet implicit assumption of many conflict resolution models dealing with reproductive skew is that subordinates have perfect knowledge of the extent of conflict between themselves and their dominants, enabling behavioural responses on an individual rather than evolutionary scale. However, a mechanism enabling subordinates to accurately assess their relative conflict levels has yet to be empirically demonstrated. Here, we show in the angelfish Centropyge bicolor that the rate of overt mild aggression from dominants to subordinates acts as a signal of increasing rank conflict. The clarity of this signal can be reduced by spatial segregation, causing subordinates to be less able to respond appropriately by regulation of their foraging rates. A reduced signal ultimately leads to a less well-defined dominance hierarchy and destabilization of the social group. Our study suggests that, contrary to previous suggestions, dominant aggression rates play a crucial role as an accurate information signal required for the evolutionary stability of skew models.

No abstract

Social groups are often structured by dominance hierarchies in which subordinates consistently defer to dominants. High‐ranking individuals benefit by gaining inequitable access to resources, and often achieve higher reproductive success; but may also suffer costs associated with maintaining dominance. We used a large‐scale field study to investigate the benefits and costs of dominance in the angelfish Centropyge bicolor, a sequential hermaphrodite. Each haremic group contains a single linear body size‐based hierarchy with the male being most dominant, followed by several females in descending size order. Compared to their subordinate females, dominant males clearly benefited from disproportionately high spawning frequencies, but bore costs in lower foraging rates and greater aggressive defence of their large territories. Within the female hierarchy, more dominant individuals benefited from higher spawning frequencies and larger home ranges, but displayed neither higher foraging rates nor spawn order priority. However, dominance in females was also linked to aggressiveness, particularly towards immediate subordinates, suggesting that females were using energetically costly aggression to maintain their high rank. We further showed by experimentally removing dominant females that the linear hierarchy was also a social queue, with subordinates growing to inherit higher rank with its attendant benefits and costs when dominants disappeared. We suggest that in C. bicolor, the primary benefit of high rank is increased reproductive success in terms of current spawning frequency and the prospect of inheriting the male position in the near future, which may be traded off against the cost of aggressively defending rank and territory.

1. The costs and benefits of behavioural care of offspring can often be easily quantified through observations and experiments. Other forms of parental investment, on the other hand, are usually less amenable to cost -benefit analysis.2. Here, the costs and benefits are estimated for protective egg coating by a chrysomelid beetle, Cryptocephalus hypochaeridis , where the female spends a considerable amount of time adding extra structural components to each of the eggs after laying them.3. Adding this protective coating was very costly, both in terms of material and energy used: the mass of the extrachorion is equivalent to half the mass of the egg, and water loss and energy expenditure while coating the egg is equivalent to half what would be lost while laying a further egg.4. Choice tests with egg predators demonstrated that these high costs are offset by benefits in terms of protection against predation: whereas uncoated eggs are readily eaten by predators, coated eggs are always rejected.