Article Processing Charges for This Open-Access Richness in Comparison to the Pre-LGM Deposits
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Biogeosciences, 14, 575–596, 2017 www.biogeosciences.net/14/575/2017/ doi:10.5194/bg-14-575-2017 © Author(s) 2017. CC Attribution 3.0 License. Sedimentary ancient DNA and pollen reveal the composition of plant organic matter in Late Quaternary permafrost sediments of the Buor Khaya Peninsula (north-eastern Siberia) Heike Hildegard Zimmermann1,2, Elena Raschke1,3, Laura Saskia Epp1, Kathleen Rosmarie Stoof-Leichsenring1, Georg Schwamborn1, Lutz Schirrmeister1, Pier Paul Overduin1, and Ulrike Herzschuh1,2 1Alfred Wegener Institute Helmholtz Centre for Polar and Marine Research, Periglacial Research Unit, Telegrafenberg A43, 14473 Potsdam, Germany 2Institute of Earth and Environmental Sciences, University of Potsdam, Karl-Liebknecht-Str. 24-25, 14476 Potsdam-Golm, Germany 3Arctic and Antarctic Research Institute, Bering St. 38, 199397 St. Petersburg, Russia Correspondence to: Heike Hildegard Zimmermann ([email protected]) and Ulrike Herzschuh ([email protected]) Received: 9 September 2016 – Discussion started: 27 September 2016 Revised: 26 December 2016 – Accepted: 11 January 2017 – Published: 7 February 2017 Abstract. Organic matter deposited in ancient, ice-rich per- decrease in taxonomic richness was less pronounced com- mafrost sediments is vulnerable to climate change and may pared to sedaDNA. Our results show the advantages of using contribute to the future release of greenhouse gases; it is sedaDNA in combination with palynological analyses when thus important to get a better characterization of the plant macrofossils are rarely preserved. The high resolution of the organic matter within such sediments. From a Late Quater- sedaDNA record provides a detailed picture of the taxonomic nary permafrost sediment core from the Buor Khaya Penin- composition of plant-derived organic matter throughout the sula, we analysed plant-derived sedimentary ancient DNA core, and palynological analyses prove valuable by allowing (sedaDNA) to identify the taxonomic composition of plant for inferences of regional environmental conditions. organic matter, and undertook palynological analysis to as- sess the environmental conditions during deposition. Using sedaDNA, we identified 154 taxa and from pollen and non- pollen palynomorphs we identified 83 taxa. In the deposits 1 Introduction dated between 54 and 51 kyr BP, sedaDNA records a diverse low-centred polygon plant community including recurring Decomposition of soil organic carbon from ancient per- aquatic pond vegetation while from the pollen record we in- mafrost deposits may augment ongoing global warming, but fer terrestrial open-land vegetation with relatively dry envi- how intensely will depend on the character of the organic ronmental conditions at a regional scale. A fluctuating dom- matter. Most soil organic carbon in permafrost was deposited inance of either terrestrial or swamp and aquatic taxa in both during the Late Pleistocene and Holocene (cf. Schirrmeister proxies allowed the local hydrological development of the et al., 2011a). Throughout the last glacial (Marine Isotope polygon to be traced. In deposits dated between 11.4 and Stages 4 to 2, ca. 71–10.5 kyr BP), much of north-eastern 9.7 kyr BP (13.4–11.1 cal kyr BP), sedaDNA shows a taxo- Siberia was non-glaciated (Hubberten et al., 2004). The re- nomic turnover to moist shrub tundra and a lower taxo- gion was exposed to extremely cold conditions, which re- nomic richness compared to the older samples. Pollen also sulted in the formation of deep permafrost with an estimated records a shrub tundra community, mostly seen as changes thickness of 500 m (Duchkov et al., 2014) and in lowlands of in relative proportions of the most dominant taxa, while a polygonal tundra environments. Ice-wedge polygons devel- oped after many cycles of wintertime frost cracking and sub- Published by Copernicus Publications on behalf of the European Geosciences Union. 576 H. H. Zimmermann et al.: SedaDNA and pollen reveal plant organic matter composition sequent springtime meltwater infiltration into these cracks. amounts to quantify its composition, but in arctic treeless Ice wedges create ridges surrounding polygonal depressions ecosystems it originates from across a variable extent from (e.g. Lachenbruch, 1962; Leffingwell, 1915; Minke et al., extra-local to local sources (Birks, 2001; van der Knaap, 2007). The cold and water-saturated conditions in the de- 1987) and taxonomic resolution is mostly limited to genus- pressions caused the accumulation of organic matter in soil and family level (Beug, 2004; Moore et al., 1991). A com- horizons, as decomposition rates were low (Davidson and paratively new proxy is the analysis of sedimentary ancient Janssens, 2006; Hugelius et al., 2014). Over time, ice- and DNA (sedaDNA), which originates from disseminated ma- organic-matter-rich permafrost was formed (Schirrmeister et terial within sediments (Haile et al., 2009; Rawlence et al., al., 2011b). These Late Pleistocene deposits are called Ice 2014). Its application to permafrost sediments was intro- Complex or Yedoma and are estimated to store 83 ± 12 Pg duced by Willerslev et al. (2003) who reported the successful organic carbon (Hugelius et al., 2014), which accounts for retrieval of plant DNA as old as 300 000 to 400 000 years. more than 10 % of the total organic carbon pool in permafrost Since then, several studies prove that DNA is exceptionally globally. well preserved in permafrost sedimentary archives with re- Permafrost is susceptible to future climate-change- spect to vascular plants (Willerslev et al., 2003), bryophytes induced increases in ground temperatures, which can lead to (Epp et al., 2012), fungi (Bellemain et al., 2013; Epp et active layer deepening (Romanovsky et al., 2010) and ther- al., 2012; Lydolph et al., 2005), bacteria (e.g. Wagner et moerosion (Grosse et al., 2011). When permafrost sections al., 2007; Willerslev et al., 2004b), invertebrates (Epp et al., thaw, microbial-driven decomposition rates increase and re- 2012), birds (Epp et al., 2012), and mammals (e.g. Arnold et lease climate-relevant greenhouse gases that further enhance al., 2011; Haile et al., 2009; Willerslev et al., 2003, 2014). climate warming in a positive feedback loop (Knoblauch The constantly cool temperatures of permafrost lead to re- et al., 2013; Wagner et al., 2007). To improve assessments duced microbial and enzymatic degradation (Levy-Booth et of the potential greenhouse gas release from organic mat- al., 2007) and limited hydrolytic damage, since up to 97 % of ter, knowledge about its composition is of great relevance the water is frozen (Willerslev et al., 2003, 2004a). SedaDNA as it allows for the inference of organic matter decompos- is supposed to be of local origin (Boessenkool et al., 2014; ability (Cornwell et al., 2008; Hobbie, 1992). Furthermore, Haile et al., 2007; Jørgensen et al., 2012; Parducci et al., knowledge about the conditions under which the organic 2013; Pedersen et al., 2016; Sjögren et al., 2016) and can be matter accumulated in the past can help us understand how preserved extracellularly, even when macrofossil evidence is it accumulates today (Lyell, 1830). Recently, the sediments, absent (Arnold et al., 2011; Willerslev et al., 2003). In com- cryolithology and stratigraphy of outcrops on the western parison to arctic pollen and macrofossils, the taxonomic reso- coast of the Buor Khaya Peninsula in north-eastern Siberia lution of sedaDNA exceeds that of pollen in almost all groups were described, including organic carbon quantity, quality of higher plants and is close to the resolution of macrofossils and degradability (Günther et al., 2013b; Schirrmeister et al., (Jørgensen et al., 2012; Pedersen et al., 2013; Sønstebø et al., 2016; Stapel et al., 2016; Strauss et al., 2012, 2013, 2015). 2010). However, the palaeobotany of the site has not been studied. The purpose of this study is to combine sedaDNA and It represents a more southerly example of highly degraded pollen analyses to reconstruct past local and regional flora Ice Complex in the central Laptev region between the Lena from a Late Quaternary permafrost sediment core, which was Delta to the west and the Indigirka lowlands to the east. The recovered from the western coast of the Buor Khaya Penin- closest palaeobotanical reconstruction was carried out at the sula, a sparsely investigated region of the Late Pleistocene north-eastern coast of the peninsula using palaeolake sedi- Ice Complex. The sediments contain moderate organic car- ments, but only of Mid-Holocene origin (Willerslev et al., bon content of generally 2–5 % (Schirrmeister et al., 2016; 2014). Hence a record which describes the floristic composi- Stapel et al., 2016; Strauss et al., 2015). Given that organic tion of organic matter from the last glacial period is needed matter is the substrate for microbial turnover and the future to complete our understanding of the source and quality of release of greenhouse gases, it is crucial to get a better char- organic carbon in these deposits. acterization of the plant organic matter within those sedi- Plant macrofossils and/or pollen are the palaeobotanical ments. At the same time it is necessary to understand the en- records usually used to study the permafrost soil organic- vironmental conditions that prevailed when the organic mat- matter composition and environmental conditions during its ter was deposited, to allow for inferences of modern and fu- deposition. Plant macrofossils, which can often be identi- ture processes under comparable environmental