Biogeosciences www.biogeosciences.net 1726-4170 1726-4189 3 4 2006 10.5194/bg-3-421-2006 http://www.biogeosciences.net/3/421/2006/ http://www.biogeosciences.net/3/421/2006/bg-3-421-2006.html http://www.biogeosciences.net/3/421/2006/bg-3-421-2006.pdf 421 437 2006-09-27 Water use strategies and ecosystem-atmosphere exchange of CO2 in two highly seasonal environments A. Arneth almut.arneth@nateko.lu.se E. M. Veenendaal C. Best W. Timmermans O. Kolle L. Montagnani O. Shibistova Department of Physical Geography and Ecosystems Analysis, Lund University, 22363 Lund, Sweden Nature Conservation and Plant Ecology Group, Wageningen University and Research Centre Max Planck Institute for Biogeochemistry, PO Box 100164, 07701 Jena, Germany The International Institute for Geo-Information Science and Earth Observation (ITC), Enschede, The Netherlands University of Tuscia, Viterbo, Italy V. N. Sukachev Forest Institute, Akademgorodok, 660036 Krasnoyarsk, Russia We compare assimilation and respiration rates, and water use strategies in four divergent ecosystems located in cold-continental central Siberia and in semi-arid southern Africa. These seemingly unrelated systems have in common a harsh and highly seasonal environment with a very sharp transition between the dormant and the active season, with vegetation facing dry air and soil conditions for at least part of the year. Moreover, the northern high latitudes and the semi-arid tropics will likely experience changes in key environmental parameters (e.g., air temperature and precipitation) in the future; indeed, in some regions marked climate trends have already been observed over the last decade or so. The magnitude of instantaneous or daily assimilation and respiration rates, derived from one to two years of eddy covariance measurements in each of the four ecosystems, was not related to the growth environment. For instance, respiration rates were clearly highest in the two deciduous systems included in the analysis (a Mopane woodland in northern Botswana and a Downy birch forest in Siberia; >300 mmol m−2 d−1), while assimilation rates in the Mopane woodland were relatively similar to a Siberian Scots pine canopy for a large part of the active season (ca. 420 mmol m−2 d−1). Acknowledging the limited number of ecosystems compared here, these data nevertheless demonstrate that factors like vegetation type, canopy phenology or ecosystem age can override larger-scale climate differences in terms of their effects on carbon assimilation and respiration rates. By far the highest rates of assimilation were observed in Downy birch, an early successional species. These were achieved at a rather conservative water use, as indicated by relatively low levels of λ, the marginal water cost of plant carbon gain. Surprisingly, the Mopane woodland growing in the semi-arid environment had significantly higher values of λ. However, its water use strategy included a very plastic response to intermittently dry periods, and values of λ were much more conservative overall during a rainy season with low precipitation and high air saturation deficits. Our comparison demonstrates that forest ecosystems can respond very dynamically in terms of water use strategy, both on interannual and much shorter time scales. But it remains to be evaluated whether and in which ecosystems this plasticity is mainly due to a short-term stomatal response, or alternatively goes hand in hand with changes in canopy photosynthetic capacity. Arneth, A., Kelliher, F. M., Gower, S. T., Scott, N. A., Byers, J. N., and McSeveny, T. M.: Environmental variables regulating soil carbon dioxide efflux following clear-cutting of a \textitPinus radiata D. Don plantation, J. Geophys. Res., 103, 5695–5705, 1998. Arneth, A., Kurbatova, J., Kolle, O., Shibistova, O., Lloyd, J., Vygodskaya, N. N., and Schulze, E.-D.: Comparative ecosystem - atmosphere exchange of energy and mass in a European Russian and a central Siberian bog II. Interseasonal and interannual variability of CO2 fluxes, Tellus, 54 B, 514–530, 2002a. Arneth, A., Lloyd, J., \vSantruková, H., Bird, M., Grigoriev, S., Gleixner, G., and Schulze, E.-D.: Response of central Siberian Scots pine in to soil water deficit and long-term trends in atmospheric CO2 concentration, Global Biogeochem. Cycles, 16, 5/1–5/13, 2002b. Arneth, A., Lloyd, J., Shibistova, O., and Kolle, O.: Snow melt in the boreal environment: exploring spring energy and CO2 fluxes in two typical central Siberian ecosystems, Boreal Environ. Res., in press, 2006. Atkin, O. K. and Tjoelker, M. G.: Thermal acclimation and the dynamic response of plant respiration to temperature, Trends in Plant Science, 8, 343, 2003. Aurela, M., Laurila, T., and Tuovinen, J. P.: The timing of snow melt controls the annual CO2 balance in a subarctic fen, Geophys. Res. Lett., 31, L16119, doi:10.1029/2004GL020315, 2004. Bhalerao, R., Keskitalo, J., Sterky, F., Erlandsson, R., Bjorkbacka, H., Birve, S. J., Karlsson, J., Gardestrom, P., Gustafsson, P., Lundeberg, J., and Jansson, S.: Gene expression in autumn leaves, Plant Physiology, 131, 430–442, 2003. Bond, W. J., Woodward, F. I., and Midgley, G. F.: The global distribution of ecosystems in a world without fire, New Phytologist, 165, 525–538, 2005. Bowman, D. and Prior, L. D.: Why do evergreen trees dominate the Australian seasonal tropics?, Australian Journal of Botany, 53, 379–399, 2005. Choudhury, B. J.: Relationships between vegetation indices, radiation absorption, and net photosynthesis evaluated by a sensitivity analysis, Remote Sens. Environ., 22, 209–233, 1987. Choudhury, B. J., Ahmed, N. U., Idso, S. B., Reginato, R. J., and Daughtry, C. S. T.: Relations between evaporation coefficients and vegetation indices studied by model simulations, Remote Sens. Environ., 50, 1–17, 1994. Clein, J. S. and Schimel, J. P.: Microbial activity of tundra and taiga soils at sub-zero temperatures, Soil Biology and Biochemistry, 27, 1231–1234, 1995. Collier, D. E. and Thibodeau, B. A.: Changes in respiration and chemical content during autumnal senescence of Populus tremuloides and Quercus rubra leaves, Tree Physiology, 15, 759–764, 1995. Cowan, I. R.: Stomatal behaviour and environment, Advances in Botanical Research, 4, 117–128, 1977. Cowan, I. R.: Regulation of water use in relation to carbon gain in higher plants, p. 589–613 in: Physiological Plant Ecology II. Water Relations and Carbon Assimilation, edited by: Lange, O. L., Nobel, P. S., Osmond, C. B., and Ziegler, H., Springer, Berlin, 1982. Cox, P. M., Betts, R. A., Jones, C. D., Spall, S. A., and Totterdell, I. J.: Acceleration of global warming due to carbon-cycle feedbacks in a coupled climate model, Nature, 408, 184–186, 2000. Dolman, A. J., Maximov, T. C., Moors, E. J., Maximov, A. P., Elbers, J. A., Kononov, V., Waterloo, M. J., and v. d. Molen, M. K.: Net ecosystem exchange of carbon dioxide and water of far eastern Siberian Larch (Larix cajanderii) on permafrost, Biogeosciences, 1, 133–146, 2004. Eklundh, L. and Olsson, L.: Vegetation index trends for the African Sahel 1982–1999, J. Geophys. Res., 30, 1430, doi:10.1029/2002GL016772, 2003. Eugster, W. and Senn, W.: A cospectral correction model for measurement of turbulent NO2 flux, Boundary-Layer Meteorol., 74, 321–340, 1995. Falge, E., Tenhunen, J., Baldocchi, D., Aubinet, M., Bakwin, P., Berbigier, P., Bernhofer, C., Bonnefond, J. M., Burba, G., Clement, R., Davis, K. J., Elbers, J. A., Falk, M., Goldstein, A. H., Grelle, A., Granier, A., Grunwald, T., Gudmundsson, J., Hollinger, D., Janssens, I. A., Keronen, P., Kowalski, A. S., Katul, G., Law, B. E., Malhi, Y., Meyers, T., Monson, R. K., Moors, E., Munger, J. W., Oechel, W., U, K. T. P., Pilegaard, K., Rannik, U., Rebmann, C., Suyker, A., Thorgeirsson, H., Tirone, G., Turnipseed, A., Wilson, K., and Wofsy, S.: Phase and amplitude of ecosystem carbon release and uptake potentials as derived from FLUXNET measurements, Agric. For. Meteorol., 113, 75–95, 2002. Foley, J. A., Costa, M. H., Delire, C., Ramankutty, N., and Snyder, P.: Green surprise? How terrestrial ecosystems could affect earth's climate, Frontiers in Ecology and Environment, 1, 38–44, 2003. Friedlingstein, P., Bopp, L., Ciais, P., Dufresne, J.-L., Fairhead, L., LeTreut, H., Monfray, P., and Orr, J.: Positive feedback between future climate change and the carbon cycle, Geophys. Res. Lett., 28, 1543–1546, 2001. Gamache, I. and Payette, S.: Height growth response of tree line black spruce to recent climate warming across the forest-tundra of eastern Canada, J. Ecology, 92, 835–845, 2004. Granier, A., Pilegaard, K., and Jensen, N. O.: Similar net ecosystem exchange of beech stands located in France and Denmark, Agric. For. Meteorol., 114, 75, 2002. Hanan, N., Kabat, P., Dolman, A. J., and Elbers, J. A.: Photosynthesis and carbon balance of a Sahelian fallow savanna, Global Change Biology, 4, 523–538, 1998. Heimann, M.: The Eurosiberian Carbonflux project, Tellus, 54B, 417–419, 2002. Hély, C., Bremond, L., Alleaume, S., Smith, B., Sykes, M. T., and Guiot, J.: Sensitivity of African biomes to changes in precipitation regime, Global Ecology and Biogeography, Global Ecol. Biogeogr., 15(3), 258–270, 2006. Hickler, T., Eklundh, L., Seaquist, J., Smith, B., Ardö, J., Olsson, L., and Sjöström, M.: Precipitation controls Sahel greening trend, Geophys. Res. Lett., 32, L21415, doi:10.1029/2005GL024370, 2005. Holtmeier, F. K. and Broll, G.: Sensitivity and response of northern hemisphere altitudinal and polar treelines to environmental change at landscape and local scales, Global Ecology and Biogeography, 14, 395–410, 2005. Jarvis, P. J. and Linder, S.: Constraints to growth of boreal forests, Nature, 405, 904–905, 2000. Jonsdottir, I. S., Virtanen, R., and Karnefelt, I.: Large-scale differentiation and dynamics in tundra plant populations and vegetation, Ambio, 28, 230–238, 1999. Justice, C. O., Vermote, E., Townshend, J. R. G., Defries, R., Roy, D. P., Hall, D. K., Salomonson, V. V., Privette, J. L., Riggs, G., Strahler, A., Lucht, W., Myeni, R. B., Knyazikhin, Y., Running, S. W., Nemani, R. R., Wan, Z., Huete, A. R., Leeuwen, W. V., Wolfe, R. E., Giglio, L., Muller, J.-P., Lewis, P., and Barnsley, M. J.: The Moderate Resolution Imaging Spectroradiometer (MODIS): Land remote sensing for global change research, IEEE Trans. Geosci. Rem. Sens., 36, 1228–1249, 1998. Kaplan, J., Bigelow, N. H., Prentice, I. C., Harrison, S., Bartlein, P. J., Christensen, T. R., Cramer, W., Matveyeva, N. V., McGuire, A. D., Murray, D. F., Razzhivin, V. Y., Smith, B., Walker, D., Anderson, P. M., Andreev, A. A., Brubaker, L. B., Edwards, M. E., and Lozhkin, A. V.: Climate change and Arctic ecosystems: 2. Modelling, paleodata-model comparisons, and future projections, J. Geophys. Res., 108, 8171, doi:10.1029/2002JD002558, 2003. Kelliher, F. M., Leuning, R., and Schulze, E.-D.: Evaporation and canopy characteristics of coniferous forests and grasslands, Oecologia, 95, 153–163, 1993. Kelliher, F. M., Lloyd, J., Arneth, A., Luhker, B., Byers, J. N., McSeveny, T. M., Milukova, I., Grigoriev, S., Panfyorov, M., Sogatchev, A., Varlargin, A., Ziegler, W., Bauer, G., Wong, S. C., and Schulze, E. D.: Carbon dioxide efflux density from the floor of a central Siberian pine forest, Agric. For. Meteorol., 94, 217–232, 1999. Kimball, B. A. and Lemon, E. R.: Air turbulence effects upon soil gas exchange, Soil Sci. Soc. Amer. J., 35, 16–21, 1971. Kurbatova, J., Arneth, A., Vygodskaya, N. N., Kolle, O., Varlargin, A. B., Milyukova, I. M., Tchebakova, N. M., Schulze, E.-D., and Lloyd, J.: Comparative ecosystem – atmosphere exchange of energy and mass in a European Russian and a central Siberian bog I. Interseasonal and interannual variability of energy and latent heat fluxes during the snowfree period, Tellus, 54 B, 497–513, 2002. Lafleur, P. M., Roulet, N. T., and Admiral, S. W.: Annual cycle of CO2 exchange at a bog peatland, J. Geophys. Res., 106, 3071–3081, 2001. Law, B. E., Ryan, M. G., and Anthoni, P. M.: Seasonal and annual respiration of a ponderosa pine ecosystem, Global Change Biology, 5, 169–182, 1999. Lloyd, J.: Modelling stomatal responses to environment in \textitMacadamia integrifolia, Aust. J. Plant Physiol., 18, 649–660, 1991. Lloyd, J., Grace, J., Miranda, A. C., Meir, P., Wong, S. C., Miranda, B. S., Wright, I. R., Gash, J. H. C., and McIntyre, J.: A simple calibrated model of Amazon rainforest productivity based on leaf biochemical properties, Plant Cell Environ., 18, 1129–1145, 1995. Lloyd, J., Shibistova, O., Zolotoukhine, D., Kolle, O., Arneth, A., Wirth, C., Styles, J. M., Tchebakova, N., and Schulze, E. D.: Seasonal and annual variations in the photosynthetic productivity and carbon balance of a central Siberian pine forest, Tellus, 54 B, 590–610, 2002. Lloyd, J. and Taylor, J. A.: On the temperature dependence of soil respiration, Functional Ecology, 8, 315–323, 1994. Lucht, W., Prentice, I. C., Myneni, R. B., Sitch, S., Friedlingstein, P., Cramer, W., Bousquet, P., Buermann, W., and Smith, B.: Climatic control of the high-latitude vegetation greening trend and Pinatubo effect, Science, 296, 1687–1689, 2002. Mellander, P.-E.: Spring water stress in Scots pine: Interaction of snow and temperature, Swedish University of Agricultural Sciences, Stockholm, 2003. Monteith, J. L. and Unsworth, M.: Principles of environmental physics, Arnold, London, 1990. Myneni, R. B., Hoffman, S., Knyazikhin, Y., Privette, J. L., Glassy, J., Tian, Y., Wang, Y., Song, X., Zhang, Y., Smith, G. R., Lotsch, A., Friedl, M., Morisette, J. T., Votava, P., Nemani, R. R., and Running, S. W.: Global products of vegetation leaf area and fraction absorbed PAR from year one of MODIS data, Remote Sens. Environ., 83, 214–231, 2002. Myneni, R. B., Keeling, C. D., Tucker, C. J., Asrar, G., and Nemani, R. R.: Increased plant growth in the northern high latitudes from 1981 to 1991, Nature, 386, 698–702, 1997. Pendall, E., Bridgham, S., Hanson, P. J., Hungate, B., Kicklighter, D. W., Johnson, D. W., Law, B. E., Luo, Y., Megonigal, J. P., Olsrud, M., Ryan, M. G., and Wan, S.: Below-ground process responses to elevated CO2 and temperature: a discussion of observations, measurement methods, and models, New Phytologist, 162, 311–322, 2004. Röser, C., Montagnani, L., Schulze, E. D., Mollicone, D., Kolle, O., Meroni, M., Papale, D., Marchesini, L. B., Federici, S., and Valentini, R.: Net CO2 exchange rates in three different successional stages of the "Dark Taiga" of central Siberia, Tellus, 54 B, 642–654, 2002. Scheffer, M., Holmgren, M., Brovkin, V., and Claussen, M.: Synergy between small- and large-scale feedbacks of vegetation on the water cycle, Global Change Biology, 11, 1003–1012, 2005. Schulze, E.-D.: Plant life forms and their carbon, water and nutrient relations, p. 616–676, in: Physiological Plant Ecology, edited by: Lange, O. L., Nobel, P. S., Osmond, C. B., and Ziegler, H., Springer Verlag, Heidelberg, 1982. Schulze, E.-D. and Hall, A. E.: Stomatal responses, water loss and CO2 assimilation rates of plants in contrasting environments, p. 181–229, in: Physiological Plant Ecology, edited by: Lange, O. L., Nobel, P. S., Osmond, C. B., and Ziegler, H., Springer Verlag, Heidelberg, 1982. Shibistova, O., Lloyd, J., Zrazewhskaya, G., Arneth, A., Kolle, O., Knohl, A., and Schmerler, J.: Annual ecosystem respiration budget for a \textitPinus sylvestris stand in central Siberia, Tellus, 54B, 568–589, 2002. Sitch, S., Smith, B., Prentice, I. C., Arneth, A., Bondeau, A., Cramer, W., Kaplan, J. O., Levis, S., Lucht, W., Sykes, M. T., Thonicke, K., and Venevsky, S.: Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model, Global Change Biology, 9, 161–185, 2003. Sitch, S., McGuire, A. D., Kimball, J., Gedney, N., Gamno, J., Engstrom, R., Wolf, A., Zhuang, Q., and Clein, J.: Assessing the circumpolar carbon balance of arctic tundra with remote sensing and process/based modelling approaches, Global Biogeochem. Cycles, in press, 2006. Skre, O., Baxter, R., Crawford, R. M. M., Callaghan, T. V., and Fedorkov, A.: How will the tundra-taiga interface respond to climate change?, Ambio, 12, 37–46, 2002. Smith, B., Prentice, I. C., and Sykes, M. T.: Representation of vegetation dynamics in the modelling of terrestrial ecosystems: comparing two contrasting approaches within European climate space, Global Ecology & Biogeography, 10, 621–637, 2001. Suni, T., Vesala, T., Markkanen, T., Berninger, F., Hari, P., Mäkelä, A., Ilvesniemi, H., Nikinmaa, E., Laurila, T., Aurela, M., Grelle, A., Lindroth, A., Arneth, A., Lloyd, J., and Shibistova, O.: Air temperature triggers the commencement of evergreen boreal forest photosynthesis in spring, Global Change Biology, 9, 1410–1426, 2003. Tchebakova, N. M., Kolle, O., Zolotoukhine, D., Arneth, A., Styles, J. M., Vygodskaya, N. N., Schulze, E. D., Shibistova, O., and Lloyd, J.: Inter-annual and seasonal variations of energy and water vapour fluxes above a Pinus sylvestris forest in the Siberian middle taiga, Tellus, 54B, 537–551, 2002. Timberlake, J. R.: Colophospermum mopane, Annotated Bibliography and Review, 1995. Veenendaal, E. M., Kolle, O., and Lloyd, J.: Seasonal variation in energy fluxes and carbon dioxide exchange for a broad-leaved semi-arid savanna (Mopane woodland) in southern Africa, Global Change Biology, 10, 318–328, 2004. Vermote, E. F. and Vermeulen, A.: MODIS Atmospheric correction algorithm: spectral reflectances (MOD09), Algorithm Technical Background Document, 1999. Whittaker, R.: Communities and Ecosystems, McMillas, New York, 1975. Williams, C. A. and Albertson, J. D.: Contrasting short- and long-timescale effects of vegetation dynamics on water and carbon fluxes in water-limited ecosystems, Water Resour. Res., 41, 2005. Woodward, F. I. and Lomas, M. R.: Vegetation dynamics – simulating responses to climatic change, Biological Reviews, 79, 643/670, 2004. Zhou, L. M., Tucker, C. J., Kaufmann, R. K., Slayback, D., Shabanov, N. V., and Myneni, R. B.: Variations in northern vegetation activity inferred from satellite data of vegetation index during 1981 to 1999, J. Geophys. Res., 106, 20 069–20 083, 2001.