Faster is not always better: selection on growth rate fluctuates across life history and environments.

Growth rate is increasingly recognized as a key life-history trait that may affect fitness directly rather than evolve as a by-product of selection on size or age. An ongoing challenge is to explain the abundant levels of phenotypic and genetic variation in growth rates often seen in natural populations, despite what is expected to be consistently strong selection on this trait. Such a paradox suggests limits to how contemporary growth rates evolve. We explored limits arising from variation in selection, based on selection differentials for age-specific growth rates expressed under different ecological conditions. We present results from a field experiment that measured growth rates and reproductive output in wild individuals of a colonial marine invertebrate (Hippopodina iririkiensis), replicated within and across the natural range of succession in its local community. Colony growth rates varied phenotypically throughout this range, but not all such variation was available for selection, nor was it always targeted by selection as expected. While the maintenance of both phenotypic and genetic variation in growth rate is often attributed to costs of growing rapidly, our study highlights the potential for fluctuating selection pressures throughout the life history and across environments to play an important role in this process.