@article{CrausbayGenderjahnHotchkissetal.2014, author = {Crausbay, Shelley and Genderjahn, Steffi and Hotchkiss, Sara and Sachse, Dirk and Kahmen, Ansgar and Arndt, Stefan K.}, title = {Vegetation dynamics at the upper reaches of a tropical montane forest are driven by disturbance over the past 7300 years}, series = {Arctic, antarctic, and alpine research : an interdisciplinary journal}, volume = {46}, journal = {Arctic, antarctic, and alpine research : an interdisciplinary journal}, number = {4}, publisher = {Institute of Arctic and Alpine Research, University of Colorado}, address = {Boulder}, issn = {1523-0430}, doi = {10.1657/1938-4246-46.4.787}, pages = {787 -- 799}, year = {2014}, abstract = {We assessed tropical montane cloud forest (TMCF) sensitivity to natural disturbance by drought, fire, and dieback with a 7300-year-long paleorecord. We analyzed pollen assemblages, charcoal accumulation rates, and higher plant biomarker compounds (average chain length [ACL] of n-alkanes) in sediments from Wai 'anapanapa, a small lake near the upper forest limit and the mean trade wind inversion ('IWI) in Hawai`i. The paleorecord of ACL suggests increased drought frequency and a lower awl elevation from 2555-1323 cal yr B.P. and 606-334 cal yr B.P. Charcoal began to accumulate and a novel fire regime was initiated ca. 880 cal yr B.P., followed by a decreased fire return interval at ca. 550 cal yr B.P. Diebacks occurred at 2931, 2161, 1162, and 306 cal yr B.P., and two of these were independent of drought or fire. Pollen assemblages indicate that on average species composition changed only 2.8\% per decade. These dynamics, though slight, were significantly associated with disturbance. The direction of species composition change varied with disturbance type. Drought was associated with significantly more vines and lianas; fire was associated with an increase in the tree fern Sadleria and indicators of open, disturbed landscapes at the expense of epiphytic ferns; whereas stand-scale dieback was associated with an increase in the tree fern Cibotium. Though this cloud forest was dynamic in response to past disturbance, it has recovered, suggesting a resilient TMCF with no evidence of state change in vegetation type (e.g., grassland or shrubland).}, language = {en} } @article{NelsonKnohlSachseetal.2017, author = {Nelson, Daniel B. and Knohl, Alexander and Sachse, Dirk and Schefulss, Enno and Kahmen, Ansgar}, title = {Sources and abundances of leaf waxes in aerosols in central Europe}, series = {Geochimica et cosmochimica acta : journal of the Geochemical Society and the Meteoritical Society}, volume = {198}, journal = {Geochimica et cosmochimica acta : journal of the Geochemical Society and the Meteoritical Society}, publisher = {Elsevier}, address = {Oxford}, issn = {0016-7037}, doi = {10.1016/j.gca.2016.11.018}, pages = {299 -- 314}, year = {2017}, abstract = {Atmospheric transport is an understudied mechanism for leaf wax hydrogen isotope applications that contributes to mobilizing and depositing these compounds on the surface of the Earth. While previous efforts have identified the importance of atmospheric leaf wax deposition in remote marine locations, the processes are not well constrained on land in temperate latitudes where lakes are common and sedimentary leaf wax hydrogen isotope values are an attractive tool for understanding past precipitation changes. This work presents results from a field study that was conducted in 2010 and 2011 at Hainich National Park, Germany in order to evaluate the quantity and sources of leaf waxes in the atmosphere. Aerosols were sampled at approximately weekly intervals inside the forest canopy, and n-alkane distributions and hydrogen isotope values were compared with those from major tree species surrounding the sampling site. Despite sampling in what was expected to be a major production center, the distribution and hydrogen isotope values of atmospheric n-alkanes bore little resemblance to those of the local vegetation. Comparison with local meteorological data and to 10-day and 36-h back air mass trajectories indicated shifting effects of winds and temperature, and that mesoscale transport processes were more important than longrange mechanisms. Back trajectories also highlighted source effects, with easterly winds coinciding with relatively lower leaf wax hydrogen isotope values from more continental regions. These results suggest that leaf wax aerosols average over spatial scales that exceed typical surface catchment areas for small lake systems, even in forested areas, yet that the area over which these compounds are derived is still relatively regional. Depositional fluxes were also estimated in order to assess the potential importance of atmospheric transport to sedimentary archives. Although difficult to constrain, these estimates suggest that atmospheric deposition may be non-negligible for lake systems in cases where inputs from rivers or surface runoff are limited. Together, these observations provide new insights on how leaf waxes from different sources are integrated during aeolian transport and the spatial scales over which these processes occur.}, language = {en} } @article{SachseKahmenGleixner2009, author = {Sachse, Dirk and Kahmen, Ansgar and Gleixner, Gerd}, title = {Significant seasonal variation in the hydrogen isotopic composition of leaf-wax lipids for two deciduous tree ecosystems (Fagus sylvativa and Acer pseudoplatanus)}, issn = {0146-6380}, doi = {10.1016/j.orggeochem.2009.02.008}, year = {2009}, abstract = {Compound specific hydrogen isotope ratios (delta D) of long chain sedimentary n-alkanes, which mostly originate from the leaf waxes of higher terrestrial plants, are increasingly employed as paleoclimate proxies. While soil water is the ultimate hydrogen source for these lipids and the isotopic fractionation during biosynthesis of lipids is thought to remain constant, environmental parameters and plant physiological processes can alter the apparent hydrogen isotopic fractionation between leaf-wax lipids and a plant's source water. However, the magnitude and timing of these effects and their influence on the isotopic composition of lipids from higher terrestrial plants are still not well understood. Therefore we investigated the seasonal variability of leaf-wax n-alkane delta D values for two different temperate deciduous forest ecosystems that are dominated by two different tree species, Beech (Fagus sylvatica) and Maple (Acer pseudoplatanus). We found significant seasonal variations for both tree species in n-alkane delta D values of up to 40\%. on timescales as short as one week. Also, the isotopic difference between different n-alkanes from the same plant species did vary significantly and reached up to 50 parts per thousand at the same time when overall n-alkane concentrations were lowest. Since delta D values of soil water at 5 and 10 cm depth, which we assume represent the delta D value of the major water source for the investigated beech trees, were enriched in autumn compared to the spring by 30 parts per thousand, whereas n-alkane delta D values increased only by 10 parts per thousand, we observed variations in the apparent fractionation between beech leaf derived n-alkanes and soil water of up to 20 parts per thousand on a seasonal scale. This observed change in the apparent fractionation was likely caused by differences in leaf water isotopic enrichment. Based on mechanistic leaf water models we conclude that changes in the isotopic difference between water vapor and soil water were the most likely reason for the observed changes in the apparent fractionation between n- alkanes and soil water. The large variability of n-alkane concentrations and delta D values over time implies a continuous de nova synthesis of these compounds over the growing season with turnover times possibly as short as weeks. The signal to reach the soil therefore represents an integrated record of the last weeks before leaf senescence. This holds true also for the sedimentary record of small catchment lakes in humid, temperate climates, where wind transport of leaf-wax lipids is negligible compared to transfer through soil and the massive input of leaves directly into the lake in autumn.}, language = {en} } @article{SachseDawsonKahmen2015, author = {Sachse, Dirk and Dawson, Todd E. and Kahmen, Ansgar}, title = {Seasonal variation of leaf wax n-alkane production and delta H-2 values from the evergreen oak tree, Quercus agrifolia}, series = {Isotopes in environmental and health studies}, volume = {51}, journal = {Isotopes in environmental and health studies}, number = {1}, publisher = {Routledge, Taylor \& Francis Group}, address = {Abingdon}, issn = {1025-6016}, doi = {10.1080/10256016.2015.1011636}, pages = {124 -- 142}, year = {2015}, abstract = {In order to understand the timing of leaf wax synthesis in higher plants, we analysed the variability in leaf wax n-alkane concentration, composition (expressed as average chain length (ACL)), and delta H-2(wax) values as well as plant source water delta H-2 values (xylem and leaf water) in the evergreen tree Quercus agrifolia over a period of 9 months, beginning with leaf flush. We identified three distinct periods of leaf development with the first month following leaf flush being characterized by de novo synthesis and possibly removal of n-alkanes. During the following 3 months, n-alkane concentrations increased sevenfold and delta H-2(wax) and ACL values increased, suggesting this period was the major leaf wax n-alkane formation period. During the remaining 4 months of the experiment, stable values suggest cessation of leaf wax n-alkane formation. We find that n-alkane synthesis in Q. agrifolia takes place over 4 months, substantially longer than that observed for deciduous trees.}, language = {en} } @misc{SachseBillaultBowenetal.2012, author = {Sachse, Dirk and Billault, Isabelle and Bowen, Gabriel J. and Chikaraishi, Yoshito and Dawson, Todd E. and Feakins, Sarah J. and Freeman, Katherine H. and Magill, Clayton R. and McInerney, Francesca A. and van der Meer, Marcel T. J. and Polissar, Pratigya and Robins, Richard J. and Sachs, Julian P. and Schmidt, Hanns-Ludwig and Sessions, Alex L. and White, James W. C. and West, Jason B. and Kahmen, Ansgar}, title = {Molecular Paleohydrology interpreting the Hydrogen- Isotopic Composition of Lipid Biomarkers from Photosynthesizing Organisms}, series = {Annual review of earth and planetary sciences}, volume = {40}, journal = {Annual review of earth and planetary sciences}, number = {1}, editor = {Jeanloz, R}, publisher = {Annual Reviews}, address = {Palo Alto}, isbn = {978-0-8243-2040-9}, issn = {0084-6597}, doi = {10.1146/annurev-earth-042711-105535}, pages = {221 -- 249}, year = {2012}, abstract = {Hydrogen-isotopic abundances of lipid biomarkers are emerging as important proxies in the study of ancient environments and ecosystems. A decade ago, pioneering studies made use of new analytical methods and demonstrated that the hydrogen-isotopic composition of individual lipids from aquatic and terrestrial organisms can be related to the composition of their growth (i.e., environmental) water. Subsequently, compound-specific deuterium/hydrogen (D/H) ratios of sedimentary biomarkers have been increasingly used as paleohydrological proxies over a range of geological timescales. Isotopic fractionation observed between hydrogen in environmental water and hydrogen in lipids, however, is sensitive to biochemical, physiological, and environmental influences on the composition of hydrogen available for biosynthesis in cells. Here we review the factors and processes that are known to influence the hydrogen-isotopic compositions of lipids-especially n-alkanes-from photosynthesizing organisms, and we provide a framework for interpreting their D/H ratios from ancient sediments and identify future research opportunities.}, language = {en} } @article{KahmenDawsonViethetal.2011, author = {Kahmen, Ansgar and Dawson, Todd E. and Vieth, Andrea and Sachse, Dirk}, title = {Leaf wax n-alkane delta D values are determined early in the ontogeny of Populus trichocarpa leaves when grown under controlled environmental conditions}, series = {Plant, cell \& environment : cell physiology, whole-plant physiology, community physiology}, volume = {34}, journal = {Plant, cell \& environment : cell physiology, whole-plant physiology, community physiology}, number = {10}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {0140-7791}, doi = {10.1111/j.1365-3040.2011.02360.x}, pages = {1639 -- 1651}, year = {2011}, abstract = {The stable hydrogen isotope ratios (delta D) of leaf wax n-alkanes record valuable information on plant and ecosystem water relations. It remains, however, unknown if leaf wax n-alkane delta D values record only environmental variation during the brief period of time of leaf growth or if leaf wax n-alkane delta D values are affected by environmental variability throughout the entire lifespan of a leaf. To resolve these uncertainties, we irrigated Populus trichocarpa trees with a pulse of deuterium-enriched water and used compound-specific stable hydrogen isotope analyses to test if the applied tracer could be recovered from leaf wax n-alkanes of leaves that were at different stages of their development during the tracer application. Our experiment revealed that only leaf wax n-alkanes from leaves that had developed during the time of the tracer application were affected, while leaves that were already fully matured at the time of the tracer application were not. We conclude from our study that under controlled environmental conditions, leaf wax n-alkanes are synthesized only early in the ontogeny of a leaf. Our experiment has implications for the interpretation of leaf wax n-alkane delta D values in an environmental context, as it suggests that these compounds record only a brief period of the environmental variability that a leaf experiences throughout its life.}, language = {en} } @article{KahmenSchefussSachse2013, author = {Kahmen, Ansgar and Schefuss, Enno and Sachse, Dirk}, title = {Leaf water deuterium enrichment shapes leaf wax n-alkane delta D values of angiosperm plants I experimental evidence and mechanistic insights}, series = {Geochimica et cosmochimica acta : journal of the Geochemical Society and the Meteoritical Society}, volume = {111}, journal = {Geochimica et cosmochimica acta : journal of the Geochemical Society and the Meteoritical Society}, publisher = {Elsevier}, address = {Oxford}, issn = {0016-7037}, doi = {10.1016/j.gca.2012.09.003}, pages = {39 -- 49}, year = {2013}, abstract = {Leaf wax n-alkanes of terrestrial plants are long-chain hydrocarbons that can persist in sedimentary records over geologic timescales. Since meteoric water is the primary source of hydrogen used in leaf wax synthesis, the hydrogen isotope composition (delta D value) of these biomarkers contains information on hydrological processes. Consequently, leaf wax n-alkane delta D values have been advocated as powerful tools for paleohydrological research. The exact kind of hydrological information that is recorded in leaf wax n-alkanes remains, however, unclear because critical processes that determine their delta D values have not yet been resolved. In particular the effects of evaporative deuterium (D)-enrichment of leaf water on the delta D values of leaf wax n-alkanes have not yet been directly assessed and quantified. Here we present the results of a study where we experimentally tested if and by what magnitude evaporative D-enrichment of leaf water affects the delta D of leaf wax n-alkanes in angiosperm C3 and C4 plants. Our study revealed that n-alkane delta D values of all plants that we investigated were affected by evaporative D-enrichment of leaf water. For dicotyledonous plants we found that the full extent of leaf water evaporative D-enrichment is recorded in leaf wax n-alkane delta D values. For monocotyledonous plants we found that between 18\% and 68\% of the D-enrichment in leaf water was recorded in the delta D values of their n-alkanes. We hypothesize that the different magnitudes by which evaporative D-enrichment of leaf water affects the delta D values of leaf wax n-alkanes in monocotyledonous and dicotyledonous plants is the result of differences in leaf growth and development between these plant groups. Our finding that the evaporative D-enrichment of leaf water affects the delta D values of leaf wax n-alkanes in monocotyledonous and dicotyledonous plants albeit at different magnitudes - has important implications for the interpretation of leaf wax n-alkane delta D values from paleohydrological records. In addition, our finding opens the door to employ delta D values of leaf wax n-alkanes as new ecohydrological proxies for evapotranspiration that can be applied in contemporary plant and ecosystem research.}, language = {en} } @article{SteinUnsickerKahmenetal.2010, author = {Stein, Claudia and Unsicker, Sybille B. and Kahmen, Ansgar and Wagner, Markus and Audorff, Volker and Auge, Harald and Prati, Daniel and Weisser, Wolfgang W.}, title = {Impact of invertebrate herbivory in grasslands depends on plant species diversity}, issn = {0012-9658}, doi = {10.1890/09-0600.1}, year = {2010}, abstract = {Invertebrate herbivores are ubiquitous in most terrestrial ecosystems, and theory predicts that their impact on plant community biomass should depend on diversity and productivity of the associated plant communities. To elucidate general patterns in the relationship between invertebrate herbivory, plant diversity, and productivity, we carried out a long-term herbivore exclusion experiment at multiple grassland sites in a mountainous landscape of central Germany. Over a period of five years, we used above-and belowground insecticides as well as a molluscicide to manipulate invertebrate herbivory at 14 grassland sites, covering a wide range of plant species diversity (13-38 species/m(2)) and aboveground plant productivity (272-1125 g.m(-2).yr(-1)), where plant species richness and productivity of the sites were not significantly correlated. Herbivore exclusion had significant effects on the plant communities: it decreased plant species richness and evenness, and it altered plant community composition. In particular, exclusion of belowground herbivores promoted grasses at the expense of herbs. In contrast to our expectation, herbivore effects on plant community biomass were not influenced by productivity. However, effect size of invertebrate herbivores was negatively correlated with plant diversity of the grasslands: the effect of herbivory on biomass tended to be negative at sites of high diversity and positive at sites of low diversity. In general, the effects of aboveground herbivores were relatively small as compared to belowground herbivores, which were important drivers of plant community composition. Our study is the first to show that variation in the effects of invertebrate herbivory on plant communities across a landscape is significantly influenced by plant species richness.}, language = {en} } @article{GarcinSchwabGleixneretal.2012, author = {Garcin, Yannick and Schwab, Valerie F. and Gleixner, Gerd and Kahmen, Ansgar and Todou, Gilbert and Sene, Olivier and Onana, Jean-Michel and Achoundong, Gaston and Sachse, Dirk}, title = {Hydrogen isotope ratios of lacustrine sedimentary n-alkanes as proxies of tropical African hydrology insights from a calibration transect across Cameroon}, series = {Geochimica et cosmochimica acta : journal of the Geochemical Society and the Meteoritical Society}, volume = {79}, journal = {Geochimica et cosmochimica acta : journal of the Geochemical Society and the Meteoritical Society}, number = {4}, publisher = {Elsevier}, address = {Oxford}, issn = {0016-7037}, doi = {10.1016/j.gca.2011.11.039}, pages = {106 -- 126}, year = {2012}, abstract = {Hydrogen isotope values (delta D) of sedimentary aquatic and terrestrial lipid biomarkers, originating from algae, bacteria, and leaf wax, have been used to record isotopic properties of ancient source water (i.e., precipitation and/or lake water) in several mid-and high-latitude lacustrine environments. In the tropics, however, where both processes associated with isotope fractionation in the hydrologic system and vegetation strongly differ from those at higher latitudes, calibration studies for this proxy are not yet available. To close this gap of knowledge, we sampled surface sediments from 11 lakes in Cameroon to identify those hydro-climatological processes and physiological factors that determine the hydrogen isotopic composition of aquatic and terrestrial lipid biomarkers. Here we present a robust framework for the application of compound-specific hydrogen isotopes in tropical Africa. Our results show that the delta D values of the aquatic lipid biomarker n-C(17) alkane were not correlated with the delta D values of lake water. Carbon isotope measurements indicate that the n-C(17) alkane was derived from multiple source organisms that used different hydrogen pools for biosynthesis. We demonstrate that the delta D values of the n-C(29) alkane were correlated with the delta D values of surface water (i.e., river water and groundwater), which, on large spatial scales, reflect the isotopic composition of mean annual precipitation. Such a relationship has been observed at higher latitudes, supporting the robustness of the leaf-wax lipid delta D proxy on a hemispheric spatial scale. In contrast, the delta D values of the n-C(31) alkane did not show such a relationship but instead were correlated with the evaporative lake water delta D values. This result suggests distinct water sources for both leaf-wax lipids, most likely originating from two different groups of plants. These new findings have important implications for the interpretation of long-chain n-alkane delta D records from ancient lake sediments. In particular, a robust interpretation of palaeohydrological data requires knowledge of the vegetation in the catchment area as different plants may utilise different water sources. Our results also suggest that the combination of carbon and hydrogen isotopes does help to differentiate between the metabolic pathway and/or growth form of organisms and therefore, the source of hydrogen used during lipid biosynthesis.}, language = {en} } @article{KahmenSachseArndtetal.2011, author = {Kahmen, Ansgar and Sachse, Dirk and Arndt, Stefan K. and Tu, Kevin P. and Farrington, Heraldo and Vitousek, Peter M. and Dawson, Todd E.}, title = {Cellulose delta O-18 is an index of leaf-to-air vapor pressure difference (VPD) in tropical plants}, series = {Proceedings of the National Academy of Sciences of the United States of America}, volume = {108}, journal = {Proceedings of the National Academy of Sciences of the United States of America}, number = {5}, publisher = {National Acad. of Sciences}, address = {Washington}, issn = {0027-8424}, doi = {10.1073/pnas.1018906108}, pages = {1981 -- 1986}, year = {2011}, abstract = {Cellulose in plants contains oxygen that derives in most cases from precipitation. Because the stable oxygen isotope composition, delta O-18, of precipitation is associated with environmental conditions, cellulose delta O-18 should be as well. However, plant physiological models using delta O-18 suggest that cellulose delta O-18 is influenced by a complex mix of both climatic and physiological drivers. This influence complicates the interpretation of cellulose delta O-18 values in a paleo-context. Here, we combined empirical data analyses with mechanistic model simulations to i) quantify the impacts that the primary climatic drivers humidity (e(a)) and air temperature (T-air) have on cellulose delta O-18 values in different tropical ecosystems and ii) determine which environmental signal is dominating cellulose delta O-18 values. Our results revealed that e(a) and T-air equally influence cellulose delta O-18 values and that distinguishing which of these factors dominates the delta O-18 values of cellulose cannot be accomplished in the absence of additional environmental information. However, the individual impacts of e(a) and T-air on the delta O-18 values of cellulose can be integrated into a single index of plant-experienced atmospheric vapor demand: the leaf-to-air vapor pressure difference (VPD). We found a robust relationship between VPD and cellulose delta O-18 values in both empirical and modeled data in all ecosystems that we investigated. Our analysis revealed therefore that delta O-18 values in plant cellulose can be used as a proxy for VPD in tropical ecosystems. As VPD is an essential variable that determines the biogeochemical dynamics of ecosystems, our study has applications in ecological-, climate-, or forensic-sciences.}, language = {en} }