Pseudoasterophyllites cretaceus from the Cenomanian (Cretaceous) of the Czech Republic: A possible link between Chloranthaceae and Ceratophyllum

Pseudoasterophyllites cretaceus from the Cenomanian of Bohemia was recently recognized as an angiosperm by association with stamens containing monosulcate pollen of the Tucanopollis type. New material indicates that the stamens were borne in short spikes, with each stamen subtended by a bract, whereas the carpels were solitary and contained a single pendent, orthotropous ovule. We have investigated the phylogenetic position of Pseudoasterophyllites by including it in a morphological analysis of extant angiosperms using backbone constraint trees that represent the current range of hypotheses on relationships of the five mesangiosperm Glades. With a backbone tree in which Chloranthaceae are linked with magnoliids and Ceratophyllum with eudicots, the most parsimonious position of Pseudoasterophyllites is sister to Chloranthaceae, but a sister-group relationship to Ceratophyllum is only one step less parsimonious. With a backbone tree in which Chloranthaceae and Ceratophyllum form a Glade, Pseudoasterophyllites is sister to Ceratophyllum, based on derived features shared with both Chloranthaceae and Ceratophyllum plus solitary female flowers (as in Ceratophyllum). Similar results are obtained when Pseudoasterophyllites is added to the analysis with other fossils inferred to be related to Chloranthaceae and/or Ceratophyllum. If the plants that produced Tucanopollis pollen in the Barremian-Aptian of Africa South America are related to Pseudoasterophyllites, these results suggest that Chloranthaceae and Ceratophyllum are relicts of one of the most important early radiations of angiosperms, which included not only colonizers of disturbed terrestrial habitats but also halophytes and aquatics.