Lessons in modularity: the evolutionary ecology of colonial invertebrates

Benthic colonial invertebrates share with higher plants a modular construction and a sessile adult life. Both types of organism show parallel evolutionary responses to common selective forces, but in contrast to the long-established focus on plants, comparable study of colonial invertebrates has developed relatively recently, largely owing to the application of new techniques in image processing and molecular biology. Species whose life cycles are readily completed under laboratory conditions and whose colonies are easily propagated from cuttings provide powerful models for experimentally investigating fundamental evolutionary problems, including metabolic allometry, the manifestation of ageing and the origin of allorecognition systems. Free of the confounding influences of behavioural manipulation and costs of copulation, colonial invertebrates whose water-borne sperm fertilize retained eggs lend themselves well to the experimental study of cryptic female choice, sperm competition and sexual conflict. In these respects, it will be productive to adopt and extend theoretical frameworks developed for flowering plants to guide experimental investigation of modular animals. Since mate choice occurs at the cellular level in modular animals, reproductive isolation is uncorrelated with morphology and cryptic speciation is likely to be widespread.