Stable carbon and oxygen isotopes in sub-fossil Sphagnum: Assessment of their applicability for palaeoclimatology
Introduction
The stable carbon and oxygen isotopic composition of plant organic matter has frequently been used for palaeoclimatic and palaeoenvironmental reconstructions. Wood is the preferred material for such studies which found empirical relationships between plant isotopic ratios and different climatic parameters, including temperature and relative humidity (e.g. DeNiro and Epstein, 1981, Edwards et al., 1985, Sternberg et al., 1986). Bulk plant tissue is composed of a number of chemical components that differ in their proportions between species. In dendroclimatological research, plant cellulose is usually used for stable isotope studies since this single chemical fraction allows linking the isotopic signal to a specific growth period in the annual cycle of wood formation (Helle and Schleser, 2004). Moreover, isolation of a single chemical component reduces problems associated with changes in the relative proportion of chemical compounds over time (Rinne et al., 2005).
Despite the great potential of peat bogs as climate archives, to date only few stable isotope studies focus on cellulose derived from continuously accumulated peat deposits. These archives provide records of environmental changes and vegetation dynamics over time, are widely distributed, and cover a large part of the earth's land surface often within human habitat (Charman, 2002). They can have relatively high accumulation rates and botanical fossils such as pollen and plant macrofossils are often well preserved due to the oxygen poor conditions in peat. Peat deposits are, therefore, an ideal archive for combining isotope-geochemical studies with palynological and palaeoecological approaches (Brenninkmeijer et al., 1982). Sphagnum is the most abundant peat forming genus in ombrotrophic bogs and fens in northern and western Europe and abundant in peat records throughout large time spans of the Holocene (Clymo, 1970). Recent studies have systematically investigated the relationship between climate parameters and cellulose isotope ratios in modern Sphagnum plants (e.g. Proctor et al., 1992, Aucour et al., 1996, Ménot and Burns, 2001, Ménot-Combes et al., 2002, Zanazzi and Mora, 2005, Skrzypek et al., 2007). Also a number of studies have attempted to use the isotopic compositions of sub-fossil peat material to infer environmental and/or climatic changes over time (Brenninkmeijer et al., 1982, Aucour et al., 1996, Turney et al., 1997, Hong et al., 2000, Hong et al., 2001, Jędrysek and Skrzypek, 2005). These studies, however, focus on bulk peat material and provided somewhat ambiguous results. Peat deposits are usually a mixture of partly decayed plant materials from different species. Use of bulk peat material, therefore, can be problematic due to contributions from multiple plants and fungi and/or selective degradation of isotopically distinct compounds (Pancost et al., 2003). A further problem must be seen in the fact that peat bogs are composed of a variety of different microenvironments that exhibit considerable variations in terms of bog wetness and moisture level. Bog vegetation, thus, typically consists of different Sphagnum species and sedges. Since different peatland species have been shown to be related to environmental- or climatic parameters in a different degree, analysing bulk peat material ignores the possible complicating factor of changes in Sphagnum species or species composition in a peat deposit (Ménot-Combes et al., 2002). Despite the great potential for environmental reconstructions, the application of sub-fossil Sphagnum cellulose of bulk organic matter from peat deposits as a valid palaeoclimate proxy, therefore, is still uncertain. An advanced contribution to test the potential of stable carbon isotopes of Sphagnum as palaeoclimate proxy was accomplished by Loader et al. (2007). The authors investigated inter- and intra-plant stable carbon isotopic variability of different physical components of individual modern Sphagnum plants. The results revealed a significant isotopic offset between branches and stems.
To explore the potential of stable isotopes of selected Sphagnum peat constituent for palaeoclimate research, we present results from a study of cellulose stable isotopes from selected Sphagnum constituents. We report the first combined δ13C and δ18O record of Sphagnum cellulose separately extracted from Sphagnum branches and stem sections manually separated from an approximately 4000 year old peat deposit. The main goals of this study are as follows. First, we sought to better understand the isotopic variability of different physical Sphagnum plant components in sub-fossil peat material. The manual separation of branches and stem sections prior to stable isotope analyses should allow determining possible stable isotope offsets between these different plant components. Second, we attempt to investigate the species composition of each individual sample down core to address the sensitivity of the stable isotope records to potential changes in Sphagnum assemblages. Since different Sphagnum species are assumed to be related to changes in bog ecology in a different degree, this approach also address the sensitivity of the records to potential changes in bog ecosystem variations. The aim of this study is, moreover, to inform sampling strategies for application of stable isotopes from Sphagnum plant material to the palaeorecord. An improved understanding of the isotope variability in sub-fossil Sphagnum plant components will significantly contribute to the evaluation of the potential of stable isotope signals from peat deposits and address to the reliability of such archives to record climatic and/or environmental information on the basis of stable isotope analysis.
Section snippets
Stable carbon isotope ratios of Sphagnum cellulose
Sphagnum does not have stomata and is therefore unable to regulate its uptake of CO2 through any physiological response. Photosynthetic cells are surrounded by large, dead so called “hyaline cells”, which form significant water reservoirs. Carbon dioxide enters the plant through pores in the hyaline cells and diffuses through the water that surrounds the chloroplast. The water filled hyaline cells build large barriers for carbon assimilation, because the diffusivity of CO2 is much lower in
Study site
In 2006 we took a core from a peat deposit located in the “Dürres Maar” (50°52'N, 6°53'E; 455 m a.s.l.), a small dry maar crater situated in the mountainous Westeifel Volcanic Field, Germany. The geological basement consists almost exclusively of Devonian shales. On this bedrock relatively poor non-calcarous soils developed in the surrounding of the maar crater without surficial inflow and outflow (Forst et al., 1997). The current mesotrophic bog has a diameter of 140–175 m with a surface area
Age-depth model, peat accumulation rate and Sphagnum peat stratigraphy
The 14C results which are given in Table 1 were used for the age-depth model of the “Dürres Maar” record (Fig. 3). All 14C results were calibrated and visualised using Oxcal (Bronk Ramsey, 1995, Bronk Ramsey, 2008). The bottom of the record was dated to ∼ 4000 cal. yr BP. Peat growth in the following two millennia was ~ 0.9 mm/yr. A considerable increase in the peat accumulation rate occurs at a depth of ~ 5.2 m (∼ 350 AD). Within the following ∼ 700 years about 3.5 m peat accumulated with a very
Implications for climate reconstructions
Our results displays that palaeoclimate interpretations based on cellulose stable isotope values derived from peat archives must take into account that there exist significant stable carbon and oxygen isotopic offset between cellulose from Sphagnum branches and stems. The results indicate that the use of bulk peat material without respect to the observed differences between the isotopic composition of Sphagnum branches and stems could clearly lead to erroneous interpretations of isotope
Conclusions
The stable carbon and oxygen isotope ratios of cellulose extracted separately from Sphagnum branches and stem sections from the “Dürres Maar” record reveal that there exist significant isotopic offset between these two different physical plant components. Isotopic differences are relatively small, however, both offsets exhibits a strong degree of correlation and are statistically highly significant. Our results indicate that the stable carbon isotopic offset between branches and stem sections
Acknowledgements
We thank Georg Heumann, Thomas Litt, Jörn Parplies, Nils Riedel and Heinz Vos for assistance in the field. Sample storage was afforded by Christa Lankes from the Institute of Crop Science and Resource Conservation, Bonn, Germany. The help of Stefanie Wagner during core sampling is greatly acknowledged. We wish to thank Bas van Geel for teaching us in Sphagnum species identification from single leaves on the basis of the characteristics of their hyaline and chlorophyllose cells. Support from
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