Do fossil plants signal palaeoatmospheric carbon dioxide concentration in the geological past?

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Abstract
Fossil, subfossil, and herbarium leaves have been shown to provide a morphological signal of the atmospheric carbon dioxide environment in which they developed by means of their stomatal density and index. An inverse relationship between stomatal density/index and atmospheric carbon dioxide concentration has been documented for all the studies to date concerning fossil and subfossil material. Furthermore, this relationship has been demonstrated experimentally by growing plants under elevated and reducedcarbon dioxide concentrations. To date, the mechanism that controls the stomatal density response to atmospheric carbon dioxide concentration remains unknown. However, stomatal parameters of fossil plants have been successfully used as a proxy indicator of palaeo–carbon dioxide levels. This paper presents new estimates of palaeo–atmospheric carbon dioxide concentrations for the Middle Eocene (Lutetian), based on the stomatal ratios of fossil Lauraceae species from Bournemouth in England. Estimates of atmospheric carbon dioxide concentrations derived from stomatal data from plants of the Early Devonian, Late Carboniferous, Early Permian and Middle Jurassic ages are reviewed in the light of new data. Semi–quantitative palaeo–carbon dioxide estimates based on the stomatal ratio (a ratio of the stomatal index of a fossil plant to that of a selected nearest living equivalent) have in the past relied on the use of a Carboniferous standard. The application of a new standard based on the present–day carbon dioxide level is reported here for comparison. The resultant ranges of palaeo–carbon dioxide estimates made from standardized fossil stomatal ratio data are in good agreement with both carbon isotopic data from terrestrial and marine sources and long–term carbon cycle modelling estimates for all the time periods studied. These data indicate elevated atmospheric carbon dioxide concentrations during the Early Devonian, Middle Jurassic and Middle Eocene, and reduced concentrations during the Late Carboniferous and Early Permian. Such data are important in demonstrating the long–term responses of plants to changing carbon dioxide concentrations and in contributing to the database needed for general circulation model climatic analogues.