The palaeoclimatic utility of terrestrial biomarkers in marine sediments
Introduction
Biomarkers have become an increasingly common tool in the reconstruction of past biological assemblages, climate and environmental conditions. As with other palaeoenvironmental proxies, the application of biomarkers has been largely but not exclusively focussed on marine sedimentary sequences from which high-resolution, continuous and geographically widespread records can be collected from a significant portion of geologic history. In contrast, lacustrine or peat bog palaeo-records are largely limited to the Holocene (at high latitudes) or Pleistocene (at lower latitudes) with older deposits lacking the temporal and spatial occurrence necessary for many palaeoenvironmental investigations. This unfortunate bias means that for some regions and significant portions of Earth history, terrestrial environmental records are less developed than marine ones.
One increasingly used approach to obtain additional high-resolution terrestrial vegetation records is to take advantage of the significant quantity of terrestrial material in marine sediments. Rivers and wind transport large amounts of vascular plant organic material to the oceans. Although only little terrestrial organic matter is thought to be preserved in marine sediments (Hedges et al., 1997), it can often be visually distinguished from marine organic matter (e.g., as wood remains or pollen), serving as the basis for palynological investigations of vegetation change (e.g., Traverse, 1989). A particular advantage of this approach is that dispersed pollen and spore assemblages broadly reflect the regional composition of terrestrial plant communities (Traverse, 1989).
Similar to pollen, biomarkers of unambiguous terrestrial origin are also common in marine sediments and can even be dominant in coastal regions with high fluvial inputs (e.g., Amazon fan; Hinrichs and Rullkötter, 1997) or open ocean settings where aeolian terrestrial inputs are greater than pelagic ones (Pagani et al., 2000). Although higher plant biomarkers are generally not as diagnostic as pollen, some biomarker proxies for specific organisms (e.g., lignin monomers as indicators for gymnosperm vs. angiosperm inputs; Hedges and Parker, 1976) or continental temperature (n-alkane average chain length;Hinrichs et al., 1998) have been proposed. Studies utilizing the carbon isotopic composition of sedimentary higher plant biomarkers, typically n-alkanes, as an indicator of relative C3 and C4 plant inputs are more common. Here we review typical higher plant biomarkers and the controls on their carbon isotopic composition and discuss how such data can be used in palaeoclimate investigations. In addition, discussed are critical constraints on the interpretation of such data and potential future research directions.
Section snippets
n-Alkyl compounds
Long-chain n-alkyl compounds (Fig. 1) are major components of epicuticular waxes from vascular plant leaves (Eglinton et al., 1962, Eglinton and Hamilton, 1967). These compounds are relatively resistant to degradation, which makes them suitable for use as higher plant biomarkers (Cranwell, 1981), and include n-alkanes (I), n-alkanols (II), n-alkanoic acids (III) and wax esters (IV). All four compound classes have been identified in recent (Rieley et al., 1991) and ancient (e.g., Bird et al.,
Delivery mechanisms of terrestrial biomarkers to marine sediments
Accumulation of terrestrially derived material in marine sediments is due to aeolian and/or riverine transport or to the presence of aquatic higher plant communities (de Leeuw et al., 1995). Because vascular higher plants only occur widely in a limited number of marine settings (e.g., shallow shelves and lagoons), higher plant material in sediments from most marine settings is assumed to be predominately terrestrially derived. However, the proportion of such material derived from aeolian as
The carbon isotopic composition of terrestrial organic matter
The carbon component of most naturally occurring carbon-containing materials contains about 1.1% 13C and 98.9% 12C, and carbon isotopic compositions are expressed as a δ13C value, in units of per mil (‰), relative to the Vienna Pee Dee Belemnite (V-PDB) standard. Plants discriminate against 13C during photosynthesis, and the magnitude of this isotope effect reflects both the plant's metabolism and the growth environment. Typically, investigations of past vegetation change have exploited the
Conclusions: deriving climatic signals from higher plant biomarkers and future research directions
Higher plant biomarkers are abundant in marine sediments; they can be readily quantified and their carbon isotopic compositions determined, providing a powerful tool to develop high-resolution records of biotic inputs to marine settings. From these records, ecosystem structure on adjacent land masses can be inferred, taking into account variations in transport mechanisms and associated but poorly understood biases. Importantly, ecosystem structure is largely driven by climatic variables such as
Acknowledgements
We would like to thank a variety of current and past collaborators, including Jaap S. Sinninghe Damsté, Enno Schefuß, Marcel Kuypers, Mark Maslin, Virginia Ettwein, Richard Evershed and Kate Freeman, who in the course of research and stimulating discussion have contributed to this review. Kate Freeman is also thanked for a helpful and constructive review of this manuscript; an anonymous reviewer also offered critical comments that helped improve the final version. Finally, thanks and admiration
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