The morphology, systematics, and ecology of the extinct juglandaceous genus Polyptera are interpreted on the basis of infructescences, fruits, staminate catkins, pollen, and compound leaves from the Paleocene of Wyoming and Montana. The elongate infructescences of Polyptera manningii bear numerous helically arranged sessile fruits. The fruit is a pyramidal nut with a vascularized husk and a multilobed disk-like wing adaptive for wind dispersal. Associated staminate catkins bear numerous helically arranged florets, each with six or more stamens. Pollen from the anthers is triporate and isopolar, with ultrastructure diagnostic of Juglandaceae, but with a pattern of exinous thinning characteristic of the extinct dispersed pollen type Maceopolipollenites anellus (Nichols et Ott) comb. nov. The correlated leaves, Juglandiphyllites glabra (Brown ex Watt) comb. nov., are deciduous, pinnately compound, and have five to seven petiolulate finely serrate leaflets. Systematically, Polyptera shows relationships both with the Hicorieae and the Juglandeae. Occurrences in the Torrejonian and Tiffanian stages of the Paleocene place Polyptera as the oldest unequivocal record of Juglandaceae. Polyptera manningii typically occurs in low-diversity assemblages of 8-15 species, and usually is a dominant or subdominant constituent of floodplain assemblages. It may have been an early-successional colonizer.