Jellyfish life histories: role of polyps in forming and maintaining scyphomedusa populations.

Large population fluctuations of jellyfish occur over a variety of temporal scales, from weekly to seasonal, inter-annual and even decadal, with some regions of the world reported to be experiencing persistent seasonal bloom events. Recent jellyfish research has focussed on understanding the causes and consequences of these population changes, with the vast majority of studies considering the effect of changing environmental variables only on the pelagic medusa. But many of the bloom-forming species are members of the Scyphozoa with complex metagenic life cycles consisting of a sexually reproducing pelagic medusa and asexually reproducing benthic polyp. Recruitment success during the juvenile (planula, polyp and ephyrae) stages of the life cycle can have a major effect on the abundance of the adult (medusa) population, but until very recently, little was known about the ecology of the polyp or scyphistoma phase of the scyphozoan life cycle. The aim of this review is to synthesise the current state of knowledge of polyp ecology by examining (1) the recruitment and metamorphosis of planulae larvae into polyps, (2) survival and longevity of polyps, (3) expansion of polyp populations via asexual propagation and (4) strobilation and recruitment of ephyrae (juvenile medusae). Where possible, comparisons are made with the life histories of other bentho-pelagic marine invertebrates so that further inferences can be made. Differences between tropical and temperate species are highlighted and related to climate change, and populations of the same species (in particular Aurelia aurita) inhabiting different habitats within its geographic range are compared. The roles that polyps play in ensuring the long-term survival of jellyfish populations as well as in the formation of bloom populations are considered, and recommendations for future research are presented.