Palaeoecology and evolution of marine hard substrate communities

Marine organisms have occupied hard substrates since the Archaean. Shells, rocks, wood and sedimentary hardgrounds offer relatively stable habitats compared to unconsolidated sediments, but the plants and animals which inhabit them must develop means to gain and defend this premium attachment space. Hard substrate communities are formed by organisms with a variety of strategies for adhering to and/or excavating the substrates they inhabit. While mobile grazers, organically attached and even soft-bodied organisms may leave evidence of their former presence in ancient hard substrate communities, a superior fossil record is left by sessile encrusters with mineralised skeletons and by borers which leave trace fossils. Furthermore, encrusters and borers are preserved in situ, retaining their spatial relationships to one another and to the substrate. Spatial competition, ecological succession, oriented growth, and differential utilisation of exposed vs. hidden substrate surfaces can all be observed or inferred. Hard substrate communities are thus excellent systems with which to study community evolution over hundreds of millions of years. Here we review the research on modern and ancient hard substrate communities, and point to some changes that have affected them over geological time scales. Such changes include a general increase in bioerosion of hard substrates, particularly carbonate surfaces, through the Phanerozoic. This is, at least in part, analogous to the infaunalisation trends seen in soft substrate communities. Encrusting forms show an increase in skeletalisation from the Palaeozoic into the Mesozoic and Cenozoic, which may be a response to increasing levels of predation. Hard substrate communities, considering borers and encrusters together, show a rough increase in tiering through the Phanerozoic which again parallels trends seen in soft substrate communities. This extensive review of the literature on living and fossil hard substrate organisms shows that many opportunities remain for large-scale studies of trends through time at the community and clade levels. Palaeontologists will especially benefit by closer integration of their work with that of neontologists, particularly in aspects of ecology such as larval recruitment, competition and succession.