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Fagales
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Evolution

Nothofagaceae, or the southern beech family, are the oldest offshoot in the order. They appear to have diversified prior to the final separation of Gondwana in the late Cretaceous Period and now have various lineages in South America, Australia, and New Zealand. All the members of Nothofagus from New Guinea and New Caledonia are closely related. Fossils of Nothofagus in Australia indicate that the family was widespread in the Southern Hemisphere, except in Africa, where it apparently never occurred. Fagaceae, or the beech and oak family, represents the next diverging lineage, which diversified primarily in north temperate regions. Of the remaining families, Juglandaceae, Myricaceae, and Rhoipteleaceae appear closely related, as do Betulaceae, Casuarinaceae, and Ticodendraceae.

Since the pollen of Fagaceae and Betulaceae shows a marked similarity to the pollen of Normapolles from the Cretaceous Period, some think that they may have originated from within that assemblage. The earliest known fossils of Fagales are pollen specimens from the Campanian Stage (about 84 million to 71 million years ago) of the Upper Cretaceous, while the Normapolles group was still in existence, but no forms transitional to modern forms are known. Various other fossils date to the end of the Cretaceous.

Pollen characteristic of Alnus from the Santonian (from about 86 million to 84 million years ago) and Campanian stages of the Upper Cretaceous in Japan and North America represents the earliest record of Betulaceae. Fossil leaves similar to those of Betuleae have been found in deposits from about 70 million years ago and from the Paleocene Epoch in the early Paleogene Period (about 60 million years ago) but are not associated with the reproductive structures that would make identification certain. Reproductive structures of Alnus have been found in Paleocene deposits in Alaska and in the former Soviet Union. The oldest fossil inflorescences and leaves to be identified with certainty as Betula are found in Eocene (from about 56 million to 34 million years ago) deposits in British Columbia and eastern Oregon. Fruits resembling those of Corylus and Carpinus from several localities in North America, Greenland, and western Europe have been found. Definite Carpinus fruits with attached bracts are known from the late Eocene of Japan, and Ostrya is first definitely recognized in Oligocene (from about 34 million to 23 million years ago) deposits in western North America.

Despite the absence of fossils showing a clear transition, it is apparent that the catkins and the reduced and simplified flowers and inflorescences in Fagales represent the culmination of an evolutionary pathway leading to wind pollination from insect-pollinated ancestors. In members of most families, numerous staminate (pollen-producing) flowers are borne in pendulous catkins, and the pistillate flowers generally show modifications, such as elongate styles and enlarged or feathery stigmas, to receive wind-transported pollen grains. All the earliest fossils attributed to Fagales already manifest these changes, and so it is impossible to know exactly how the changes took place.

David E. Boufford Kenneth J. Sytsma