Rensch's rule inverted--female-driven gigantism in nine-spined stickleback Pungitius pungitius

1. Allometric scaling of sexual size dimorphism (SSD) with body size is a commonplace occurrence in intraspecific or interspecific comparisons. Typically, SSD increases with body size when males, and decreases when females are the larger sex--a pattern known as Rensch's rule. Intraspecific studies of Rensch's rule in vertebrates are extremely scarce. 2. In an allometric SSD-body size relationship, the sex with the larger body size variation is the driver of size divergence whereas the other sex is following it owing to correlational selection. Hence, one can test which sex is responsible for the observed body size divergence within this framework. 3. Nine-spined stickleback (Pungitius pungitius) provides an excellent model to study intraspecific variation in SSD owing to the large interpopulation variation in mean body size. Using data on body size variation in 11 nine-spined stickleback populations covering the full known size range of the species, we investigated: (i) whether variation in SSD scales allometrically with mean body size across the populations; (ii) which sex is driving the allometric relationship and (iii) whether the observed pattern is likely to have a genetic component. In addition, we analysed the size dependency of female reproductive output. 4. We found strong support for an inverse of Rensch's rule: level of female-biased SSD increased with increasing mean size while females were the more variable sex. Results from a common garden experiment supported the pattern found in the wild. Females from giant populations had 2-3 times larger reproductive output than normal-sized females. 5. The fact that females were the more variable sex indicates that the evolution of gigantism in nine-spined sticklebacks is driven by females, and the 2-3 times larger reproductive output per clutch of giant vs. normal-sized females suggests fecundity selection to have an important role in it. Our results oppose the commonly held view that males drive the evolution of SSD as a result of sexual selection favouring larger males.