Sperm competition and the evolution of alternative reproductive tactics in the swordtail Xiphophorus nigrensis (Poeciliidae)
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Darwin identified sexual selection as an important evolutionary process resulting from differences among males in their ability to secure mates. In the latter half of the 20th century, it became apparent that females often mate with multiple partners within the same reproductive cycle, leading to the overlap of ejaculates from multiple males and sperm competition for the fertilization of the eggs. Here, I examine how sperm competition has influenced the evolution of Xiphophorus nigrensis, an internally fertilized, livebearing fish with alternative male mating tactics that are dependent upon male size. I find that variation in male tactic is correlated with variation in traits relevant to sperm competition: small males that sneak copulations produce ejaculates with a greater proportion of fertilization-capable sperm (sperm viability) and sperm that is longer-lived following activation compared to large males that court females. Sperm morphology is also divergent between tactics and correlated with sperm performance: smaller males have larger midpieces and midpiece size is positively correlated with sperm velocity and longevity. Social environment also affects ejaculate quality, with sperm velocity rapidly increasing when a small male is exposed to another small male compared to when he is exposed to a large male. Large male ejaculates were invariant across social environments. Next, I demonstrate experimentally that the observed variation in sperm quality has important consequences for the outcome of sperm competition. Males with superior sperm viability sire more offspring, while sperm velocity is negatively associated with sperm competitive ability when sperm are stored within the female prior to fertilization. Finally, I show that sperm competition is likely to have important effects on male reproductive success in the wild by characterizing the genetic mating system of X. nigrensis. I find that 61% of females collected from the Nacimiento Río Choy produce offspring sired by 2-4 males. Paternity is strongly skewed among sires, with an average of 70% of offspring sired by one of the males represented in the brood. These studies illustrate sperm competition can have potent effects on the evolution of animals.