Lifetime selection on a hypoallometric size trait in the spotted hyena

Size-related traits are common targets of natural selection, yet there is a relative paucity of data on selection among mammals, particularly from studies measuring lifetime reproductive success (LRS). We present the first phenotypic selection analysis using LRS on size-related traits in a large terrestrial carnivore, the spotted hyena, which displays a rare pattern of female-biased sexual size dimorphism (SSD). Using path analysis, we investigate the operation of selection to address hypotheses proposed to explain SSD in spotted hyenas. Ideal size measures are elusive, and allometric variation often obfuscates interpretation of size proxies. We adopt a novel approach integrating two common methods of assessing size, and demonstrate lifetime selection on size-related traits that scale hypoallometrically with overall body size. Our data support selection on hypoallometric traits in hyenas, but not on traits exhibiting isometric or hyperallometric scaling relationships, or on commonly used measures of overall body size. Our results represent the first estimate of lifetime selection on a large carnivore, and suggest a possible route for maintenance of female-biased SSD in spotted hyenas. Finally, our results highlight the importance of choosing appropriate measures when estimating animal body size, and suggest caution in interpreting selection on size-related traits as selection on size itself.