Pelagic species diversity, biogeography, and evolution

Pelagic (open-ocean) species have enormous population sizes and broad, even global, distributions. These characteristics should damp rates of speciation in allopatric and vicariant evolutionary models since dispersal should swamp diverging populations and prevent divergence. Yet the fossil record suggests that rates of evolutionary turnover in pelagic organisms are often quite rapid, comparable to rates observed in much more highly fragmented terrestrial and shallow-marine environments. Furthermore, genetic and ecological studies increasingly suggest that species diversity is considerably higher in the pelagic realm than inferred from many morphological taxonomies.Zoogeographic evidence suggests that ranges of many pelagic groups are much more limited by their ability to maintain viable populations than by any inability to disperse past tectonic and hydrographic barriers to population exchange. Freely dispersing pelagic taxa resemble airborne spores or wind-dispersed seeds that can drift almost anywhere but complete the entire life cycle only in favorable habitats. It seems likely that vicariant and allopatric models for speciation are far less important in pelagic evolution than sympatric or parapatric speciation in which dispersal is not limiting. Nevertheless, speciation can be quite rapid and involve cladogenesis even in cases where morphological data suggest gradual species transitions. Indeed, recent paleoecological and molecular studies increasingly suggest that classic examples of “phyletic gradualism” involve multiple, cryptic speciation events.Paleoceanographic and climatic change seem to influence rates of turnover by modifying surface water masses and environmental gradients between them to create new habitats rather than by preventing dispersal. Changes in the vertical structure and seasonality of water masses may be particularly important since these can lead to changes in the depth and timing of reproduction. Long-distance dispersal may actually promote evolution by regularly carrying variants of a species across major oceanic fronts and exposing them to very different selection pressures than occur in their home range. High dispersal in pelagic taxa also implies that extinction should be difficult to achieve except though global perturbations that prevent populations from reestablishing themselves following local extinction. High rates of extinction in some pelagic groups suggests either that global perturbations are common, or that the species are much more narrowly adapted than we would infer from current taxonomies.