Predator-prey interactions are commonly studied with an interest in determining the optimal strategy for prey. However, the implications of deviating from optimal strategy are often unclear. The present study considered these consequences by studying how the direction of an escape response affects the strategy of prey fish. We simulated these interactions with numerical and analytical mathematics and compared our predictions with measurements in zebrafish larvae (Danio rerio), which are preyed upon by adults of the same species. Consistent with existing theory, we treated the minimum distance between predator and prey as the strategic payoff that prey aim to maximize. We found that these interactions may be characterized by three strategic domains that are defined by the speed of predator relative to the prey. The "fast predator" domain occurs when the predator is more than an order of magnitude faster than the prey. The escape direction of the prey had only a small effect on the minimum distance under these conditions. For the "slow predator" domain, when the prey is faster than the predator, we found that differences in direction had no effect on the minimum distance for a broad range of escape angles. This was the regime in which zebrafish were found to operate. In contrast, the optimal escape angle offers a large benefit to the minimum distance in the intermediate strategic domain. Therefore, optimal strategy is most meaningful to prey fish when predators are faster than prey by less than a factor of 10. This demonstrates that the strategy of a prey animal does not matter under certain conditions that are created by the behavior of the predator.

Predation is a fundamental interaction between species, yet it is unclear what escape strategies are effective for prey survival. Classical theory proposes that prey should either escape in a direction that conforms to a performance optimum or that is random and therefore unpredictable. Here, we show that larval zebrafish (Danio rerio) instead use a mixed strategy that may be either random or directed. This was determined by testing classic theory with measurements of the escape direction in response to a predator robot. We found that prey consistently escaped in a direction contralateral to the robot when approached from the side of the prey's body. At such an orientation, the predator appeared in the prey's central visual field and the contralateral response was consistent with a model of strategy that maximizes the distance from the predator. By contrast, when the robot approached the rostral or caudal ends of the body, and appeared in the prey's peripheral vision, the escape showed an equal probability of a contralateral or ipsilateral direction. At this orientation, a contralateral response offered little strategic advantage. Therefore, zebrafish larvae adopt an escape strategy that maximizes distance from the threat when strategically beneficial and that is otherwise random. This sensory-mediated mixed strategy may be employed by a diversity of animals and offers a new paradigm for understanding the factors that govern prey survival.

Speciation is a multifaceted process that involves numerous aspects of the biological sciences and occurs for multiple reasons. Ecology plays a major role, including both abiotic and biotic factors. Whether populations experience similar or divergent ecological environments, they often adapt to local conditions through divergence in biomechanical traits. We investigate the role of biomechanics in speciation using fish predator-prey interactions, a primary driver of fitness for both predators and prey. We highlight specific groups of fishes, or specific species, that have been particularly valuable for understanding these dynamic interactions and offer the best opportunities for future studies that link genetic architecture to biomechanics and reproductive isolation (RI). In addition to emphasizing the key biomechanical techniques that will be instrumental, we also propose that the movement towards linking biomechanics and speciation will include (i) establishing the genetic basis of biomechanical traits, (ii) testing whether similar and divergent selection lead to biomechanical divergence, and (iii) testing whether/how biomechanical traits affect RI. Future investigations that examine speciation through the lens of biomechanics will propel our understanding of this key process.