Information on fossil Nothofagus, and related genera, is reviewed based on pollen grains, leaves, woods, and flowers and fruits. The available data suggest that ancestors of Nothofagus and related families probably originated in West Gondwanaland. By the Santonian, Nothofagus was the only living genus already differentiated in the southern Hemisphere. Northern counterparts included several species in living genera of the Betulaceae, Myricaceae, and Ulmaceae. Rapid evolution and speciation of Nothofagus took place in the Eocene. After the pollen types and the main leaf forms were established in the Cretaceous and Eocene, respectively, the rate of morphological change was slow. More speciation was possible after the Miocene in New Guinea and also in other areas during the Plio-Pleistocene. Several lines of evidence support former suggestions of giving Nothofagus a higher taxonomic rank. Nothofagus Blume has frequently captured the interest of botanists and paleobotanists. It plays a dominant role in the temperate forests of the Southern Hemisphere, has economic importance as a timber source, and has an intriguing distribution as a vicariant of the other genera in Fagaceae. It comprises 34 species living in South America, Australia, New Zealand, New Caledonia, and New Guinea. It certainly lived in Antarctica, but perhaps never in South Africa. The Fagaceae are usually associated with the Betulaceae in the order Fagales (Thorne, 1983; Takhtajan, 1980), sometimes including also the Balanopaceae (Cronquist, 1968). Other related orders are Didymelales, Casuarinales, Myricales, and Hamamelidales, all included in the subclass Hamamelidae. Although this subclass has been subjected to considerable systematic rearrangement (Thorne, 1973), the taxa mentioned above were not very much affected. The Fagaceae have been segregated into three subfamilies since De Candolle (1864), and this classification has been supported by modern studies such as those of Forman (1966) and Hjelmqvist (1948) on the female cupule. The only objection has been the proposal to segregateNothofagus as a new family, made by Kuprianova (1965), on the basis of the pollen grains. More recent support for some kind of segregation was advanced by Crepet and Daghlian (1980), Smiley and Huggins (1981), Thorne (1983), Nixon (1984), and Jones (1984b). The classification of the species of Nothofagus into two sections and four series was suggested by van Steenis (1953) and Soepadmo (1972) on the basis of the female cupule and vegetative morphology. Recent information does not support fully such an arrangement (Elias, 1971; Philipson & Philipson, 1979; Hill, 1983), but no alternative system has been proposed. In recent years a growing body of information has accumulated about fossil materials of Nothofagus and related genera, shedding some light on its history and phylogenetical development. This information has not been fully used, and speculation about the evolution and paleobiogeography of the genera has been based mainly on the evidence of living plants (van Steenis, 1971; Darlington, 1965; Melville, 1973, 1981; Cracraft, 1975; Humphries, 1981, 1985; Heads, 1985). Therefore, it seems reasonable to review information concerning fossil forms. Modern and fossil data are reviewed first in this paper, followed by a discussion. The review of fossil data is arranged by type of fossil (pollen, leaves, woods, and flowers and fruits). In each case, a brief statement about their morphology in living plants is given. The discussion starts with the Cretaceous history of the genus, and also deals with the fossil record of related families. Then, the Paleogene account comprises the differentiation of species within the genera, so the morphology of the living ones is also considered. Finally, the Neogene part deals mainly with biogeography, and the information concerns primarily living plants. The fossil evidence was compiled by Wolfe (1973) and Muller (1981). When some information is given without menI Departamento de Ciencias Biol6gicas, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Intendente Guiraldes 2620, 1428 Buenos Aires, Repfiblica Argentina. Member of Consejo Nacional de Investigaciones Cientificas (CONICET). ANN. MISSOURI BOT. GARD. 73: 276-283. 1986. This content downloaded from 157.55.39.211 on Tue, 09 Aug 2016 05:15:09 UTC All use subject to http://about.jstor.org/terms 1986] ROMERO-NOTHOFAGUS EVOLUTION 277 tioning the source, it may be found in these reviews. However, in important cases the original publication is given. Other more recent papers, or ones not considered by Wolfe or Muller, are quoted as well. The references to paleogeography are based on Smith et al. (1981) and those to age are based on Harland et al. (1982).
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