Ulmus leaves and fruits from the Early–Middle Eocene of northwestern North America: systematics and implications for character evolution within Ulmaceae

Abstract

Leaves and fruits of Ulmus from the Early–Middle Eocene of British Columbia and Washington are assigned to two species. Ulmus okanaganensis sp.nov. is based on leaves attached to flowering and fruiting twigs and isolated leaves and fruits. Leaves display a polymorphism ranging from large leaves with compound teeth with a blunt apex to small ones with simple teeth resembling those of Zelkova. In extant Ulmus, sucker-shoot leaves, elongation-shoot leaves, and leaves on short annual branches often display a very similar polymorphism. In the fossil, flowers are arranged in fascicles having short pedicels. Fascicles are formed in the axils of leaves of current-year shoots and appear together with the leaves. This is uncommon in modern species of Ulmus, where leaves appear either in spring on previous-year shoots or in autumn in the axils of leaves of current-year shoots. Fruits of U. okanaganensis are samaras with extremely reduced or absent wings. Unwinged fruits of modern Ulmus are typically ciliate along the margin of the endocarp and the persistent styles but only a single fruit of U. okanaganensis has been found preserving hairs. The small, shallowly lobed perianth is situated below the endocarp. A second type of foliage is assigned to Ulmus chuchuanus (Berry) LaMotte. This foliage is wider than that of U. okanaganensis and has more densely spaced secondary veins. It also has characteristic compound teeth with primary and subsidiary teeth displaying conspicuously different orientations. Leaves of U. chuchuanus co-occur with a second type of fruit but have not been found in attachment. These fruits are larger than in U. okanaganensis, with a narrow wing, persistent styles, and a large and wide persistent perianth that tapers abruptly into the perianth tube. A cladistic analysis suggests that U. okanaganensis is nested within the subgenus Ulmus, which is a paraphyletic grade basal to the subgenus Oreoptelea. Ulmus chuchuanus foliage shows affinities to the subgenus Ulmus, while the associated fruits display affinities to the subgenus Oreoptelea.