Adaptive radiation, nonadaptive radiation, ecological speciation and nonecological speciation

Radiations of ecologically and morphologically differentiated sympatric species can exhibit the pattern of a burst of diversification, which might be produced by ecological divergence between populations, together with the acquisition of reproductive isolation (‘ecological speciation’). Here we suggest that this pattern could also arise if speciation precedes significant ecological differentiation (i.e. through geographical isolation and nonadaptive radiation). Subsequently, species ecologically differentiate and spread into sympatry. Alternative routes to producing ecologically differentiated sympatric species are difficult to detect in old radiations. However, nonadaptive radiations are common and might therefore regularly be responsible for currently ecologically differentiated sympatric species (e.g. among groups that are not susceptible to ecological speciation). Species evolving nonadaptively over long periods might eventually replace young, ecologically produced species.