Size dependence of offspring production in isopods: a synthesis

In isopods, parental care takes the form of offspring brooding in marsupial pouches. Marsupial brooding was an important step towards the origin of terrestrial lifestyles among isopods, but its potential role in shaping isopod life histories remains unknown. It is here considered that marsupial brooding imposes costs and creates a temporary association between the survival of mothers and that of their offspring. Integrating findings from different life history models, we predicted that the effects of marsupial brooding set selective conditions for the continuation of growth after maturation, which leads to indeterminate growth, and the production of larger offspring by larger females. Based on this perspective, a study on the size dependence of offspring production in the woodlouse Porcellioscaber was performed and the generality of the results was tested by reviewing the literature on offspring production in other isopods. In P.scaber and almost all the other studied isopods, clutch size is positively related to female size. Such dependence is a necessary pre-condition for the evolution of indeterminate growth. The body mass of P.scaber differed six-fold between the largest and smallest brooding females, indicating a high potential for post-maturation growth. Our review showed that offspring size is a rarely studied trait in isopods and that it correlates negatively with offspring number but positively with female size in nearly half of the studied species. Our study of P.scaber revealed similar patterns, but the positive effect of female size on offspring size occurred only in smaller broods, and the negative relation between clutch size and offspring size occurred only in larger females. We conclude that the intraspecific patterns of offspring production in isopods agree with theoretical predictions regarding the role of offspring brooding in shaping the adaptive patterns of female investment in growth, reproduction, and the parental care provided to individual offspring.