Evidence on vegetative and inflorescence morphology of Chloranthaceae (Angiospermae) from the Early Cretaceous (middle–late Albian) of Spain

The near-basal angiosperm family Chloranthaceae (with four living genera) is prominently represented in the Early Cretaceous fossil record by pollen and flowers, but its leaves, other vegetative parts and inflorescences are less well known. Here, we report impressions of leaves, stems and inflorescences from the middle–late Albian Escucha Formation of Teruel Province (NE Spain) that help redress this imbalance and bring into clearer focus the morphological and systematic diversity of this key group in the early angiosperm radiation. We used parsimony analysis of a morphological dataset of living and fossil Chloranthaceae to evaluate the position of the fossils on a molecular-based tree of extant taxa. <i>Todziaphyllum elongatum</i> gen. et sp. nov., with festooned semicraspedodromous venation and asymmetrical chloranthoid teeth, is most parsimoniously placed on the stem lineage of the living genus <i>Hedyosmum</i>. Leaves of <i>Alcainea eklundiae</i> gen. et sp. nov., with festooned semicraspedodromous venation and symmetric teeth, occur attached to stems in opposite pairs at swollen nodes with sheathing leaf bases and interpetiolar stipules, along with compound spikes of flowers with a monosymmetric androecium of three stamens, each flower subtended by a bract. This represents the first time that fossil leaves, stems and inflorescences of Chloranthaceae have been found in organic connection. <i>Alcainea</i> may be sister to <i>Sarcandra</i>, <i>Chloranthus</i> or the clade consisting of both genera, like the mesofossil <i>Canrightiopsis</i>. Leaf Type A, with festooned craspedodromous venation and symmetrical teeth, may be related to <i>Ascarina</i>. These observations provide direct evidence for the distinctive vegetative and inflorescence morphology of Chloranthaceae in the Early Cretaceous. They are consistent with the level of diversification inferred from coeval chloranthaceous pollen and floral mesofossils, including lines nested in crown group Chloranthaceae but not in any of living genera, and exhibiting character combinations not retained in living Chloranthaceae. http://zoobank.org/urn:lsid:zoobank.org:pub:08B96BA0-3B27-48A1-9E8F-CB5811AE8C07