A Theoretical Comparison of Alternative Male Mating Strategies in Cephalopods and Fishes.

We used computer simulations of growth, mating and death of cephalopods and fishes to explore the effect of different life-history strategies on the relative prevalence of alternative male mating strategies. Specifically, we investigated the consequences of single or multiple matings per lifetime, mating strategy switching, cannibalism, resource stochasticity, and altruism towards relatives. We found that a combination of single (semelparous) matings, cannibalism and an absence of mating strategy changes in one lifetime led to a more strictly partitioned parameter space, with a reduced region where the two mating strategies co-exist in similar numbers. Explicitly including Hamilton's rule in simulations of the social system of a Cichlid led to an increase of dominant males, at the expense of both sneakers and dwarf males ("super-sneakers"). Our predictions provide general bounds on the viable ratios of alternative male mating strategies with different life-histories, and under possibly rapidly changing ecological situations.

Fisheries-induced evolution of alternative male life history tactics in Coho salmon.

Fisheries-induced evolution (FIE) can result when harvest imposes artificial selection on variation in heritable phenotypic traits. While there is evidence for FIE, it remains difficult to disentangle the contributions of within-generation demographic adjustment, phenotypic plasticity, and genetic adaption to observed changes in life history traits. We present evidence for FIE using dozens of Coho salmon (Oncorhynchus kisutch) populations in which males adopt one of two age-invariant, heritable life history tactics: most mature as large three-year-old "hooknose" and typically fight for spawning opportunities, while some mature as small two-year-old "jacks" and fertilize eggs through sneaking. The closure of a fishery targeting three-year-old fish provided an experimental test of the prediction that fishery-imposed selection against hooknose males drives an evolutionary increase in the proportion of males adopting the jack tactic. The data support the prediction: 43 of 46 populations had higher jack proportions during than after the fishery. The data further suggest that changes in jack proportion were not solely the result of demographic adjustments to harvest. We suggest that systems where fisheries differentially exploit phenotypically discrete, age-invariant life histories provide excellent opportunities for detecting FIE.

Sperm maturation and male tactic-specific differences in ejaculates in the plainfin midshipman fish Porichthys notatus.

Using the plainfin midshipman fish Porichthys notatus, a species with alternative reproductive tactics (ARTs), we investigated how sperm maturation shapes sperm competitive abilities. We compared sperm performance and morphology before and after final sperm maturation by sampling sperm from the testes and stripped ejaculates of guarders and sneakers. In accordance with sperm competition risk theory, ejaculates from sneaker males had three times as much sperm as ejaculates from guarder males and sneaker males produced faster swimming sperm than guarder males, but this was only the case after final sperm maturation had occurred. Additionally, fully mature sperm found in ejaculates had larger heads and midpieces than sperm found in the testes. These results emphasize the important role played by non-sperm components of an ejaculate in mediating sperm performance and potentially also morphology.

Recruitment variation disrupts the stability of alternative life histories in an exploited salmon population.

Males of many fish species exhibit alternative reproductive tactics, which can influence the maturation schedules, fishery productivity, and resilience to harvest of exploited populations. While alternative mating phenotypes can persist in stable equilibria through frequency-dependent selection, shifts in tactic frequencies have been observed and can have substantial consequences for fisheries. Here, we examine the dynamics of precocious sneaker males called "jacks" in a population of sockeye salmon (Oncorhynchus nerka) from Frazer Lake, Alaska. Jacks, which are of little commercial value due to their small body sizes, have recently been observed at unusually high levels in this stock, degrading the value of regional fisheries. To inform future strategies for managing the prevalence of jacks, we used long-term monitoring data to identify what regulates the frequencies of alternative male phenotypes in the population over time. Expression of the jack life history could not be explained by environmental factors expected to influence juvenile body condition and maturation probability. Instead, we found a strong positive association between the proportion of individuals maturing as jacks within a cohort and the prevalence of jacks among the males that sired that cohort. Moreover, due to differences in age-at-maturity between male phenotypes, and large interannual variability in recruitment strength, jacks from strong year-classes often spawn among older males from the weaker recruitments of earlier cohorts. Through such "cohort mismatches," which are amplified by size-selective harvest on older males, jacks frequently achieve substantial representation in the breeding population, and likely high total fertilizations. The repeated occurrence of these cohort mismatches appears to disrupt the stabilizing influence of frequency-dependent selection, allowing the prevalence of jacks to exceed what might be expected under equilibrium conditions. These results emphasize that the dynamics of alternative life histories can profoundly influence fishery performance and should be explicitly considered in the management of exploited populations.

Sneaker "jack" males outcompete dominant "hooknose" males under sperm competition in Chinook salmon (Oncorhynchus tshawytscha).

In a variety of taxa, males deploy alternative reproductive tactics to secure fertilizations. In many species, small "sneaker" males attempt to steal fertilizations while avoiding encounters with larger, more aggressive, dominant males. Sneaker males usually face a number of disadvantages, including reduced access to females and the higher likelihood that upon ejaculation, their sperm face competition from other males. Nevertheless, sneaker males represent an evolutionarily stable strategy under a wide range of conditions. Game theory suggests that sneaker males compensate for these disadvantages by investing disproportionately in spermatogenesis, by producing more sperm per unit body mass (the "fair raffle") and/or by producing higher quality sperm (the "loaded raffle"). Here, we test these models by competing sperm from sneaker "jack" males against sperm from dominant "hooknose" males in Chinook salmon. Using two complementary approaches, we reject the fair raffle in favor of the loaded raffle and estimate that jack males were ∼1.35 times as likely as hooknose males to fertilize eggs under controlled competitive conditions. Interestingly, the direction and magnitude of this skew in paternity shifted according to individual female egg donors, suggesting cryptic female choice could moderate the outcomes of sperm competition in this externally fertilizing species.