Why Dominant Individuals Cooperate — Fitness Consequences of Cooperative Courtship in a System with Variable Cooperative Display Coalitions

Understanding the evolution of cooperative behaviors is a major goal of evolutionary biology, but the majority of research in this field has focused on why helpers assist others. Helpers’ reproductive costs introduce a clear paradox to our understanding of natural selection as helpers in cooperative systems apparently sacrifice reproductive opportunities to increase others’ fitness. This puzzle in cooperative behaviors has led to significant advances in our understanding of indirect and delayed fitness benefits for helpers. However, as cooperation results from the interaction of individuals that may have very different incentives for participation it is equally important to understand whether and how cooperation benefits the dominant recipients of this help. There has been relatively little attention paid to why the recipient of the apparent help participates in the cooperative relationship, in part because the advantage to the dominant individual seems apparent in many systems. Existing work reveals a variety of potential benefits for dominant individuals and that the benefits for dominants may be less obvious than assumed. To date investigations into costs and benefits of cooperation to dominant individuals have been largely limited to cooperative breeding behavior. My dissertation research investigates the fitness consequences of cooperative courtship display for dominant individuals, in the White-ruffed Manakin, Corapipo altera. Manakins (Aves: Pipridae) are small, primarily lekking passerines, and, in some species, males cooperate in their courtship displays. Previous work on manakin cooperative display behavior has focused on benefits to subordinate males. The fitness consequences of cooperation for dominant individuals has not yet addressed in a system with variation in cooperative strategies. I found strong evidence of cooperation among male C. altera. I also found that, within a single population of C. altera on the Atlantic slope of Costa Rica, some males participate in coordinated display with other males (45.4±20% were classified as cooperative in any given year), while other males appear to only display singly. My dissertation research investigated the causes and consequences of cooperation by dominant C. altera males by quantifying aspects of the males' fitness including how inclusive fitness benefits may facilitate the maintenance of cooperative display coalitions and the consequences of cooperative display coalitions for males’ annual reproductive success, survival, and social status — important parts of lifetime fitness for long-lived, iteroparous species including C. altera. I found that cooperative males were not more closely related than expected at random from the population. Males that cooperated did not have higher annual reproductive success than males that displayed solo nor was there a significant difference in the frequency of copulations after a solo courtship display and a courtship display by multiple males. In a survival analysis, cooperation did not significantly affect the survival of dominant males. There was no consistent pattern of cooperation (or non-cooperation) among males across their tenure as dominant male: some were always cooperative, some always non-cooperative, but many males with multi-year tenures switched between cooperative and non-cooperative statuses. However, more males than expected employed strictly solo strategies across their tenure as dominant individuals, given the population-wide rates of survival and cooperation. The degree to which males cooperated, defined as the proportion of tenure classified as cooperative, was unrelated to variation in lifespan or length of tenure as a dominant male. Additionally, the proportion of total tenure classified as cooperative did not explain the patterns of lifetime reproductive success. Together, these results reject the hypotheses that dominant males in cooperative partnerships gain indirect or direct fitness benefits from their associations with subordinate males. Seeking to understand processes underlying patterns of fitness consequences from cooperative behaviors, I conducted three experiments to determine if males at sites where the dominant male was cooperative were faster or more intense in their response to an experimental stimulus. Cooperative males were not faster to respond to a female at the display site nor were they faster to respond to the vocalization of an unknown male conspecific at the display site. Cooperative males were not significantly more likely to respond to a predator model, however, they were significantly more likely to spend time near the snake and lizard models. There could be benefit of sociality in the detection of terrestrial predators. This research addresses previously unexplored aspects of cooperative courtship display, and therefore represents a significant contribution to the more general understanding of the costs and benefits of cooperation. The variation in the amount of cooperation expressed by different individuals of this species offers a unique opportunity to separate the fitness consequences of cooperation by comparing differences in success not only among individuals, but also those among displays in different cooperative contexts by the same individual.