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We present the oldest known occurrences of crown-group Trochodendraceae based on new material from the Palaeocene of Wyoming, USA. Two genera are recognized, Trochodendron and Eotrochion gen. nov. The fossil fruit of Trochodendron infernense sp. nov. is represented by a pedicellate, apically dehiscent capsular fruit composed of nine follicle-like units, each bearing a persistent convex style. The basal part is ornamented with numerous raised stamen scars. From the same deposits, Eotrochion is represented by infructescences, fruits and associated leaves. The infructescences are racemes of numerous apically dehiscent capsules, each with c. 14–16 styles, each with an underlying nectary and receptacles lacking stamen scars, but possessing a prominent perianth scar. A phylogenetic assessment of the modern species, plus representatives of four extinct genera of fossil Trochodendraceae based on available morphological characters, yields a favoured topology of Trochodendron(Eotrochion(Concavistylon kvacekii(C. wehrii (Pentacentron, Tetracentron)))). A parsimony analysis of currently available characters indicates that C. wehrii renders Concavistylon non-monophyletic. Accordingly, we transfer it to Paraconcavistylon gen. nov., characterized by pendent, rather than erect infructescences. We also reconsider the extinct Nordenskioeldia (Late Cretaceous to Miocene), the prior placement of which in Trochodendraceae has been challenged, and we consider it to fall outside the crown group of the family.

Morphology, anatomy, phylogenetics and distribution of fossil and extant Trochodendraceae in the Northern Hemisphere