<?xml version="1.0" encoding="utf-8" standalone="no"?>
<!DOCTYPE article SYSTEM "http://www.clim-past-discuss.net/inc/cpd/copernicus.dtd">
<article language="en">
	<journal>
		<journal_title>Climate of the Past Discussions</journal_title>
		<journal_url>www.clim-past-discuss.net</journal_url>
		<issn>1814-9340</issn>
		<eissn>1814-9359</eissn>
		<volume_number>5</volume_number>
		<issue_number>2</issue_number>
		<publication_year>2009</publication_year>
	</journal>
	<doi>10.5194/cpd-5-1187-2009</doi>
	<article_url>http://www.clim-past-discuss.net/5/1187/2009/</article_url>
	<abstract_html>http://www.clim-past-discuss.net/5/1187/2009/cpd-5-1187-2009.html</abstract_html>
	<fulltext_pdf>http://www.clim-past-discuss.net/5/1187/2009/cpd-5-1187-2009.pdf</fulltext_pdf>
	<start_page>1187</start_page>
	<end_page>1213</end_page>
	<publication_date>2009-03-31</publication_date>
	<article_title content_type="html">Climate and CO&lt;sub&gt;2&lt;/sub&gt; modulate the C&lt;sub&gt;3&lt;/sub&gt;-C&lt;sub&gt;4&lt;/sub&gt; balance and &amp;delta;&lt;sup&gt;13&lt;/sup&gt;C signal in simulated vegetation</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>O. Flores</name>
			<email>olivier.flores@cefe.cnrs.fr</email>
		</author>
		<author numeration="2" affiliations="1,2">
			<name>E. S. Gritti</name>
		</author>
		<author numeration="3" affiliations="2">
			<name>D. Jolly</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">CEFE, UMR 5175 CNRS, 1919, route de Mende, 34293, Montpellier cedex 5, France</affiliation>
		<affiliation numeration="2" content_type="html">ISEM, UMR 5554 CNRS/Univ. Montpellier II, Case 61, 34095 Montpellier cedex 5, France</affiliation>
	</affiliations>
	<abstract content_type="html">Fossil pollen data and &amp;delta;&lt;sup&gt;13&lt;/sup&gt;C measurements from cores collected in peatbogs
or lakes have shown major changes in the terrestrial vegetation during Late
Quaternary. Although the effect of climate on the C&lt;sub&gt;3&lt;/sub&gt;-C&lt;sub&gt;4&lt;/sub&gt; balance has been
discussed for 50 years, the impact of a low atmospheric CO&lt;sub&gt;2&lt;/sub&gt; during the Last
Glacial Maximum (LGM) was emphasized recently and conflicting evidence
exists. In this paper, we use a physiologically-based biome model (BIOME4) in
an iterative mode to simulate vegetation response to changing mean climate
conditions and atmospheric CO&lt;sub&gt;2&lt;/sub&gt; partial pressure (&lt;i&gt;p&lt;/i&gt;&lt;sub&gt;CO&lt;sub&gt;2&lt;/sub&gt;&lt;/sub&gt;). In particular, we
investigate the transition from LGM to present conditions in two sites which
changed from either a C&lt;sub&gt;4&lt;/sub&gt;- or a C&lt;sub&gt;3&lt;/sub&gt;-dominated vegetation to the opposite
pole, respectively at Kuruyange (Burundi) and Lingtaï (Central Loess
Plateau, China). The response of the C&lt;sub&gt;3&lt;/sub&gt;-C&lt;sub&gt;4&lt;/sub&gt; balance and &amp;delta;&lt;sup&gt;13&lt;/sup&gt;C signal in the
simulated vegetation are investigated. The results show that the vegetation
is primarily sensitive to temperature and &lt;i&gt;p&lt;/i&gt;&lt;sub&gt;CO&lt;sub&gt;2&lt;/sub&gt;&lt;/sub&gt;. Rainfall impacted the
simulated variables below a threshold which decreased with higher &lt;i&gt;p&lt;/i&gt;&lt;sub&gt;CO&lt;sub&gt;2&lt;/sub&gt;&lt;/sub&gt;.
Climate and &lt;i&gt;p&lt;/i&gt;&lt;sub&gt;CO&lt;sub&gt;2&lt;/sub&gt;&lt;/sub&gt; interacted differently between the two sites showing
indirect effects on the &amp;delta;&lt;sup&gt;13&lt;/sup&gt;C signal. Moreover, the plant functional types
(PFTs) differed in their composition and in their response between the two
sites, emphasizing that the competition between C&lt;sub&gt;3&lt;/sub&gt; and C&lt;sub&gt;4&lt;/sub&gt; plants cannot
be hardly considered as a simple binary scheme. Our results confirm the
advantages of using process-based models to understand past vegetation
changes and the need to take account of multiple drivers when the C&lt;sub&gt;3&lt;/sub&gt;-C&lt;sub&gt;4&lt;/sub&gt;
balance is reconstructed from a palaeo-&amp;delta;&lt;sup&gt;13&lt;/sup&gt;C signal.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Aucour, A. and Hillaire-Marcel, C.: Late Quaternary biomass changes from $^13$C measurements in a highland peatbog from equatorial Africa (Burundi), Quaternary Res., 41, 225–233, 1994. </reference>
		<reference numeration="2" content_type="text"> Aucour, A., Bonnefille, R., and Hillaire-Marcel, C.: Sources and accumulation rates of organic carbon in an equatorial peatbog (Burundi, East Africa) during the Holocene: carbon isotope constraints, Palaeogeogr. Palaeocl., 150, 179–189, 1999. </reference>
		<reference numeration="3" content_type="text"> Balsam, W., Ji, J., and Chen, J.: Climatic interpretation of the Luochuan and Lingta\&quot;i loess sections, China, based on changing iron oxide mineralogy and magnetic susceptibility, Earth Planet. Sci. Lett., 223, 335–348, \doi10.1016/j.epsl.2004.04.023, online availabel at: prefixhttp://www.sciencedirect.com/science/article/B6V61-4CMHW4H-1/% 2/f0b75eda3057d9e7caf9cb6e3559d74c, 2004. </reference>
		<reference numeration="4" content_type="text"> Bonnefille, R. and Chalié, F.: Pollen-inferred precipitation time-series from equatorial mountains, Africa, the last 40 kyr BP, Global Planet. Change, 26, 25–50, \doi10.1016/S0921-8181(00)00032-1, online available at: prefixhttp://www.sciencedirect.com/science/article/B6VF0-41NK8KV-4/% 2/b9e6b181bc5605dabde493f8d445dd47, 2000. </reference>
		<reference numeration="5" content_type="text"> Bonnefille, R. and Riollet, G.: The Kashiru pollen sequence (Burundi). Palaeoclimatic implications for the last 40000 yr BP in tropical Africa, Quaternary Res., 30, 19–35, 1988. </reference>
		<reference numeration="6" content_type="text"> Bonnefille, R., Roeland, J C., and Guiot, J.: Temperature and rainfall estimates for the past 40,000 years in equatorial Africa, Nature, 346, 347–349, \doi10.1038/346347a0, online available at: prefixhttp://dx.doi.org/10.1038/346347a0, 1990. </reference>
		<reference numeration="7" content_type="text"> Bonnefille, R., Riollet, G., and Buchet, G.: Nouvelle séquence pollinique d&apos;une tourbière de la crête Zaire-Nil (Burundi), Rev. Palaeobot. palyno., 67, 315–330, 1991. </reference>
		<reference numeration="8" content_type="text"> Bonnefille, R., Riollet, G., Buchet, G., Icole, M., Lafont, R., Arnold, M., and Jolly, D.: Glacial/interglacial record from intertropical Africa, high resolution pollen and carbon data at Rusaka, Burundi, Quaternary Sci. Rev., 14, 917–936, \doi10.1016/0277-3791(95)00071-2, online available at: prefixhttp://www.sciencedirect.com/science/article/B6VBC-3Y45T0F-G/% 2/935d72141738130a956b4197008cc7e9, 1995. </reference>
		<reference numeration="9" content_type="text"> Boom, A., Marchant, R., Hooghiemstra, H., and Damst, J. S S.: CO&lt;sub&gt;2&lt;/sub&gt;- and temperature-controlled altitudinal shifts of C&lt;sub&gt;4&lt;/sub&gt;- and C&lt;sub&gt;3&lt;/sub&gt;-dominated grasslands allow reconstruction of palaeoatmospheric $p$CO&lt;sub&gt;2&lt;/sub&gt;, Palaeogeogr. Palaeocl., 177, 151–168, \doi10.1016/S0031-0182(01)00357-1, online available at: prefixhttp://www.sciencedirect.com/science/article/B6V6R-44HWS2G-5/% 2/de7e43d6ba4f0823157de9e10feaed84, 2002. </reference>
		<reference numeration="10" content_type="text"> Coetzee, J.: Pollen analytical studies in east and southern Africa, Palaeoecol. Afr., 3, 1–146, 1967. </reference>
		<reference numeration="11" content_type="text"> Cole, D R. and Monger, H C.: Influence of atmospheric CO&lt;sub&gt;2&lt;/sub&gt; on the decline of C&lt;sub&gt;4&lt;/sub&gt; plants during the last deglaciation, Nature, 368, 533–536, \doi10.1038/368533a0, online available at: prefixhttp://dx.doi.org/10.1038/368533a0, 1994. </reference>
		<reference numeration="12" content_type="text"> Deines, P.: The isotopic composition of reduced inorganic carbon, in: Handbook of Environmental Isotope Chemistry: the terrestrial environment, edited by: Fritz, P. and Fontes, J., vol. 1, 329–406, Elsevier, Amsterdam, The Netherlands, 1980. </reference>
		<reference numeration="13" content_type="text"> Gu, Z Y., Liu, Q., Xu, B., Han, J M., Yang, S L., Ding, Z L., and Liu, T S.: Climate as the dominant control on C-3 and C-4 plant abundance in the Loess Plateau: Organic carbon isotope evidence from the last glacial-interglacial loess-soil sequences, Chinese Sci. Bull., 48, 1271–1276, 2003. </reference>
		<reference numeration="14" content_type="text"> Guiot, J.: Statistical analyses of biospherical variability, in: Long-Term Climatic Variations: Data and Modelling, edited by: Duplessy, J. and Spyridakis, M., vol 22 of NATO ASI Series I, 299–334, Springer-Verlag, New-York, USA, 1994. </reference>
		<reference numeration="15" content_type="text"> Guiot, J., Torre, F., Jolly, D., Peyron, O., Boreux, J J., and Cheddadi, R.: Inverse vegetation modeling by Monte Carlo sampling to reconstruct palaeoclimates under changed precipitation seasonality and CO&lt;sub&gt;2&lt;/sub&gt; conditions: application to glacial climate in Mediterranean region, Ecol. Modell., 127, 119–140, \doi10.1016/S0304-3800(99)00219-7, online available at: prefixhttp://www.sciencedirect.com/science/article/B6VBS-3YRVDGH-2/% 2/d93a6c4f34f7fa03462b434dfe4fd213, 2000. </reference>
		<reference numeration="16" content_type="text"> Hamilton, A.: The interpretation of pollen diagrams from highland Uganda, Palaeoecol. Afr., 7, 45–149, 1972. </reference>
		<reference numeration="17" content_type="text"> Hamilton, A.: Environmental history of East Africa: a study of the Quaternary, Academic press, New York, USA, 328~pp., 1982. </reference>
		<reference numeration="18" content_type="text"> Hatté, C. and Guiot, J.: Palaeoprecipitation reconstruction by inverse modelling using the isotopic signal of loess organic matter: application to the Nußloch loess sequence (Rhine Valley, Germany), Clim. Dynam., 25, 315–327, \doi10.1007/s00382-005-0034-3, online available at: prefixhttp://dx.doi.org/10.1007/s00382-005-0034-3, 2005. </reference>
		<reference numeration="19" content_type="text"> Haxeltine, A. and Prentice, I.: BIOME3: An equilibrium terrestrial biosphere model based on ecophysiological constraints, availability, and competition among plant functional types, Global Biogeochem. Cy., 10, 693–709, 1996. </reference>
		<reference numeration="20" content_type="text"> Haxeltine, A., Prentice, I., and Cresswell, I.: A coupled carbon and water flux model to predict vegetation structure, J. Veg. Sci., 7, 651–666, 1996. </reference>
		<reference numeration="21" content_type="text"> Huang, Y., Street-Perrott, F A., Metcalfe, S E., Brenner, M., Moreland, M., and Freeman, K H.: Climate Change as the Dominant Control on Glacial-Interglacial Variations in C&lt;sub&gt;3&lt;/sub&gt; and C&lt;sub&gt;4&lt;/sub&gt; Plant Abundance, Science, 293, 1647–1651, \doi10.1126/science.1060143, online available at: prefixhttp://www.sciencemag.org/cgi/content/abstract/293/5535/1647, 2001. </reference>
		<reference numeration="22" content_type="text"> Jackson, R. B., Canadell, J., Ehleringer, J. R., Mooney, H. A., Sala, O. E., and Schulze, E. D.: A global analysis of root distributions for terrestrial biomes Oecologia, 108, 389–411, 1996. </reference>
		<reference numeration="23" content_type="text"> Jolly, D. and Haxeltine, A.: Effect of low glacial atmopsheric CO&lt;sub&gt;2&lt;/sub&gt; on tropical African montane vegetation, Science, 276, 786–788, online available at: prefixhttp://www.sciencemag.org/cgi/content/abstract/276/5313/786, 1997. </reference>
		<reference numeration="24" content_type="text"> Jolly, D., Bonnefille, R., and Roux, M.: Numerical interpretation of a high resolution Holocene pollen record from Burundi, Palaeogeogr. Palaeocl., 109, 357–370, 1994. </reference>
		<reference numeration="25" content_type="text"> Jolly, D., Taylor, D., Marchant, R., Hamilton, A., Bonnefille, R., Buchet, G., and Riollet, G.: Vegetation dynamics in central Africa since 18000 yr BP: pollen records from the interlacustinterlacustrine highlands of Burundi, Rwanda and western Uganda, J. Biogeogr., 24, 495–512, online available at: prefixhttp://www3.interscience.wiley.com.gate1.inist.fr/journal/119% 154764/abstract, 1997. </reference>
		<reference numeration="26" content_type="text"> Kaplan, J O., Prentice, I C., and Buchmann, N.: The stable carbon isotope composition of the terrestrial biosphere: Modeling at scales from the leaf to the globe, Global Biogeochem. Cy., 16, 1–11, \doi10.1029/2001GB001403, 2002. </reference>
		<reference numeration="27" content_type="text"> Liu, W., Huang, Y., An, Z., Clemens, S C., Li, L., Prell, W L., and Ning, Y.: Summer monsoon intensity controls C&lt;sub&gt;4&lt;/sub&gt;/C&lt;sub&gt;3&lt;/sub&gt; plant abundance during the last 35 ka in the Chinese Loess Plateau: Carbon isotope evidence from bulk organic matter and individual leaf waxes, Palaeogeogr. Palaeocl., 220, 243–254, \doi10.1016/j.palaeo.2005.01.001, online available at: prefixhttp://www.sciencedirect.com.gate1.inist.fr/science/article/B% 6V6R-4FPX303-2/2/7e137eeb5316e97855f61d0cfbc23b10, 2005a. </reference>
		<reference numeration="28" content_type="text"> Liu, W., Xiahong, F., Youfeng, N., Qingle, Z., Yunning, C., and Zhisheng, A N.: $^13C$ variation of C&lt;sub&gt;3&lt;/sub&gt; and C&lt;sub&gt;4&lt;/sub&gt; plants across an Asian monsoon rainfall gradient in arid northwestern China, Glob. Change Biol., 11, 1094–1100, \doi10.1111/j.1365-2486.2005.00969.x, online available at: prefixhttp://dx.doi.org/10.1111/j.1365-2486.2005.00969.x, 2005b. </reference>
		<reference numeration="29" content_type="text"> Lloyd, J. and Farquhar, G D.: $^13$C discrimination during CO&lt;sub&gt;2&lt;/sub&gt; assimilation by the terrestrial biosphere, Oecol., 99, 201–215, \doi10.1007/BF00627732, online available at: prefixhttp://dx.doi.org/10.1007/BF00627732, 1994. </reference>
		<reference numeration="30" content_type="text"> Meyers, P A.: Preservation of elemental and isotopic source identification of sedimentary organic matter, Chem. Geol., 114, 289–302, \doi10.1016/0009-2541(94)90059-0, online available at: prefixhttp://www.sciencedirect.com/science/article/B6V5Y-488G61V-97% /2/1d2f736e88f649cb9896848d6a51d6da, 1994. </reference>
		<reference numeration="31" content_type="text"> Monnin, E., Indermuhle, A., Dallenbach, A., Fluckiger, J., Stauffer, B., Stocker, T F., Raynaud, D., and Barnola, J.-M.: Atmospheric CO&lt;sub&gt;2&lt;/sub&gt; Concentrations over the Last Glacial Termination, Science, 291, 112–114, \doi10.1126/science.291.5501.112, online available at: prefixhttp://www.sciencemag.org/cgi/content/abstract/291/5501/112, 2001. </reference>
		<reference numeration="32" content_type="text"> New, M., Lister, D., Hulme, M., and Makin, I.: A high-resolution data set of surface climate over global land areas, Climate Res., 21, 1–25, 2002. </reference>
		<reference numeration="33" content_type="text"> Prentice, I C., Cramer, W., Harrison, S P., Leemans, R., Monserud, R A., and Solomon, A M.: A global biome model based on plant physiology and dominance, soil properties and climate, J. Biogeogr., 19, 117–134, 1992. </reference>
		<reference numeration="34" content_type="text"> Sage, R F.: The evolution of C&lt;sub&gt;4&lt;/sub&gt; photosynthesis, New Phytol., 161, 341–370, \doi10.1111/j.1469-8137.2004.00974.x, online available at: prefixhttp://www.blackwell-synergy.com/doi/abs/10.1111/j.1469-8137.% 2004.00974.x, 2004. </reference>
		<reference numeration="35" content_type="text"> Schefusz, E., Schouten, S., Jansen, J. H F., and Damste, J. S S.: African vegetation controlled by tropical sea surface temperatures in the mid-Pleistocene period, Nature, 422, 418–421, \doi10.1038/nature01500, online available at: prefixhttp://dx.doi.org/10.1038/nature01500, 2003. </reference>
		<reference numeration="36" content_type="text"> Street-Perrott, F A., Huang, Y., Perrott, R A., Eglinton, G., Barker, P., Khelifa, L B., Harkness, D D., and Olago, D O.: Impact of Lower Atmospheric Carbon Dioxide on Tropical Mountain Ecosystems, Science, 278, 1422–1426, \doi10.1126/science.278.5342.1422, online available at: prefixhttp://www.sciencemag.org.gate1.inist.fr/cgi/content/abstract% /278/5342/1422, 1997. </reference>
		<reference numeration="37" content_type="text"> Taylor, D.: Late Quaternary pollen records from two Ugandan mires: evidence for environmental change in the Rukiga Highlands of southwest Uganda, Palaeogeogr. Palaeocl., 80, 283–300, 1990. </reference>
		<reference numeration="38" content_type="text"> Taylor, D.: Pollen evidence from Muchoya swamp, Rukiga Highlands (Uganda), for abrupt changes in vegetation during the last ca. 21000 years, B. Soc. Geol. Fr., 163, 77–82, 1992. </reference>
		<reference numeration="39" content_type="text"> Taylor, D.: Environmental change in montane southwest Uganda: a pollen record for the Holocene from Ahakagyezi swamp, The Holocene, 3, 324–332, 1993. </reference>
		<reference numeration="40" content_type="text"> Vincens, A.: Late Quaternary vegetation history of the South-Tanganyika Basin. Climatic implications in south central Africa, Palaeogeogr. Palaeocl., 86, 207–226, 1991. </reference>
		<reference numeration="41" content_type="text"> Wand, S. J E., Midgley, G F., Jones, M H., and Curtis, P S.: Responses of wild C&lt;sub&gt;4&lt;/sub&gt; and C&lt;sub&gt;3&lt;/sub&gt; grass (Poaceae) species to elevated atmospheric CO&lt;sub&gt;2&lt;/sub&gt; concentration: a meta-analytic test of current theories and perceptions, Global Change Biology, 5, 723–741, \doi10.1046/j.1365-2486.1999.00265.x, online available at: prefixhttp://www3.interscience.wiley.com/journal/119098780/abstract, 1999. </reference>
		<reference numeration="42" content_type="text"> Wang, G., Feng, X., Han, J., Zhou, L., Tan, W., and Su, F.: Paleovegetation reconstruction using d13C of Soil Organic Matter, Biogeosciences, 5, 1325–1337, 2008. </reference>
		<reference numeration="43" content_type="text"> Ward, J K., Tissue, D T., Thomas, R B., and Strain, B R.: Comparative responses of model C&lt;sub&gt;3&lt;/sub&gt; and C&lt;sub&gt;4&lt;/sub&gt; plants to drought in low and elevated CO&lt;sub&gt;2&lt;/sub&gt;, Glob. Change Biol., 5, 857–867, 1999. </reference>
		<reference numeration="44" content_type="text"> Yu, G., Chen, X., Ni, J., Cheddadi, R., Guiot, J., Han, H., Harrison, S P., Huang, C., Ke, M., Kong, Z., Li, S., Li, W., Liew, P., Liu, G., Liu, J., Liu, Q., Liu, K., Prentice, I C., Qui, W., Ren, G., Song, C., Sugita, S., Sun, X., Tang, L., Campo, E V., Xia, Y., Xu, Q., Yan, S., Yang, X., Zhao, J., and Zheng, Z.: Palaeovegetation of China: a pollen data-based synthesis for the mid-Holocene and last glacial maximum, J. Biogeogr., 27, 635–664, \doi10.1046/j.1365-2699.2000.00431.x, online available at: prefixhttp://dx.doi.org/10.1046/j.1365-2699.2000.00431.x, 2000. </reference>
		<reference numeration="45" content_type="text"> Zhang, Z., Zhao, M., Lu, H., and Faiia, A M.: Lower temperature as the main cause of C4 plant declines during the glacial periods on the Chinese Loess Plateau, Earth Planet. Sci. Lett., 214, 467–481, \doi10.1016/S0012-821X(03)00387-X, online available at: prefixhttp://www.sciencedirect.com.gate1.inist.fr/science/article/B% 6V61-49CR9VN-3/2/5c68ec345fcd900c4a6afe8cfba8b157, 2003. </reference>
		<reference numeration="46" content_type="text"> Zheng, Z., Baoyin, Y., and Petit-Maire, N.: Paleoenvironments in China during the Last Glacial Maximum and the Holocene optimum, Episodes, 21, 152–158, 1998. </reference>
	</references>
</article>

