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<article language="en">
	<journal>
		<journal_title>Biogeosciences Discussions</journal_title>
		<journal_url>www.biogeosciences-discuss.net</journal_url>
		<issn>1810-6277</issn>
		<eissn>1810-6285</eissn>
		<volume_number>6</volume_number>
		<issue_number>2</issue_number>
		<publication_year>2009</publication_year>
	</journal>
	<doi>10.5194/bgd-6-3707-2009</doi>
	<article_url>http://www.biogeosciences-discuss.net/6/3707/2009/</article_url>
	<abstract_html>http://www.biogeosciences-discuss.net/6/3707/2009/bgd-6-3707-2009.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences-discuss.net/6/3707/2009/bgd-6-3707-2009.pdf</fulltext_pdf>
	<start_page>3707</start_page>
	<end_page>3769</end_page>
	<publication_date>2009-04-06</publication_date>
	<article_title content_type="html">Basin-wide variations in foliar properties of Amazonian forest: phylogeny, soils and climate</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>N. M. Fyllas</name>
			<email>n.fyllas@leeds.ac.uk</email>
		</author>
		<author numeration="2" affiliations="2,3">
			<name>S. PatiÃ±o</name>
		</author>
		<author numeration="3" affiliations="1">
			<name>T. R. Baker</name>
		</author>
		<author numeration="4" affiliations="4">
			<name>G. Bielefeld Nardoto</name>
		</author>
		<author numeration="5" affiliations="4">
			<name>L. A. Martinelli</name>
		</author>
		<author numeration="6" affiliations="1,5,10">
			<name>C. A. Quesada</name>
		</author>
		<author numeration="7" affiliations="5,6">
			<name>R. Paiva</name>
		</author>
		<author numeration="8" affiliations="7">
			<name>M. Schwarz</name>
		</author>
		<author numeration="9" affiliations="8">
			<name>V. Horna</name>
		</author>
		<author numeration="10" affiliations="9">
			<name>L. M. Mercado</name>
		</author>
		<author numeration="11" affiliations="5,10">
			<name>A. Santos</name>
		</author>
		<author numeration="12" affiliations="11">
			<name>L. Arroyo</name>
		</author>
		<author numeration="13" affiliations="12">
			<name>E. M. JimÃ©nez</name>
		</author>
		<author numeration="14" affiliations="5">
			<name>F. J. LuizÃ£o</name>
		</author>
		<author numeration="15" affiliations="13">
			<name>D. A. Neill</name>
		</author>
		<author numeration="16" affiliations="14">
			<name>N. Silva</name>
		</author>
		<author numeration="17" affiliations="2">
			<name>A. Prieto</name>
		</author>
		<author numeration="18" affiliations="15">
			<name>A. Rudas</name>
		</author>
		<author numeration="19" affiliations="16">
			<name>M. Silviera</name>
		</author>
		<author numeration="20" affiliations="17">
			<name>I. C. G. Vieira</name>
		</author>
		<author numeration="21" affiliations="1">
			<name>G. Lopez-Gonzalez</name>
		</author>
		<author numeration="22" affiliations="18">
			<name>Y. Malhi</name>
		</author>
		<author numeration="23" affiliations="1">
			<name>O. L. Phillips</name>
		</author>
		<author numeration="24" affiliations="1">
			<name>J. Lloyd</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Earth and Biosphere Institute, School of Geography, University of Leeds, LS2 9JT, UK</affiliation>
		<affiliation numeration="2" content_type="html">Instituto de InvestigaciÃ³n de Recursos BiolÃ³gicos, Alexander von Humboldt, BogotÃ¡, Colombia</affiliation>
		<affiliation numeration="3" content_type="html">UMR-ECOFOG, INRA, 97310, Korou, French Guiana</affiliation>
		<affiliation numeration="4" content_type="html">CENA-USP, Av CentenÃ¡rio 303, 13416-000 Piracicaba, SP, Brazil</affiliation>
		<affiliation numeration="5" content_type="html">Instituto National de Pesquisas AmazÃ´nicas, Manaus, Brazil</affiliation>
		<affiliation numeration="6" content_type="html">SecretÃ¡ria Municipal de Desenvolvimento e Meio Ammbiente ma Prefeturia Municipal de MauÃ©s, MauÃ©s, Brazil</affiliation>
		<affiliation numeration="7" content_type="html">Max-Planck-Institut fuer Biogeochemie, Jena, Germany</affiliation>
		<affiliation numeration="8" content_type="html">Abteilung Ã–kologie und Ã–kosystemforschung, Albrecht-von-Haller-Institut fÃ¼r Pflanzenwissenschaften, UniversitÃ¤t GÃ¶ttingen, GÃ¶ttingen, Germany</affiliation>
		<affiliation numeration="9" content_type="html">Centre for Ecology and Hydrology, Wallingford, UK</affiliation>
		<affiliation numeration="10" content_type="html">Departamento de Ecologia, Universidade de BrasÃ­lia, Brazil</affiliation>
		<affiliation numeration="11" content_type="html">Museo Noel Kempff Mercado, Santa Cruz, Bolivia</affiliation>
		<affiliation numeration="12" content_type="html">Universidad Nacional de Colombia, Leticia,Colombia</affiliation>
		<affiliation numeration="13" content_type="html">Herbario Nacional del Ecuador, Quito, Ecuador</affiliation>
		<affiliation numeration="14" content_type="html">Empera Brasileira de Pesquisas AgropecuÃ¡ria, Belem, Brazil</affiliation>
		<affiliation numeration="15" content_type="html">Instituto de Ciencias Naturales, Universidad Nacional de Colombia, BogotÃ¡, Colombia</affiliation>
		<affiliation numeration="16" content_type="html">Depto de CiÃªncias da Natureza, Universidade Federal do Acre, Rio Branco, Brazil</affiliation>
		<affiliation numeration="17" content_type="html">Museu Paraense Emilio Goeldi Caixa Postal 399 BelÃ©m, ParÃ¡ 66040-170, Brazil</affiliation>
		<affiliation numeration="18" content_type="html">University of Oxford, Oxford, UK</affiliation>
	</affiliations>
	<abstract content_type="html">We analysed 1040 individual trees, positioned in sixty three plots across
the Amazon Basin for leaf mass per area (&lt;i&gt;M&lt;sub&gt;A&lt;/sub&gt;&lt;/i&gt;), leaf carbon isotope
composition (&amp;delta;&lt;sup&gt;13&lt;/sup&gt;C) and leaf level concentrations of C, N, P,
Ca, Mg, K and Al. All trees were identified to the species with the dataset
containing 58 families, 236 genera and 508 species, distributed across a
wide range of soil types and precipitation regimes. Some foliar characters
such as &lt;i&gt;M&lt;sub&gt;A&lt;/sub&gt;&lt;/i&gt;, [C], [N] and [Mg] emerge as highly constrained by the taxonomic
affiliation of tree species, but with others such as [P], [K], [Ca] and
&amp;delta;&lt;sup&gt;13&lt;/sup&gt;C also strongly influenced by site growing conditions. By
removing the environmental contribution to trait variation, we find that
intrinsic values of most trait pairs coordinate, although different species
(characterised by different trait suites) are found at discrete locations
along a common axis of coordination. Species that tend to occupy higher
fertility soils are characterised by a lower &lt;i&gt;M&lt;sub&gt;A&lt;/sub&gt;&lt;/i&gt; and have a higher
intrinsic [N], [P], [K], [Mg] and &amp;delta;&lt;sup&gt;13&lt;/sup&gt;C than their lower
fertility counterparts. Despite this consistency, different scaling patterns
were observed between low and high fertility sites. Inter-relationships are
thus substantially modified by growth environment. Analysing the
environmental component of trait variation, we found soil fertility to be
the most important predictor, influencing all leaf nutrient concentrations
and &amp;delta;&lt;sup&gt;13&lt;/sup&gt;C composition and reducing &lt;i&gt;M&lt;sub&gt;A&lt;/sub&gt;&lt;/i&gt;. Mean annual
temperature was negatively associated with leaf level [N], [P] and [K]
concentrations. Total annual precipitation positively influences &lt;i&gt;M&lt;sub&gt;A&lt;/sub&gt;&lt;/i&gt;, [C]
and &amp;delta;&lt;sup&gt;13&lt;/sup&gt;C, but with a negative impact on [Mg]. These results
provide a first basis for understanding the relationship between the
physiological functioning and distribution of tree species across Amazonia.</abstract>
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