Modes and mechanisms of speciation in pteridophytes : Implications of contrasting patterns in ferns representing temperate and tropical habitats

Discovering how biological diversification results in species is one of the primary challenges facing evolutionary biologists. In the ferns, evidence indicates that dissimilar speciation modes and mechanisms may differentiate some temperate and tropical groups. The Polypodium sibiricum group contains three related diploid species that all inhabit rock outcrops in temperate forests. Although differing lettle in gross leaf morphology and joined by the distinctive morphological synapomorphy of sporangiasters, these three species have an average interspecific genetic identity developed from isozymic com-parisons of only 0.460. A likely mode of speciation is that periodic glaciation pushed Po. sibiricum populations south and, with the retreat of the glaciers, southern populations persisted, evolved diagnostic traits, and ultimately erected postzygotic barriers to interbreeding. This hypothesis follows a classic allopatric speciation model and interspecific distinctions may have been reinforced through contact mediated by subsequent ice ages. In contrast, a monophyletic group of four diploid, epiphytic Pleopeltis species centered in Mexico has an isozymically-determined average interspecific genetic identity value of 0.849. In spite of this high value, these species show greater morphological discrimination than do the Polypodium species. Although the species ranges overlap, they appear to occupy ecologically discrete habitats. These Pleopeltis species may have originated through adaptation to different ecological zones and developed individual morphologies in the process. The high interspecific genetic identity values among the Pleopeltis species suggest a relatively recent and/or rapid process. These hypotheses should be tested by further biosystematic investigations and the discovery of additional monophyletic assemblages with similar patterns of speciation.