Invited review: climate change impacts in polar regions: lessons from Antarctic moss bank archives.

Mosses are the dominant plants in polar and boreal regions, areas which are experiencing rapid impacts of regional warming. Long-term monitoring programmes provide some records of the rate of recent climate change, but moss peat banks contain an unrivalled temporal record of past climate change on terrestrial plant Antarctic systems. We summarise the current understanding of climatic proxies and determinants of moss growth for contrasting continental and maritime Antarctic regions, as informed by 13C and 18O signals in organic material. Rates of moss accumulation are more than three times higher in the maritime Antarctic than continental Antarctica with growing season length being a critical determinant of growth rate, and high carbon isotope discrimination values reflecting optimal hydration conditions. Correlation plots of 13C and 18O values show that species (Chorisodontium aciphyllum / Polytrichum strictum) and growth form (hummock / bank) are the major determinants of measured isotope ratios. The interplay between moss growth form, photosynthetic physiology, water status and isotope composition are compared with developments of secondary proxies, such as chlorophyll fluorescence. These approaches provide a framework to consider the potential impact of climate change on terrestrial Antarctic habitats as well as having implications for future studies of temperate, boreal and Arctic peatlands. There are many urgent ecological and environmental problems in the Arctic related to mosses in a changing climate, but the geographical ranges of species and life-forms are difficult to track individually. Our goal was to translate what we have learned from the more simple systems in Antarctica, for application to Arctic habitats.JR is funded by Natural Environment Research Council grant NE/H014896/1.This is the author accepted manuscript. The final version is available from Wiley via http://dx.doi.org/10.1111/gcb.1277

Stable isotope signals provide seasonal climatic markers for moss functional groups.

Living moss biomass and archival peat deposits represent key indicators of present and past climatic conditions, but prediction of future climatic impacts requires appropriate marker species to be characterized under a range of contemporary conditions. Stable isotope signals in high latitude moss deposits offer potential climatic proxies. Seasonal changes in δ13C and δ18O of organic material (cellulose) in representative functional groups, and associated photosynthetic activity (as chlorophyll fluorescence) have been compared across East Anglia, UK, as a function of tissue water content. Representative species from contrasting acid bog, heathland, and fen woodland habitats were selected for monthly sampling of recent growth tissues between spring 2017 and autumn 2018, with isotopic signals in purified cellulose compared with tissue water, precipitation, and nearby groundwater signals. Sphagnum and Polytrichum groups, which tend to dominate peat formation, provided contrasting and complementary indicators of seasonal variations in carbon assimilation. Cellulose δ18O signals from Sphagnum spp. demonstrate seasonal variations in source precipitation inputs; carbon isotope signals in Polytrichum spp. indicate evaporative demand and photosynthetic limitation

Interpreting bryophyte stable carbon isotope composition: Plants as temporal and spatial climate recorders

Bryophytes are unable to control tissue water content although physiological adaptations allow growth in a wide range of habitats. Carbon isotope signals in two mosses (Syntrichia ruralis and Chorisodontium aciphyllum) and two liverworts (Conocephalum conicum and Marchantia polymorpha), whether instantaneous (real-time, Δ13C), or organic matter (as δ13COM), provide an assimilation-weighted summary of bryophyte environmental adaptations. In mosses, δ13COM is within the measured range of Δ13C values, which suggests that other proxies, such as compound specific organic signals will be representative of historical photosynthetic and growth conditions. The liverworts were photosynthetically active over a wider range of relative water contents (RWC) than the mosses. There was a consistent 5‰ offset between Δ13C values in C. conicum and M. polymorpha, suggestive of greater diffusion limitation in the latter. Analysis of a C. aciphyllum moss-peat core showed the isotopic composition over the past 200 years reflects recent anthropogenic CO2 emissions. Once corrected for source-CO2 inputs, the seasonally integrated ∆13COM between 1350 and 2000 AD varied by 1.5‰ compared with potential range of the 12‰ measured experimentally, demonstrating the relatively narrow range of conditions under which the majority of net assimilation takes place. Carbon isotope discrimination also varies spatially, with a 4‰ shift in epiphytic bryophyte organic matter found between lowland Amazonia and upper montane tropical cloud forest in the Peruvian Andes; associated with increased diffusion limitation

Leaf water δ18O reflects water vapour exchange and uptake by C3 and CAM epiphytic bromeliads in Panama.

The distributions of CAM and C3 epiphytic bromeliads across an altitudinal gradient in western Panama were identified from carbon isotope (δ13C) signals, and epiphyte water balance was investigated via oxygen isotopes (δ18O) across wet and dry seasons. There were significant seasonal differences in leaf water (δ18Olw), precipitation, stored 'tank' water and water vapour. Values of δ18Olw were evaporatively enriched at low altitude in the dry season for the C3 epiphytes, associated with low relative humidity (RH) during the day. Crassulacean acid metabolism (CAM) δ18Olw values were relatively depleted, consistent with water vapour uptake during gas exchange under high RH at night. At high altitude, cloudforest locations, C3 δ18Olw also reflected water vapour uptake by day. A mesocosm experiment with Tillandsia fasciculata (CAM) and Werauhia sanguinolenta (C3) was combined with simulations using a non-steady-state oxygen isotope leaf water model. For both C3 and CAM bromeliads, δ18Olw became progressively depleted under saturating water vapour by day and night, although evaporative enrichment was restored in the C3 W. sanguinolenta under low humidity by day. Source water in the overlapping leaf base 'tank' was also modified by evaporative δ18O exchanges. The results demonstrate how stable isotopes in leaf water provide insights for atmospheric water vapour exchanges for both C3 and CAM systems

Electrical output of bryophyte microbial fuel cell systems is sufficient to power a radio or an environmental sensor.

Plant microbial fuel cells are a recently developed technology that exploits photosynthesis in vascular plants by harnessing solar energy and generating electrical power. In this study, the model moss species Physcomitrella patens, and other environmental samples of mosses, have been used to develop a non-vascular bryophyte microbial fuel cell (bryoMFC). A novel three-dimensional anodic matrix was successfully created and characterized and was further tested in a bryoMFC to determine the capacity of mosses to generate electrical power. The importance of anodophilic microorganisms in the bryoMFC was also determined. It was found that the non-sterile bryoMFCs operated with P. patens delivered over an order of magnitude higher peak power output (2.6 ± 0.6 µW m-2) than bryoMFCs kept in near-sterile conditions (0.2 ± 0.1 µW m-2). These results confirm the importance of the microbial populations for delivering electrons to the anode in a bryoMFC. When the bryoMFCs were operated with environmental samples of moss (non-sterile) the peak power output reached 6.7 ± 0.6 mW m-2. The bryoMFCs operated with environmental samples of moss were able to power a commercial radio receiver or an environmental sensor (LCD desktop weather station).The authors are grateful for funding provided by the UK Engineering and Physical Sciences Research Council (EPSRC) (P.B., A.G.S. and C.J. Howe), EnAlgae (http://www.enalgae.eu/, INTERREG IVB NWE) (P.B. and C.J. Howe), the Royal Society URF (C.J. Harrison), the Gatsby Charitable Foundation (Fellowship GAT2962) (C.J. Harrison), the Leverhulme Trust (P.B. and C.J. Howe), the Shuttleworth Foundation (P.B.) and the Department of Science and Technology and the National Research Foundation of South Africa through the South African Research Chair Initiative Chair in Bioprocess Engineering (UID 64778) (S.T.L.H. and D.M.R.I.).This is the final version of the article. It first appeared from the Royal Society Publishing via https://doi.org/10.1098/rsos.16024

Stable isotopes and Antarctic moss banks: plants and soil microbes respond to recent warming on the Antarctic Peninsula [abstract only]

The Antarctic Peninsula is one of the most rapidly warming regions on Earth, with air temperature increases of as much as 3°C recorded since the 1950s. However, the longer-term context of this change is limited and existing records, largely relying on ice core data, are not suitably located to be able to trace the spatial signature of change over time. We are working on a project exploiting stable isotope records preserved in moss peat banks spanning 10 degrees of latitude along the Antarctic Peninsula as an archive of late Holocene climate variability. Here we present a unique time series of past moss growth and soil microbial activity that has been produced from a 150 year old moss bank at Lazarev Bay, Alexander Island (69°S), a site at the southern limit of significant plant growth in the Antarctic Peninsula region. These moss banks are ideal archives for palaeoclimate research as they are well-preserved by freezing, generally monospecific, easily dated by radiocarbon techniques, and have sufficiently high accumulation rates to permit decadal resolution. We use accumulation rates, cellulose δ13C and fossil testate amoebae to show that growth rates, assimilation and microbial productivity rose rapidly in the 1960s, consistent with temperature change, although recently may have stalled, concurrent with other evidence. The increase in biological activity is unprecedented in the last 150 years. Along with work completed on Signy Island (60°S), in the South Orkney Islands, in which we used carbon isotope evidence to show recent climate-related enhancement of CO2 assimilation and peat accumulation rates in Antarctica, the observed relationships between moss growth, microbial activity and climate suggests that moss bank records have the potential to test the regional expression of temperature variability shown by instrumental data on the Antarctic Peninsula over centennial to millennial timescales, by providing long-term records of summer growth conditions, complementing the more distant and widely dispersed ice core records. We will conclude by placing the records into the wider context of the latest progress of analysis of moss bank cores obtained along the length of the Antarctic Peninsula and Scotia arc. Royles, J., M. J. Amesbury, P. Convey, H. Griffiths, D. A. Hodgson, M. J. Leng and D. J. Charman (2013). Plants and soil microbes respond to recent warming on the Antarctic Peninsula. Current Biology 23(17): 1702-1706. Royles, J., J. Ogée, L. Wingate, D. A. Hodgson, P. Convey and H. Griffiths (2012). Carbon isotope evidence for recent climate-related enhancement of CO2 assimilation and peat accumulation rates in Antarctica. Global Change Biology 18(10): 3112-3124

Moss stable isotopes (carbon-13, oxygen-18) and testate amoebae reflect environmental inputs and microclimate along a latitudinal gradient on the Antarctic Peninsula.

The stable isotope compositions of moss tissue water (δ(2)H and δ(18)O) and cellulose (δ(13)C and δ(18)O), and testate amoebae populations were sampled from 61 contemporary surface samples along a 600-km latitudinal gradient of the Antarctic Peninsula (AP) to provide a spatial record of environmental change. The isotopic composition of moss tissue water represented an annually integrated precipitation signal with the expected isotopic depletion with increasing latitude. There was a weak, but significant, relationship between cellulose δ(18)O and latitude, with predicted source water inputs isotopically enriched compared to measured precipitation. Cellulose δ(13)C values were dependent on moss species and water content, and may reflect site exposure to strong winds. Testate amoebae assemblages were characterised by low concentrations and taxonomic diversity, with Corythion dubium and Microcorycia radiata types the most cosmopolitan taxa. The similarity between the intra- and inter-site ranges measured in all proxies suggests that microclimate and micro-topographical conditions around the moss surface were important determinants of proxy values. Isotope and testate amoebae analyses have proven value as palaeoclimatic, temporal proxies of climate change, whereas this study demonstrates that variations in isotopic and amoeboid proxies between microsites can be beyond the bounds of the current spatial variability in AP climate.The research was funded by the Natural Environment Research Council Antarctic Funding Initiative grant NE/H014896/ to DJC, PC, DAH and HG. PC, DAH and JR contribute to the BAS ‘Polar Science for Planet Earth’ research programme. Carbon isotope analyses were undertaken by Chris Kendrick at the NERC Isotope Geosciences Laboratory. Sample collection was supported by HMS Protector and HMS Endurance. Thanks to Iain Rudkin and Ashly Fusiarski for fieldwork support, to Adrian Dahood for water sample collection and to Sue Rouillard in the University of Exeter Geography drawing office for Figure 1.This is the final version of the article. It first appeared from Springer via https://doi.org/10.1007/s00442-016-3608-

Bryophyte stable isotope composition, diversity and biomass define tropical montane cloud forest extent.

Liverworts and mosses are a major component of the epiphyte flora of tropical montane forest ecosystems. Canopy access was used to analyse the distribution and vertical stratification of bryophyte epiphytes within tree crowns at nine forest sites across a 3400 m elevational gradient in Peru, from the Amazonian basin to the high Andes. The stable isotope compositions of bryophyte organic material (13C/12C and 18O/16O) are associated with surface water diffusive limitations and, along with C/N content, provide a generic index for the extent of cloud immersion. From lowland to cloud forest δ13C increased from -33‰ to -27‰, while δ18O increased from 16.3‰ to 18.0‰. Epiphytic bryophyte and associated canopy soil biomass in the cloud immersion zone was estimated at up to 45 t dry mass ha-1, and overall water holding capacity was equivalent to a 20 mm precipitation event. The study emphasizes the importance of diverse bryophyte communities in sequestering carbon in threatened habitats, with stable isotope analysis allowing future elevational shifts in the cloud base associated with changes in climate to be tracked.NERC PhD Studentshi

Can oxygen stable isotopes be used to track precipitation moisture

Variations in the isotopic composition of precipitation are determined by fractionation processes which occur during temperature- and humidity-dependent phase changes associated with evaporation and condensation. Oxygen stable isotope ratios have therefore been frequently used as a source of palaeoclimate data from a variety of proxy archives, which integrate this signal over time. Applications from ombrotrophic peatlands, where the source water used in cellulose synthesis is derived solely from precipitation, have been mostly limited to Northern Hemisphere Sphagnum-dominated bogs, with few in the Southern Hemisphere or in peatlands dominated by vascular plants. New Zealand (NZ) provides an ideal location to undertake empirical research into oxygen isotope fractionation in vascular peatlands because single taxon analysis can be easily carried out, in particular using the preserved root matrix of the restionaceous wire rush (Empodisma spp.) that forms deep Holocene peat deposits throughout the country. Furthermore, large gradients are observed in the mean isotopic composition of precipitation across NZ, caused primarily by the relative influence of different climate modes. Here, we test whether δ18O of Empodisma α-cellulose from ombrotrophic restiad peatlands in NZ can provide a methodology for developing palaeoclimate records of past precipitation δ18O. Surface plant, water and precipitation samples were taken over spatial (six sites spanning >10◦ latitude) and temporal (monthly measurements over one year) gradients. A link between the isotopic composition of root-associated water, the most likely source water for plant growth, and precipitation in both datasets was found. Back-trajectory modelling of precipitation moisture source for rain days prior to sampling showed clear seasonality in the temporal data that was reflected in root-associated water. The link between source water and plant cellulose was less clear, although mechanistic modelling predicted mean cellulose values within published error margins for both datasets. Improved physiological understanding and modelling of δ18O in restiad peatlands should enable use of this approach as a new source of palaeoclimate data to reconstruct changes in past atmospheric circulation