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1135 Devore.Vp Floristic composition and comparison of middle Eocene to late Eocene and Oligocene floras in North America MELANIE L. DEVORE & KATHLEEN B. PIGG In comparison to the early and middle Eocene, the late Eocene and particularly the Oligocene floral record is sparse in North America. Changing tectonic, environmental and climatic conditions during these times resulted in the develop- ment of fewer depositional systems favorable for fossil preservation. Floras are known from the Southeast, the Pacific Northwest and the Rocky Mountains. Each area has a distinct geological history that shaped both the vegetation adjacent to sites of deposition as well as the depositional environments themselves. The floristic change from middle to late Eocene, and then to Oligocene reflects a changing paleoclimate from the thermal maximum to cooler and drier condi- tions in the late Paleogene. In the present paper, major middle, and then late Eocene and finally Oligocene floras of North America are summarized, with an emphasis on their regional geology, depositional setting, paleoclimate and significant floral elements. The North American occurrences of coryphoid palms (Sabal) and cycads are reviewed in relationship to their biogeographic history. Finally, we suggest several directions for future research that will further illuminate the floristic changes from middle, to late Eocene and Oligocene that occurred in North America. • Key words: Claiborne, Eocene/Oligocene transition, Florissant, Okanogan Highlands, Tertiary floras. DEVORE, M.L. &. PIGG, K.B. 2010. Floristic composition and comparison of middle Eocene to late Eocene and Oligocene floras in North America. Bulletin of Geosciences 85(1), 111–134 (1 figure, 2 tables). Czech Geological Sur- vey, Prague. ISSN 1214-1119. Manuscript received April 7, 2009; accepted in revised form October 5, 2009; published online January 8, 2010; issued xxxx xx, 2010. Melanie L. DeVore (corresponding author), Georgia College & State University, 135 Herty Hall, Milledgeville, GA 31062-0001, USA; [email protected] • Kathleen B. Pigg, School of Life Sciences, Arizona State University, PO Box 874501, Tempe AZ 85287-4501 USA; [email protected] Assemblages of plant-bearing fossiliferous sedimentary Eocene toward the cooler and drier conditions that continue rocks of the late Eocene and especially Oligocene ages are to the present day. Floral response to climate change has rare in North America (Wing 1987). There are several rea- been studied by both the Nearest Living Relative method sons why this is so. First, the extensive Paleogene fluvial (NLR) and leaf physiognomy. The NLR uses the distribu- plain covering the region once occupied by the Cretaceous tion of closely related living plants to infer climate in fossil Intercontinental Seaway became diminished, limiting the floras (e.g., Axelrod 1957, 1966a). This method is valuable, areas with the appropriate conditions necessary for plant however fossil forms may not have occupied identical preservation. Secondly, climate became drier, leading to fe- niches or had equal physiological tolerances as their current wer environments of deposition available for preserving me- relatives. Leaf physiognomic methods such as Leaf Margin gafossils. Thirdly, the ash-producing volcanism responsible Analysis (LMA), Leaf Area (LA) and Climate-Leaf Analy- for blocking drainage systems in the western Rocky Moun- sis Multivariate Program (CLAMP) are based on assem- tains to create depositional basins decreased during the Oli- blages of extant leaves characterized by characters of size gocene (Graham 1999). Thus the combination of changing and shape correlated to their occurrence in modern environ- geomorphology, drying climate and less volcanic activity all ments. These taxon-free approaches provide a means inde- contributed to the decreased number and extent of fossil pendent of systematic affinities for estimating climate plant beds of this age. As a result, the transition from the ro- (Wolfe 1987, Graham 1999). Easily quantifiable leaf physi- bust fossil records of the early and middle Eocene into the ognomy techniques have become a powerful tool for provid- late Eocene and Oligocene is only broadly understood. ing numerical data that can be compared readily with other Despite the limited late Eocene and Oligocene plant bed proxies, such as carbon isotopes. These two types of meth- exposures in North America, this is an important time period ods often are used in combination to establish estimates of to study for several reasons. First, these floras mark a major paleoclimate and paleoelevation parameters (e.g., Green- shift in climate from the warmer and more humid middle wood et al. 2005). DOI 10.3140/bull.geosci.1135 111 Bulletin of Geosciences Vol. 85, 1, 2009 A second major area of interest is the origin and diversi- Determination of precise ages and depositional environ- fication of major taxonomic groups. As taxon-free meth- ments within the Claiborne Formation is limited because ods have become de rigueur for climate reconstruction, tra- stratigraphic units are difficult to correlate. The most ex- ditional systematic studies of fossil plants have shifted tensively studied plants of the Claiborne occur in localized toward novel applications that relate to molecular clay pits (e.g., Puryear, Warman, Lamkin). These outcrops phylogenetics and biogeography (Manchester 1999, are small, discontinuous units not found in conjunction Manos & Stanford 2001, Tiffney & Manchester 2001, with marine deposits or other biostratigraphically informa- Manos et al. 2007, Manchester et al. 2009). Fossil occur- tive sediments. In contrast to localities in the West that are rences act as an independent proxies for dating nodes of di- commonly associated with intense orogenic activity during vergence for molecular and/or combined data sets. They the Cenozoic, the Mississippian Embayment is on the pas- also provide a means for examining character evolution, sive margin of the North American plate and typically and broad responses of lineages to environmental change, lacks datable ashes and other igneous units. Since radio- as well as demonstrating past distributions. The middle to metric dating is not available, floras in the region are corre- late Eocene and then Oligocene floras mark important lated primarily by pollen zonation (Elsik & Dilcher 1974, chapters in the evolutionary and biogeographic history of Potter & Dilcher 1980, Table 1). major plant groups as they adapt, diversify, and wax or The depositional environments of the Claiborne For- wane in the fossil record. mation have been interpreted in several ways. Originally, In this contribution the major floras of middle to late Berry (1916) considered the Claiborne to be the result of a Eocene and Oligocene ages in North America are re- series of fluvial and lacustrine deposits in back-beach areas viewed, with a brief description of the geological setting, of the Gulf of Mexico. Dilcher (1971) attributed the clay depositional environment, and paleoclimate of each, fol- lenses of Henry and Weakley Counties of Tennessee to de- lowed by a description of significant floral elements (Ta- posits in abandoned river channels of oxbow lakes occupy- ble 1). This study presents more detail about several floras ing a low-lying floodplain. In one study Moore et al. that to date have not been comprehensively summarized. (2001) suggested that the section exposed at the Wilbanks Fossil occurrences of cycads and coryphoid palms are re- Clay Pit in western Tennessee may have been deposited in viewed for North America in relationship to their one to several seasons. biogeographic history. Lastly, directions for future re- Multiple overlapping clay lenses with truncated mar- search are suggested that will further illuminate an under- gins, cut-and-fills, and cross-bedded sandstones containing standing of the late Eocene and Oligocene floras of North clay rip-ups were found in northeastern outcrops of the America. Claiborne. Some of these plant-bearing clays are inter- preted as being deposited within a braided stream system (Moore et al. 2003). Based on the three-dimensional struc- Overview of the floras ture of several clay lenses characterized by distinct lower contacts and fine laminated bedding, Moore et al. (2003) Middle Eocene to Early Oligocene in the later proposed that rates of infilling probably occurred in Southeast Mississippi Embayment these deposits over a 1,000–2,000 year period. Future stud- ies, particularly within the context of basin analysis, may One of the best glimpses of the late Eocene and Oligocene help to clarify the apparent contrasting modes of deposition floras comes from southeastern North America in the re- within the Claiborne. gion of the Mississippian Embayment. These floras were The Mean Annual Temperature (MAT) for Claiborne initially surveyed by Berry (1916, 1922, 1924, 1925, 1930, has been estimated variously as between 20–28 °C or 1941) who, over the course of 25 years, described more 22–30 °C (Graham 1999). Dilcher (1973) suggested that than 340 species of Eocene and Oligocene fossil plants, the climate was similar to coastal Louisiana (21 °C) or to many from this floristic region (Pigg & DeVore 2007). southern peninsular Florida (24 °C). These estimates, Many of Berry’s identifications have been reinterpreted, so with MAT above 20 °C, indicate a megathermal flora subsequent papers should be consulted. A sustained effort (Wolfe 1987). The Claiborne is characterized as a dry in studying Mississippian Embayment floras
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