Systematics and Leaf Architecture of the Gunneraceae

Cladistic and phenetic analyses of leaf and other morphological characters ofGunnera strongly support monophyly of the genus, with the Saxifragaceae s.str. as the closest sister group. This morphologically based phylogeny provides a more coherent understanding of the evolutionary history ofGunnera than do recent phylogenetic hypotheses based on genetic data sets with Myrothamnaceae as the sister group. Simple, crenate, palinactinodromously veined leaves lacking freely ending veinlets and tricolpate, tectate-perforate pollen with a reticulate exine indicate a shared ancestry. Within the genusGunnera all six traditionally recognized subgenera are monophyletic, as supported by leaf architectural apomorphies. The monotypic subgenusOstenigunnera is the sister group to the other five subgenera, which can be divided into two principal lineages. One lineage includes the subgeneraMilligania andMisandra, characterized by a prostate stoloniferous habit with small, low-rank leaves and exclusively unisexual flowers, whereas the other lineage includes the subgeneraPerpensum, Pseudo Gunnera, andPanke, all of which possess at least some hermaphroditic flowers and larger, high-rank leaves. When the phylogeny of the subgenera is considered in light of biogeography and the fossil record, a number of cladogenetic events can be explained by continental vicariance in the Late Cretaceous. The AfricanPerpensum became distinct from the other large-leafed lineage with the separation of the African continent ca. 90 Ma. The two small-leafed lineages, the subgeneraMilligania andMisandra, split with the separation of New Zealand from Western Gondwana, about 80 Ma.Pseudo-Gunnera became isolated fromPanke prior to this time, whenPanke fossils occur in North America.Gunnera probably arose out of an early herbaceous radiation of tricolpate eudicots having close affinity to the basal Saxifragaceae, espethe genusChrysosplenium.