Phylogenies and angiosperm diversification

Abstract

Approaches to patterns of diversification based on counting taxa at a given rank can be misleading, even when all taxa are monophyletic. Such “rank-based” approaches are unable to reflect a hierarchy of evolutionary events because taxa of the same rank cannot be nested within one another. Phylogenetic trees specify an order of origination of characters and clades and can therefore be used in some cases to test hypotheses on causal relationships between characters and changes in diversity. “Tree-thinking” also clarifies discussions of the age of groups, by distinguishing between splitting of the stem-lineage from its sister group and splitting of the crown-group into extant clades.Cladistic evidence thatPentoxylon, Bennettitales, and Gnetales are the sister group of angiosperms implies that the angiosperm line (angiophytes) existed by the Late Triassic. The presence of primitive members of five basic angiosperm clades indicates that the crown-group (angiosperms) had begun to diversify by the mid-Early Cretaceous (Barremian-Aptian), but not necessarily much earlier. The greatest unresolved issue raised by cladistic analyses concerns the fact that the angiosperm tree can be rooted in two almost equally parsimonious positions. Trees rooted near Magnoliales (among “woody magnoliids”) suggest that the angiosperm radiation may have been triggered by the origin of intrinsic traits, e.g., a fast-growing, rhizomatous habit in the paleoherb and eudicot subgroup. However, trees rooted among paleoherbs, which are favored by rRNA data, imply that these traits are basic for angiosperms as a whole. This could mean that the crown-group originated not long before its radiation, or, if it did originate earlier, that its radiation was delayed due to extrinsic factors. Such factors could be a trend from environmental homogeneity and stability in the Jurassic to renewed tectonic activity and disturbance in the Early Cretaceous. Potentially relevant pre-Cretaceous fossils cannot be placed with confidence, but may be located along the stem-lineage (stem angiophytes); their generally paleoherb-like features favor the paleoherb rooting. The history of angiophytes may parallel that of Gnetales: some diversification of the stem-lineage in the Late Triassic, near disappearance in the Jurassic, and vigorous radiation of the crown-group in the Early Cretaceous.