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Testate Amoebae As Proxies for Mean Annual Water-Table Depth in Sphagnum-Dominated Peatlands of North America

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Testate Amoebae As Proxies for Mean Annual Water-Table Depth in Sphagnum-Dominated Peatlands of North America JOURNAL OF QUATERNARY SCIENCE (2008) 23(1) 43–57 Copyright ß 2007 John Wiley & Sons, Ltd. Published online 9 August 2007 in Wiley InterScience (www.interscience.wiley.com) DOI: 10.1002/jqs.1114 Testate amoebae as proxies for mean annual water-table depth in Sphagnum-dominated peatlands of North America ROBERT K. BOOTH* Department of Earth and Environmental Science, Lehigh University, Bethlehem, Pennsylvania, USA Booth, R. K. 2007. Testate amoebae as proxies for mean annual water-table depth in Sphagnum-dominated peatlands of North America. J. Quaternary Sci., Vol 23 pp. 43–57. ISSN 0267-8179. Received 6 September 2006; Revised 22 December 2006; Accepted 8 January 2007 ABSTRACT: Peatland-inhabiting testate amoebae are sensitive indicators of substrate-moisture conditions and have increasingly been used in palaeohydrological studies. However, to improve accuracy of testate-amoeba-based hydrological inferences, baseline ecological data on rare taxa, a larger geographic network of calibration sites, and incorporation of long-term estimates of water-table depth are needed. Species–environment relationships at 369 sites from 31 peatlands in eastern North America were investigated. Long-term estimates of water-table depth were obtained using the method of polyvinyl (PVC) tape-discolouration. Transfer functions were developed using a variety of models, and validated through jackknifing techniques and with an independent dataset where water-table depths were directly measured throughout the growing season. Results indicate that mean annual water-table depth can be inferred from testate amoeba assemblages with a mean error of 6 to 8 cm, although there is a slight systematic bias. All transfer function models performed similarly and produced similar reconstructions on a fossil sequence. In a preliminary effort towards development of a comprehensive North American calibration dataset, data from this study were combined with previous studies in Michigan and the Rocky Mountains (n ¼ 650). This combined dataset had slightly larger mean errors of prediction (8–9 cm) but includes data for several rare taxa. Copyright # 2007 John Wiley & Sons, Ltd. KEYWORDS: testate amoebae; Rhizopoda; peatlands; hydrology; palaeoclimate. Introduction events (Booth et al., 2006), monitoring and informing bog restoration efforts (Buttler et al., 1996; Davis and Wilkinson, 2004), and assessing the responses of terrestrial vegetation to Testate amoebae are a polyphyletic group of protozoa that past climate variability and change (Wilmhurst et al., 2002; produce decay-resistant and morphologically distinct shells, or Booth and Jackson, 2003; Booth et al., 2004). tests. Within oligotrophic peatlands (i.e. nutrient-poor) the Although the utility of testate amoebae as environmental distribution of taxa is primarily controlled by substrate mois- indicators has been clearly demonstrated, improving the pre- ture, making them sensitive proxies for past and ongoing cision and accuracy of transfer functions should increase the hydrological change (Charman, 2001). In the past decade, applicability of testate amoebae in addressing a wider range of transfer functions to infer water-table depth from testate environmental issues. Several problems with current cali- amoeba assemblages have been developed and validated in bration datasets need to be addressed to improve accuracy, various regions of the world (e.g. Charman, 1997; Charman and precision, and confidence. A primary concern has been that Warner, 1997; Woodland et al., 1998; Bobrov et al., 1999; most calibration studies have not collected environmental and Mitchell et al., 1999; Booth, 2002; Lamentowicz and Mitchell, community data at the same temporal scale. For example, most 2005; Charman et al., 2007; Payne et al., 2006). These training sets have been based on one-time measurements of transfer functions have been used to address a wide range of water-table depth (i.e. water-table depth measured only at time environmental and palaeoenvironmental issues. For example, of sampling) even though modern collections of testate amoeba they have played an important role in understanding temporal assemblages integrate the last several years of accumulation. relationships between vegetation and hydrology within raised Therefore, comparison of species–environment relationships bogs (McMullen et al., 2004), delimiting spatial and temporal among studies carried out at different times of year in different patterns of past centennial and sub-centennial scale drought regions has been problematic. Also, reconstructions can only be made along relative moisture gradients. The inclusion of annually averaged water-table depth measurements can * Correspondence to: R. K. Booth, Department of Earth and Environmental Science, Lehigh University, 31 Williams Drive, Bethlehem, PA 18015, USA. provide more meaningful reconstructions (Woodland et al., E-mail: [email protected] 1998), although long-term hydrological data are rarely 44 JOURNAL OF QUATERNARY SCIENCE available at sufficient spatial and temporal density and usually In this study, I attempt to address some of these problems by are prohibitably expensive to obtain. investigating relationships among testate amoeba assemblages The recent development and validation of a method of and hydrology from an array of 31 peatlands in eastern North estimating water-table depth using polyvinyl chloride (PVC) American, extending from the mid-continent to the east coast at tape may provide an inexpensive means of obtaining more mid-latitudes. I use the PVC-tape discolouration method to comparable and meaningful estimates of water-table depth for provide long-term estimates of water-table depth at the calibration studies (Belyea, 1999; Booth et al., 2005). The sampling locations, and develop transfer functions for water- PVC-tape discolouration method is based on the observation table depth. These transfer functions are validated using that PVC-tape changes colour when exposed to reducing jackknifing (leave-one-out) techniques (Birks, 1998). To test conditions (Belyea, 1999). Validation of the PVC-tape the assumption that the transfer functions, which are based on discolouration method in peatlands of northern Wisconsin PVC-discolouration estimates of water-table depth, predict indicates that the highest point of PVC-tape discolouration on mean annual water-table depths, I apply the transfer functions tape-lined stakes inserted into peatlands is a good approxi- to an independent dataset where mean annual water-table mation of the mid-summer or mean position of the water-table depths were directly measured. Water-table depths also were depth during the growing season (Booth et al., 2005). Other reconstructed for a sequence of fossil assemblages, allowing studies also have indicated strong relationships between comparison of reconstructions based on different transfer peatland water-table depth and the height of PVC-tape function models. Finally, I attempt to combine this newly discolouration (Bragazza, 1996; Belyea, 1999; Navra´tilova developed dataset with data from previous studies in Michigan and Ha´jek, 2005). However, precise relationships with water and the Rocky Mountains, in a preliminary effort toward the tables are not completely clear, with the position of development of a comprehensive North American calibration discolouration ranging from the mean height of the water dataset. table, to the depth that experiences inundation 70% of the time, to a few centimetres below the highest water levels (Bragazza, 1996; Belyea, 1999; Booth et al., 2005). The method may be problematic at fen sites with strongly fluctuating water tables Methods (Schnitchen et al., 2006). Another problem with many calibration datasets has been the relatively small geographic range of sampling. Even though Field methods many taxa are cosmopolitan in distribution and occupy similar ecological niches in different regions (e.g. Booth and Zygmunt, The sampled peatlands included raised bogs and kettle 2005), application of transfer functions developed from peatlands in continental and eastern North America (Fig. 1). assemblages in one region to fossil datasets from another All peatlands were Sphagnum-dominated, and characteristics region has sometimes been problematic (Charman et al., 2006). of the sites are shown in Table 1. Samples of testate amoeba Limited geographic sampling also may explain the lack of good communities were collected from within each peatland in an modern analogues for some fossil assemblages (Charman, effort to capture the range of hydrologic variability (e.g. 2001). Lack of good modern analogues appears to be primarily hummocks, hollows, pools). Methods of sampling were due to poor representation of several taxa in modern calibration modelled after previous studies (Booth, 2001, 2002). At each datasets (e.g. Difflugia pulex), but in some cases may be caused sampling site, 10 cm3 of Sphagnum moss was collected. The by taphonomic biases in highly decomposed peat (Wilmshurst samples consisted of the upper 3–5 cm of Sphagnum after the et al., 2003). Larger sampling networks and collection of upper 1–2 centimetres of moss were removed, which typically long-term environmental data should improve our under- corresponded to the brown area of the stem directly below the standing of testate amoeba ecology and biogeography, leading green portion. Vertical variation in assemblage composition to more accurate and precise environmental and palaeoenvir- occurs along the Sphagnum stem, and samples collected from onmental
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