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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Evolutionary heritage influences Amazon tree ecology</title><author><name sortKey="Coelho De Souza, Fernanda" sort="Coelho De Souza, Fernanda" uniqKey="Coelho De Souza F" first="Fernanda" last="Coelho De Souza">Fernanda Coelho De Souza</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Dexter, Kyle G" sort="Dexter, Kyle G" uniqKey="Dexter K" first="Kyle G." last="Dexter">Kyle G. Dexter</name><affiliation><nlm:aff id="af2"><addr-line>School of Geosciences</addr-line>,<institution>University of Edinburgh</institution>,<addr-line>201 Crew Building, King's Buildings, Edinburgh EH9 3FF</addr-line>,<country>UK</country></nlm:aff></affiliation><affiliation><nlm:aff id="af3"><addr-line>Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Phillips, Oliver L" sort="Phillips, Oliver L" uniqKey="Phillips O" first="Oliver L." last="Phillips">Oliver L. Phillips</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Brienen, Roel J W" sort="Brienen, Roel J W" uniqKey="Brienen R" first="Roel J. W." last="Brienen">Roel J. W. Brienen</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Chave, Jerome" sort="Chave, Jerome" uniqKey="Chave J" first="Jerome" last="Chave">Jerome Chave</name><affiliation><nlm:aff id="af4"><institution>Université Paul Sabatier CNRS, UMR 5174 Evolution et Diversité Biologique</institution>,<addr-line>bâtiment 4R1, Toulouse 31062</addr-line>,<country>France</country></nlm:aff></affiliation></author><author><name sortKey="Galbraith, David R" sort="Galbraith, David R" uniqKey="Galbraith D" first="David R." last="Galbraith">David R. Galbraith</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Lopez Gonzalez, Gabriela" sort="Lopez Gonzalez, Gabriela" uniqKey="Lopez Gonzalez G" first="Gabriela" last="Lopez Gonzalez">Gabriela Lopez Gonzalez</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Monteagudo Mendoza, Abel" sort="Monteagudo Mendoza, Abel" uniqKey="Monteagudo Mendoza A" first="Abel" last="Monteagudo Mendoza">Abel Monteagudo Mendoza</name><affiliation><nlm:aff id="af5"><addr-line>Jardín Botánico de Missouri</addr-line>,<institution>Prolongacion Bolognesi Mz. E</institution>,<addr-line>Lote 6, Oxapampa, Pasco</addr-line>,<country>Peru</country></nlm:aff></affiliation><affiliation><nlm:aff id="af40"><institution>Universidad Nacional de San Antonio Abad del Cusco</institution>,<addr-line>Av. de la Cultura N° 733, Cusco</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="Pennington, R Toby" sort="Pennington, R Toby" uniqKey="Pennington R" first="R. Toby" last="Pennington">R. Toby Pennington</name><affiliation><nlm:aff id="af3"><addr-line>Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Poorter, Lourens" sort="Poorter, Lourens" uniqKey="Poorter L" first="Lourens" last="Poorter">Lourens Poorter</name><affiliation><nlm:aff id="af6"><addr-line>Forest Ecology and Forest Management Group</addr-line>,<institution>Wageningen University and Research</institution>,<addr-line>PO Box 47, 6700 AA Wageningen</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Alexiades, Miguel" sort="Alexiades, Miguel" uniqKey="Alexiades M" first="Miguel" last="Alexiades">Miguel Alexiades</name><affiliation><nlm:aff id="af7"><addr-line>School of Anthropology and Conservation</addr-line>,<institution>University of Kent</institution>,<addr-line>Marlowe Building, Canterbury, Kent CT2 7NR</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Alvarez Davila, Esteban" sort="Alvarez Davila, Esteban" uniqKey="Alvarez Davila E" first="Esteban" last="Álvarez-Dávila">Esteban Álvarez-Dávila</name><affiliation><nlm:aff id="af8"><institution>Fundación Con Vida</institution>,<addr-line>Medellin Cra 48: 20-114</addr-line>,<country>Colombia</country></nlm:aff></affiliation></author><author><name sortKey="Andrade, Ana" sort="Andrade, Ana" uniqKey="Andrade A" first="Ana" last="Andrade">Ana Andrade</name><affiliation><nlm:aff id="af9"><addr-line>Biological Dynamics of Forest Fragment Project (INPA & STRI), C.P. 478, Manaus, Amazonas 69.011-970</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Aragao, Luis E O C" sort="Aragao, Luis E O C" uniqKey="Aragao L" first="Luis E. O. C." last="Aragão">Luis E. O. C. Aragão</name><affiliation><nlm:aff id="af10"><addr-line>Geography, College of Life and Environmental Sciences</addr-line>,<institution>University of Exeter, Drive, Exeter</institution>,<addr-line>Rennes EX4 4RJ</addr-line>,<country>UK</country></nlm:aff></affiliation><affiliation><nlm:aff id="af11"><institution>National Institute for Space Research (INPE)</institution>,<addr-line>São José dos Campos, São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Araujo Murakami, Alejandro" sort="Araujo Murakami, Alejandro" uniqKey="Araujo Murakami A" first="Alejandro" last="Araujo-Murakami">Alejandro Araujo-Murakami</name><affiliation><nlm:aff id="af12"><addr-line>Museo de Historia Natural Noel Kempff Mercado</addr-line>,<institution>Universidad Autonoma Gabriel Rene Moreno</institution>,<addr-line>Casilla 2489, Av. Irala 565, Santa Cruz</addr-line>,<country>Bolivia</country></nlm:aff></affiliation></author><author><name sortKey="Arets, Eric J M M" sort="Arets, Eric J M M" uniqKey="Arets E" first="Eric J. M. M." last="Arets">Eric J. M. M. Arets</name><affiliation><nlm:aff id="af13"><addr-line>Alterra</addr-line>,<institution>Wageningen University and Research Centre</institution>,<addr-line>PO Box 47, Wageningen 6700 AA</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Aymard C, Gerardo A" sort="Aymard C, Gerardo A" uniqKey="Aymard C G" first="Gerardo A." last="Aymard C">Gerardo A. Aymard C</name><affiliation><nlm:aff id="af14"><addr-line>UNELLEZ-Guanare, Programa del Agro y del Mar</addr-line>,<institution>Herbario Universitario (PORT)</institution>,<addr-line>Mesa de Cavacas, Estado Portuguesa 3350</addr-line>,<country>Venezuela</country></nlm:aff></affiliation></author><author><name sortKey="Baraloto, Christopher" sort="Baraloto, Christopher" uniqKey="Baraloto C" first="Christopher" last="Baraloto">Christopher Baraloto</name><affiliation><nlm:aff id="af15"><addr-line>International Center for Tropical Botany, Department of Biological Sciences</addr-line>,<institution>Florida International University</institution>,<addr-line>Miami, FL 33199</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Barroso, Jorcely G" sort="Barroso, Jorcely G" uniqKey="Barroso J" first="Jorcely G." last="Barroso">Jorcely G. Barroso</name><affiliation><nlm:aff id="af16"><institution>Universidade Federal do Acre, Campus de Cruzeiro do Sul</institution>,<country>Acre, Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Bonal, Damien" sort="Bonal, Damien" uniqKey="Bonal D" first="Damien" last="Bonal">Damien Bonal</name><affiliation><nlm:aff id="af17"><institution>INRA, UMR 1137 ‘Ecologie et Ecophysiologie Forestiere’</institution>,<addr-line>Champenoux 54280</addr-line>,<country>France</country></nlm:aff></affiliation></author><author><name sortKey="Boot, Rene G A" sort="Boot, Rene G A" uniqKey="Boot R" first="Rene G. A." last="Boot">Rene G. A. Boot</name><affiliation><nlm:aff id="af18"><addr-line>Tropenbos International, PO Box 232, Wageningen 6700 AE</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Camargo, Jose L C" sort="Camargo, Jose L C" uniqKey="Camargo J" first="José L. C." last="Camargo">José L. C. Camargo</name><affiliation><nlm:aff id="af9"><addr-line>Biological Dynamics of Forest Fragment Project (INPA & STRI), C.P. 478, Manaus, Amazonas 69.011-970</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Comiskey, James A" sort="Comiskey, James A" uniqKey="Comiskey J" first="James A." last="Comiskey">James A. Comiskey</name><affiliation><nlm:aff id="af19"><addr-line>National Park Service, 120 Chatham Lane, Fredericksburg, VA 22405</addr-line>,<country>USA</country></nlm:aff></affiliation><affiliation><nlm:aff id="af20"><institution>Smithsonian Institution</institution>,<addr-line>1100 Jefferson Dr, SW, Washington, DC 20560</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Valverde, Fernando Cornejo" sort="Valverde, Fernando Cornejo" uniqKey="Valverde F" first="Fernando Cornejo" last="Valverde">Fernando Cornejo Valverde</name><affiliation><nlm:aff id="af21"><addr-line>Proyecto Castaña, Madre de Dios</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="De Camargo, Plinio B" sort="De Camargo, Plinio B" uniqKey="De Camargo P" first="Plínio B." last="De Camargo">Plínio B. De Camargo</name><affiliation><nlm:aff id="af22"><addr-line>Centro de Energia Nuclear na Agricultura</addr-line>,<institution>Universidade de São Paulo</institution>,<addr-line>São Paulo, Sao Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Di Fiore, Anthony" sort="Di Fiore, Anthony" uniqKey="Di Fiore A" first="Anthony" last="Di Fiore">Anthony Di Fiore</name><affiliation><nlm:aff id="af23"><addr-line>Department of Anthropology</addr-line>,<institution>University of Texas at Austin</institution>,<addr-line>SAC Room 5.150, 2201 Speedway Stop C3200, Austin, TX 78712</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Elias, Fernando" sort="Elias, Fernando" uniqKey="Elias F" first="Fernando" last="Elias">Fernando Elias</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Erwin, Terry L" sort="Erwin, Terry L" uniqKey="Erwin T" first="Terry L." last="Erwin">Terry L. Erwin</name><affiliation><nlm:aff id="af25"><addr-line>Department of Entomology</addr-line>,<institution>Smithsonian Institution</institution>,<addr-line>PO Box 37012</addr-line>, MRC 187,<addr-line>Washington, DC 20013-7012</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Feldpausch, Ted R" sort="Feldpausch, Ted R" uniqKey="Feldpausch T" first="Ted R." last="Feldpausch">Ted R. Feldpausch</name><affiliation><nlm:aff id="af10"><addr-line>Geography, College of Life and Environmental Sciences</addr-line>,<institution>University of Exeter, Drive, Exeter</institution>,<addr-line>Rennes EX4 4RJ</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Ferreira, Leandro" sort="Ferreira, Leandro" uniqKey="Ferreira L" first="Leandro" last="Ferreira">Leandro Ferreira</name><affiliation><nlm:aff id="af26"><addr-line>Museu Paraense Emilio Goeldi, C.P. 399, 66.040-170, Belém, Pará</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Fyllas, Nikolaos M" sort="Fyllas, Nikolaos M" uniqKey="Fyllas N" first="Nikolaos M." last="Fyllas">Nikolaos M. Fyllas</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Gloor, Emanuel" sort="Gloor, Emanuel" uniqKey="Gloor E" first="Emanuel" last="Gloor">Emanuel Gloor</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Herault, Bruno" sort="Herault, Bruno" uniqKey="Herault B" first="Bruno" last="Herault">Bruno Herault</name><affiliation><nlm:aff id="af27"><institution>Cirad, UMR EcoFoG (AgroParisTech, CNRS, Inra, U Antilles, U Guyane)</institution>,<addr-line>Campus Agronomique, Kourou 97310</addr-line>,<country>French Guiana</country></nlm:aff></affiliation></author><author><name sortKey="Herrera, Rafael" sort="Herrera, Rafael" uniqKey="Herrera R" first="Rafael" last="Herrera">Rafael Herrera</name><affiliation><nlm:aff id="af28"><institution>Centro de Ecología IVIC</institution>,<addr-line>Caracas</addr-line>,<country>Venezuela</country></nlm:aff></affiliation><affiliation><nlm:aff id="af29"><addr-line>Institut für Geographie und Regionalforschung, University of Vienna, Wien</addr-line>,<country>Austria</country></nlm:aff></affiliation></author><author><name sortKey="Higuchi, Niro" sort="Higuchi, Niro" uniqKey="Higuchi N" first="Niro" last="Higuchi">Niro Higuchi</name><affiliation><nlm:aff id="af30"><institution>INPA</institution>,<addr-line>Av. André Araújo, 2.936 - Petrópolis – 69.067-375, Manaus, Amazonas</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Honorio Coronado, Euridice N" sort="Honorio Coronado, Euridice N" uniqKey="Honorio Coronado E" first="Eurídice N." last="Honorio Coronado">Eurídice N. Honorio Coronado</name><affiliation><nlm:aff id="af31"><institution>Instituto de Investigaciones de la Amazonia Peruana</institution>,<addr-line>Apartado 784, Iquitos</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="Killeen, Timothy J" sort="Killeen, Timothy J" uniqKey="Killeen T" first="Timothy J." last="Killeen">Timothy J. Killeen</name><affiliation><nlm:aff id="af32"><institution>AGTECA - Amazónica</institution>,<addr-line>Santa Cruz</addr-line>,<country>Bolivia</country></nlm:aff></affiliation></author><author><name sortKey="Laurance, William F" sort="Laurance, William F" uniqKey="Laurance W" first="William F." last="Laurance">William F. Laurance</name><affiliation><nlm:aff id="af33"><addr-line>Centre for Tropical Environmental and Sustainability Science (TESS) and College of Science and Engineering</addr-line>,<institution>James Cook University</institution>,<addr-line>Cairns, Queensland 4878</addr-line>,<country>Australia</country></nlm:aff></affiliation></author><author><name sortKey="Laurance, Susan" sort="Laurance, Susan" uniqKey="Laurance S" first="Susan" last="Laurance">Susan Laurance</name><affiliation><nlm:aff id="af33"><addr-line>Centre for Tropical Environmental and Sustainability Science (TESS) and College of Science and Engineering</addr-line>,<institution>James Cook University</institution>,<addr-line>Cairns, Queensland 4878</addr-line>,<country>Australia</country></nlm:aff></affiliation></author><author><name sortKey="Lloyd, Jon" sort="Lloyd, Jon" uniqKey="Lloyd J" first="Jon" last="Lloyd">Jon Lloyd</name><affiliation><nlm:aff id="af34"><addr-line>Department of Life Sciences</addr-line>,<institution>Imperial College London</institution>,<addr-line>Silwood Park Campus, Buckhurst, Road, Ascot, Berkshire SL5 7PY</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Lovejoy, Thomas E" sort="Lovejoy, Thomas E" uniqKey="Lovejoy T" first="Thomas E." last="Lovejoy">Thomas E. Lovejoy</name><affiliation><nlm:aff id="af35"><addr-line>Environmental Science and Policy, and the Department of Public and International Affairs</addr-line>,<institution>George Mason University (GMU)</institution>,<addr-line>Washington, DC</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Malhi, Yadvinder" sort="Malhi, Yadvinder" uniqKey="Malhi Y" first="Yadvinder" last="Malhi">Yadvinder Malhi</name><affiliation><nlm:aff id="af36"><addr-line>Environmental Change Institute, School of Geography and the Environment</addr-line>,<institution>University of Oxford</institution>,<addr-line>Oxford</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Maracahipes, Leandro" sort="Maracahipes, Leandro" uniqKey="Maracahipes L" first="Leandro" last="Maracahipes">Leandro Maracahipes</name><affiliation><nlm:aff id="af37"><addr-line>Programa de Pós-graduação em Ecologia e Evolução</addr-line>,<institution>Universidade Federal de Goias</institution>,<addr-line>Goiânia, Goias</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Marimon, Beatriz S" sort="Marimon, Beatriz S" uniqKey="Marimon B" first="Beatriz S." last="Marimon">Beatriz S. Marimon</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Marimon Junior, Ben H" sort="Marimon Junior, Ben H" uniqKey="Marimon Junior B" first="Ben H." last="Marimon-Junior">Ben H. Marimon-Junior</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Mendoza, Casimiro" sort="Mendoza, Casimiro" uniqKey="Mendoza C" first="Casimiro" last="Mendoza">Casimiro Mendoza</name><affiliation><nlm:aff id="af38"><addr-line>Escuela de Ciencias Forestales, Unidad Académica del Trópico</addr-line>,<institution>Universidad Mayor de San Simón</institution>,<addr-line>Sacta</addr-line>,<country>Bolivia</country></nlm:aff></affiliation></author><author><name sortKey="Morandi, Paulo" sort="Morandi, Paulo" uniqKey="Morandi P" first="Paulo" last="Morandi">Paulo Morandi</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Neill, David A" sort="Neill, David A" uniqKey="Neill D" first="David A." last="Neill">David A. Neill</name><affiliation><nlm:aff id="af39"><addr-line>Universidad Estatal Amazónica, Puyo, Pastaza</addr-line>,<country>Ecuador</country></nlm:aff></affiliation></author><author><name sortKey="Vargas, Percy Nu Ez" sort="Vargas, Percy Nu Ez" uniqKey="Vargas P" first="Percy Nú Ez" last="Vargas">Percy Nú Ez Vargas</name><affiliation><nlm:aff id="af40"><institution>Universidad Nacional de San Antonio Abad del Cusco</institution>,<addr-line>Av. de la Cultura N° 733, Cusco</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="Oliveira, Edmar A" sort="Oliveira, Edmar A" uniqKey="Oliveira E" first="Edmar A." last="Oliveira">Edmar A. Oliveira</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Lenza, Eddie" sort="Lenza, Eddie" uniqKey="Lenza E" first="Eddie" last="Lenza">Eddie Lenza</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Palacios, Walter A" sort="Palacios, Walter A" uniqKey="Palacios W" first="Walter A." last="Palacios">Walter A. Palacios</name><affiliation><nlm:aff id="af41"><institution>Universidad Técnica del Norte and Herbario Nacional del Ecuador</institution>,<addr-line>Casilla 17-21-1787, Av. Río Coca E6-115, Quito</addr-line>,<country>Ecuador</country></nlm:aff></affiliation></author><author><name sortKey="Pe Uela Mora, Maria C" sort="Pe Uela Mora, Maria C" uniqKey="Pe Uela Mora M" first="Maria C." last="Pe Uela-Mora">Maria C. Pe Uela-Mora</name><affiliation><nlm:aff id="af42"><institution>Universidad Regional Amazónica IKIAM</institution>,<addr-line>Tena</addr-line>,<country>Ecuador</country></nlm:aff></affiliation></author><author><name sortKey="Pipoly, John J" sort="Pipoly, John J" uniqKey="Pipoly J" first="John J." last="Pipoly">John J. Pipoly</name><affiliation><nlm:aff id="af43"><addr-line>Broward County Parks and Recreation Division, 950 NW 38th St., Oakland Park, FL 33309</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Pitman, Nigel C A" sort="Pitman, Nigel C A" uniqKey="Pitman N" first="Nigel C. A." last="Pitman">Nigel C. A. Pitman</name><affiliation><nlm:aff id="af44"><addr-line>Center for Tropical Conservation</addr-line>,<institution>Duke University</institution>,<addr-line>PO Box 90381, Durham, NC 27708</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Prieto, Adriana" sort="Prieto, Adriana" uniqKey="Prieto A" first="Adriana" last="Prieto">Adriana Prieto</name><affiliation><nlm:aff id="af45"><addr-line>Doctorado Instituto de Ciencias Naturales</addr-line>,<institution>Universidad ciol de Colombia</institution>,<country>Colombia</country></nlm:aff></affiliation></author><author><name sortKey="Quesada, Carlos A" sort="Quesada, Carlos A" uniqKey="Quesada C" first="Carlos A." last="Quesada">Carlos A. Quesada</name><affiliation><nlm:aff id="af30"><institution>INPA</institution>,<addr-line>Av. André Araújo, 2.936 - Petrópolis – 69.067-375, Manaus, Amazonas</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Ramirez Angulo, Hirma" sort="Ramirez Angulo, Hirma" uniqKey="Ramirez Angulo H" first="Hirma" last="Ramirez-Angulo">Hirma Ramirez-Angulo</name><affiliation><nlm:aff id="af46"><addr-line>Instituto de Investigaciones para el Desarrollo Forestal (INDEFOR), Facultad de Ciencias Forestales y Ambientales</addr-line>,<institution>Universidad de Los Andes</institution>,<addr-line>Conjunto Forestal, C.P. 5101, Mérida</addr-line>,<country>Venezuela</country></nlm:aff></affiliation></author><author><name sortKey="Rudas, Agustin" sort="Rudas, Agustin" uniqKey="Rudas A" first="Agustin" last="Rudas">Agustin Rudas</name><affiliation><nlm:aff id="af45"><addr-line>Doctorado Instituto de Ciencias Naturales</addr-line>,<institution>Universidad ciol de Colombia</institution>,<country>Colombia</country></nlm:aff></affiliation></author><author><name sortKey="Ruokolainen, Kalle" sort="Ruokolainen, Kalle" uniqKey="Ruokolainen K" first="Kalle" last="Ruokolainen">Kalle Ruokolainen</name><affiliation><nlm:aff id="af47"><addr-line>Department of Geography and Geology</addr-line>,<institution>University of Turku, 20014 Turku</institution>,<country>Finland</country></nlm:aff></affiliation></author><author><name sortKey="Salomao, Rafael P" sort="Salomao, Rafael P" uniqKey="Salomao R" first="Rafael P." last="Salomão">Rafael P. Salomão</name><affiliation><nlm:aff id="af26"><addr-line>Museu Paraense Emilio Goeldi, C.P. 399, 66.040-170, Belém, Pará</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Silveira, Marcos" sort="Silveira, Marcos" uniqKey="Silveira M" first="Marcos" last="Silveira">Marcos Silveira</name><affiliation><nlm:aff id="af48"><institution>Museu Universitário, Universidade Federal do Acre</institution>,<addr-line>Rio Branco, AC 69910-900</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Stropp, Juliana" sort="Stropp, Juliana" uniqKey="Stropp J" first="Juliana" last="Stropp">Juliana Stropp</name><affiliation><nlm:aff id="af49"><addr-line>Institute of Biological and Health Sciences (ICBS)</addr-line>,<institution>Federal University of Alagoas</institution>,<addr-line>Maceió, AL</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Ter Steege, Hans" sort="Ter Steege, Hans" uniqKey="Ter Steege H" first="Hans" last="Ter Steege">Hans Ter Steege</name><affiliation><nlm:aff id="af50"><institution>Naturalis Biodiversity Center</institution>,<addr-line>Vondellaan 55, Postbus 9517, Leiden 2300 RA</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Thomas Caesar, Raquel" sort="Thomas Caesar, Raquel" uniqKey="Thomas Caesar R" first="Raquel" last="Thomas-Caesar">Raquel Thomas-Caesar</name><affiliation><nlm:aff id="af51"><institution>Iwokrama Intertiol Centre for Rainforest Conservation and Development</institution>,<addr-line>77 High Street Kingston, Georgetown</addr-line>,<country>Guyana</country></nlm:aff></affiliation></author><author><name sortKey="Van Der Hout, Peter" sort="Van Der Hout, Peter" uniqKey="Van Der Hout P" first="Peter" last="Van Der Hout">Peter Van Der Hout</name><affiliation><nlm:aff id="af52"><addr-line>Van der Hout Forestry Consulting, Jan Trooststraat 6, Rotterdam 3078 HP</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Van Der Heijden, Geertje M F" sort="Van Der Heijden, Geertje M F" uniqKey="Van Der Heijden G" first="Geertje M. F." last="Van Der Heijden">Geertje M. F. Van Der Heijden</name><affiliation><nlm:aff id="af53"><addr-line>School of Geography</addr-line>,<institution>University of Nottingham</institution>,<addr-line>University Park, Nottingham NG7 2RD</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Van Der Meer, Peter J" sort="Van Der Meer, Peter J" uniqKey="Van Der Meer P" first="Peter J." last="Van Der Meer">Peter J. Van Der Meer</name><affiliation><nlm:aff id="af54"><institution>Van Hall Larenstein University of Applied Sciences</institution>,<addr-line>PO Box 9001, 6880 GB Velp</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Vasquez, Rodolfo V" sort="Vasquez, Rodolfo V" uniqKey="Vasquez R" first="Rodolfo V." last="Vasquez">Rodolfo V. Vasquez</name><affiliation><nlm:aff id="af5"><addr-line>Jardín Botánico de Missouri</addr-line>,<institution>Prolongacion Bolognesi Mz. E</institution>,<addr-line>Lote 6, Oxapampa, Pasco</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="Vieira, Simone A" sort="Vieira, Simone A" uniqKey="Vieira S" first="Simone A." last="Vieira">Simone A. Vieira</name><affiliation><nlm:aff id="af55"><addr-line>Universidade Estadual de Campinas</addr-line>,<institution>Núcleo de Estudos e Pesquisas Ambientais – NEPAM</institution>,<addr-line>Campinas, São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Vilanova, Emilio" sort="Vilanova, Emilio" uniqKey="Vilanova E" first="Emilio" last="Vilanova">Emilio Vilanova</name><affiliation><nlm:aff id="af56"><addr-line>Facultad de Ciencias Forestales y Ambientales</addr-line>,<institution>Universidad de Los Andes</institution>,<addr-line>Mérida</addr-line>,<country>Venezuela</country></nlm:aff></affiliation></author><author><name sortKey="Vos, Vincent A" sort="Vos, Vincent A" uniqKey="Vos V" first="Vincent A." last="Vos">Vincent A. Vos</name><affiliation><nlm:aff id="af57"><institution>Centro de Investigación y Promoción del Campesinado - regional Norte Amazónico</institution>,<addr-line>C/ Nicanor Gonzalo Salvatierra N° 362, Casilla 16, Riberalta</addr-line>,<country>Bolivia</country></nlm:aff></affiliation><affiliation><nlm:aff id="af58"><institution>Universidad Autónoma del Beni</institution>,<addr-line>Campus Universitario, Riberalta</addr-line>,<country>Bolivia</country></nlm:aff></affiliation></author><author><name sortKey="Wang, Ophelia" sort="Wang, Ophelia" uniqKey="Wang O" first="Ophelia" last="Wang">Ophelia Wang</name><affiliation><nlm:aff id="af59"><institution>Northern Arizona University</institution>,<addr-line>Flagstaff, AZ 86011</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Young, Kenneth R" sort="Young, Kenneth R" uniqKey="Young K" first="Kenneth R." last="Young">Kenneth R. Young</name><affiliation><nlm:aff id="af60"><addr-line>Department of Geography and the Environment</addr-line>,<institution>University of Texas at Austin</institution>,<addr-line>Austin, TX 78712</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Zagt, Roderick J" sort="Zagt, Roderick J" uniqKey="Zagt R" first="Roderick J." last="Zagt">Roderick J. Zagt</name><affiliation><nlm:aff id="af18"><addr-line>Tropenbos International, PO Box 232, Wageningen 6700 AE</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Baker, Timothy R" sort="Baker, Timothy R" uniqKey="Baker T" first="Timothy R." last="Baker">Timothy R. Baker</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">27974517</idno><idno type="pmc">5204144</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5204144</idno><idno type="RBID">PMC:5204144</idno><idno type="doi">10.1098/rspb.2016.1587</idno><date when="2016">2016</date><idno type="wicri:Area/Pmc/Corpus">002C92</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">002C92</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Evolutionary heritage influences Amazon tree ecology</title><author><name sortKey="Coelho De Souza, Fernanda" sort="Coelho De Souza, Fernanda" uniqKey="Coelho De Souza F" first="Fernanda" last="Coelho De Souza">Fernanda Coelho De Souza</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Dexter, Kyle G" sort="Dexter, Kyle G" uniqKey="Dexter K" first="Kyle G." last="Dexter">Kyle G. Dexter</name><affiliation><nlm:aff id="af2"><addr-line>School of Geosciences</addr-line>,<institution>University of Edinburgh</institution>,<addr-line>201 Crew Building, King's Buildings, Edinburgh EH9 3FF</addr-line>,<country>UK</country></nlm:aff></affiliation><affiliation><nlm:aff id="af3"><addr-line>Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Phillips, Oliver L" sort="Phillips, Oliver L" uniqKey="Phillips O" first="Oliver L." last="Phillips">Oliver L. Phillips</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Brienen, Roel J W" sort="Brienen, Roel J W" uniqKey="Brienen R" first="Roel J. W." last="Brienen">Roel J. W. Brienen</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Chave, Jerome" sort="Chave, Jerome" uniqKey="Chave J" first="Jerome" last="Chave">Jerome Chave</name><affiliation><nlm:aff id="af4"><institution>Université Paul Sabatier CNRS, UMR 5174 Evolution et Diversité Biologique</institution>,<addr-line>bâtiment 4R1, Toulouse 31062</addr-line>,<country>France</country></nlm:aff></affiliation></author><author><name sortKey="Galbraith, David R" sort="Galbraith, David R" uniqKey="Galbraith D" first="David R." last="Galbraith">David R. Galbraith</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Lopez Gonzalez, Gabriela" sort="Lopez Gonzalez, Gabriela" uniqKey="Lopez Gonzalez G" first="Gabriela" last="Lopez Gonzalez">Gabriela Lopez Gonzalez</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Monteagudo Mendoza, Abel" sort="Monteagudo Mendoza, Abel" uniqKey="Monteagudo Mendoza A" first="Abel" last="Monteagudo Mendoza">Abel Monteagudo Mendoza</name><affiliation><nlm:aff id="af5"><addr-line>Jardín Botánico de Missouri</addr-line>,<institution>Prolongacion Bolognesi Mz. E</institution>,<addr-line>Lote 6, Oxapampa, Pasco</addr-line>,<country>Peru</country></nlm:aff></affiliation><affiliation><nlm:aff id="af40"><institution>Universidad Nacional de San Antonio Abad del Cusco</institution>,<addr-line>Av. de la Cultura N° 733, Cusco</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="Pennington, R Toby" sort="Pennington, R Toby" uniqKey="Pennington R" first="R. Toby" last="Pennington">R. Toby Pennington</name><affiliation><nlm:aff id="af3"><addr-line>Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Poorter, Lourens" sort="Poorter, Lourens" uniqKey="Poorter L" first="Lourens" last="Poorter">Lourens Poorter</name><affiliation><nlm:aff id="af6"><addr-line>Forest Ecology and Forest Management Group</addr-line>,<institution>Wageningen University and Research</institution>,<addr-line>PO Box 47, 6700 AA Wageningen</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Alexiades, Miguel" sort="Alexiades, Miguel" uniqKey="Alexiades M" first="Miguel" last="Alexiades">Miguel Alexiades</name><affiliation><nlm:aff id="af7"><addr-line>School of Anthropology and Conservation</addr-line>,<institution>University of Kent</institution>,<addr-line>Marlowe Building, Canterbury, Kent CT2 7NR</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Alvarez Davila, Esteban" sort="Alvarez Davila, Esteban" uniqKey="Alvarez Davila E" first="Esteban" last="Álvarez-Dávila">Esteban Álvarez-Dávila</name><affiliation><nlm:aff id="af8"><institution>Fundación Con Vida</institution>,<addr-line>Medellin Cra 48: 20-114</addr-line>,<country>Colombia</country></nlm:aff></affiliation></author><author><name sortKey="Andrade, Ana" sort="Andrade, Ana" uniqKey="Andrade A" first="Ana" last="Andrade">Ana Andrade</name><affiliation><nlm:aff id="af9"><addr-line>Biological Dynamics of Forest Fragment Project (INPA & STRI), C.P. 478, Manaus, Amazonas 69.011-970</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Aragao, Luis E O C" sort="Aragao, Luis E O C" uniqKey="Aragao L" first="Luis E. O. C." last="Aragão">Luis E. O. C. Aragão</name><affiliation><nlm:aff id="af10"><addr-line>Geography, College of Life and Environmental Sciences</addr-line>,<institution>University of Exeter, Drive, Exeter</institution>,<addr-line>Rennes EX4 4RJ</addr-line>,<country>UK</country></nlm:aff></affiliation><affiliation><nlm:aff id="af11"><institution>National Institute for Space Research (INPE)</institution>,<addr-line>São José dos Campos, São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Araujo Murakami, Alejandro" sort="Araujo Murakami, Alejandro" uniqKey="Araujo Murakami A" first="Alejandro" last="Araujo-Murakami">Alejandro Araujo-Murakami</name><affiliation><nlm:aff id="af12"><addr-line>Museo de Historia Natural Noel Kempff Mercado</addr-line>,<institution>Universidad Autonoma Gabriel Rene Moreno</institution>,<addr-line>Casilla 2489, Av. Irala 565, Santa Cruz</addr-line>,<country>Bolivia</country></nlm:aff></affiliation></author><author><name sortKey="Arets, Eric J M M" sort="Arets, Eric J M M" uniqKey="Arets E" first="Eric J. M. M." last="Arets">Eric J. M. M. Arets</name><affiliation><nlm:aff id="af13"><addr-line>Alterra</addr-line>,<institution>Wageningen University and Research Centre</institution>,<addr-line>PO Box 47, Wageningen 6700 AA</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Aymard C, Gerardo A" sort="Aymard C, Gerardo A" uniqKey="Aymard C G" first="Gerardo A." last="Aymard C">Gerardo A. Aymard C</name><affiliation><nlm:aff id="af14"><addr-line>UNELLEZ-Guanare, Programa del Agro y del Mar</addr-line>,<institution>Herbario Universitario (PORT)</institution>,<addr-line>Mesa de Cavacas, Estado Portuguesa 3350</addr-line>,<country>Venezuela</country></nlm:aff></affiliation></author><author><name sortKey="Baraloto, Christopher" sort="Baraloto, Christopher" uniqKey="Baraloto C" first="Christopher" last="Baraloto">Christopher Baraloto</name><affiliation><nlm:aff id="af15"><addr-line>International Center for Tropical Botany, Department of Biological Sciences</addr-line>,<institution>Florida International University</institution>,<addr-line>Miami, FL 33199</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Barroso, Jorcely G" sort="Barroso, Jorcely G" uniqKey="Barroso J" first="Jorcely G." last="Barroso">Jorcely G. Barroso</name><affiliation><nlm:aff id="af16"><institution>Universidade Federal do Acre, Campus de Cruzeiro do Sul</institution>,<country>Acre, Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Bonal, Damien" sort="Bonal, Damien" uniqKey="Bonal D" first="Damien" last="Bonal">Damien Bonal</name><affiliation><nlm:aff id="af17"><institution>INRA, UMR 1137 ‘Ecologie et Ecophysiologie Forestiere’</institution>,<addr-line>Champenoux 54280</addr-line>,<country>France</country></nlm:aff></affiliation></author><author><name sortKey="Boot, Rene G A" sort="Boot, Rene G A" uniqKey="Boot R" first="Rene G. A." last="Boot">Rene G. A. Boot</name><affiliation><nlm:aff id="af18"><addr-line>Tropenbos International, PO Box 232, Wageningen 6700 AE</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Camargo, Jose L C" sort="Camargo, Jose L C" uniqKey="Camargo J" first="José L. C." last="Camargo">José L. C. Camargo</name><affiliation><nlm:aff id="af9"><addr-line>Biological Dynamics of Forest Fragment Project (INPA & STRI), C.P. 478, Manaus, Amazonas 69.011-970</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Comiskey, James A" sort="Comiskey, James A" uniqKey="Comiskey J" first="James A." last="Comiskey">James A. Comiskey</name><affiliation><nlm:aff id="af19"><addr-line>National Park Service, 120 Chatham Lane, Fredericksburg, VA 22405</addr-line>,<country>USA</country></nlm:aff></affiliation><affiliation><nlm:aff id="af20"><institution>Smithsonian Institution</institution>,<addr-line>1100 Jefferson Dr, SW, Washington, DC 20560</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Valverde, Fernando Cornejo" sort="Valverde, Fernando Cornejo" uniqKey="Valverde F" first="Fernando Cornejo" last="Valverde">Fernando Cornejo Valverde</name><affiliation><nlm:aff id="af21"><addr-line>Proyecto Castaña, Madre de Dios</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="De Camargo, Plinio B" sort="De Camargo, Plinio B" uniqKey="De Camargo P" first="Plínio B." last="De Camargo">Plínio B. De Camargo</name><affiliation><nlm:aff id="af22"><addr-line>Centro de Energia Nuclear na Agricultura</addr-line>,<institution>Universidade de São Paulo</institution>,<addr-line>São Paulo, Sao Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Di Fiore, Anthony" sort="Di Fiore, Anthony" uniqKey="Di Fiore A" first="Anthony" last="Di Fiore">Anthony Di Fiore</name><affiliation><nlm:aff id="af23"><addr-line>Department of Anthropology</addr-line>,<institution>University of Texas at Austin</institution>,<addr-line>SAC Room 5.150, 2201 Speedway Stop C3200, Austin, TX 78712</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Elias, Fernando" sort="Elias, Fernando" uniqKey="Elias F" first="Fernando" last="Elias">Fernando Elias</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Erwin, Terry L" sort="Erwin, Terry L" uniqKey="Erwin T" first="Terry L." last="Erwin">Terry L. Erwin</name><affiliation><nlm:aff id="af25"><addr-line>Department of Entomology</addr-line>,<institution>Smithsonian Institution</institution>,<addr-line>PO Box 37012</addr-line>, MRC 187,<addr-line>Washington, DC 20013-7012</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Feldpausch, Ted R" sort="Feldpausch, Ted R" uniqKey="Feldpausch T" first="Ted R." last="Feldpausch">Ted R. Feldpausch</name><affiliation><nlm:aff id="af10"><addr-line>Geography, College of Life and Environmental Sciences</addr-line>,<institution>University of Exeter, Drive, Exeter</institution>,<addr-line>Rennes EX4 4RJ</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Ferreira, Leandro" sort="Ferreira, Leandro" uniqKey="Ferreira L" first="Leandro" last="Ferreira">Leandro Ferreira</name><affiliation><nlm:aff id="af26"><addr-line>Museu Paraense Emilio Goeldi, C.P. 399, 66.040-170, Belém, Pará</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Fyllas, Nikolaos M" sort="Fyllas, Nikolaos M" uniqKey="Fyllas N" first="Nikolaos M." last="Fyllas">Nikolaos M. Fyllas</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Gloor, Emanuel" sort="Gloor, Emanuel" uniqKey="Gloor E" first="Emanuel" last="Gloor">Emanuel Gloor</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Herault, Bruno" sort="Herault, Bruno" uniqKey="Herault B" first="Bruno" last="Herault">Bruno Herault</name><affiliation><nlm:aff id="af27"><institution>Cirad, UMR EcoFoG (AgroParisTech, CNRS, Inra, U Antilles, U Guyane)</institution>,<addr-line>Campus Agronomique, Kourou 97310</addr-line>,<country>French Guiana</country></nlm:aff></affiliation></author><author><name sortKey="Herrera, Rafael" sort="Herrera, Rafael" uniqKey="Herrera R" first="Rafael" last="Herrera">Rafael Herrera</name><affiliation><nlm:aff id="af28"><institution>Centro de Ecología IVIC</institution>,<addr-line>Caracas</addr-line>,<country>Venezuela</country></nlm:aff></affiliation><affiliation><nlm:aff id="af29"><addr-line>Institut für Geographie und Regionalforschung, University of Vienna, Wien</addr-line>,<country>Austria</country></nlm:aff></affiliation></author><author><name sortKey="Higuchi, Niro" sort="Higuchi, Niro" uniqKey="Higuchi N" first="Niro" last="Higuchi">Niro Higuchi</name><affiliation><nlm:aff id="af30"><institution>INPA</institution>,<addr-line>Av. André Araújo, 2.936 - Petrópolis – 69.067-375, Manaus, Amazonas</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Honorio Coronado, Euridice N" sort="Honorio Coronado, Euridice N" uniqKey="Honorio Coronado E" first="Eurídice N." last="Honorio Coronado">Eurídice N. Honorio Coronado</name><affiliation><nlm:aff id="af31"><institution>Instituto de Investigaciones de la Amazonia Peruana</institution>,<addr-line>Apartado 784, Iquitos</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="Killeen, Timothy J" sort="Killeen, Timothy J" uniqKey="Killeen T" first="Timothy J." last="Killeen">Timothy J. Killeen</name><affiliation><nlm:aff id="af32"><institution>AGTECA - Amazónica</institution>,<addr-line>Santa Cruz</addr-line>,<country>Bolivia</country></nlm:aff></affiliation></author><author><name sortKey="Laurance, William F" sort="Laurance, William F" uniqKey="Laurance W" first="William F." last="Laurance">William F. Laurance</name><affiliation><nlm:aff id="af33"><addr-line>Centre for Tropical Environmental and Sustainability Science (TESS) and College of Science and Engineering</addr-line>,<institution>James Cook University</institution>,<addr-line>Cairns, Queensland 4878</addr-line>,<country>Australia</country></nlm:aff></affiliation></author><author><name sortKey="Laurance, Susan" sort="Laurance, Susan" uniqKey="Laurance S" first="Susan" last="Laurance">Susan Laurance</name><affiliation><nlm:aff id="af33"><addr-line>Centre for Tropical Environmental and Sustainability Science (TESS) and College of Science and Engineering</addr-line>,<institution>James Cook University</institution>,<addr-line>Cairns, Queensland 4878</addr-line>,<country>Australia</country></nlm:aff></affiliation></author><author><name sortKey="Lloyd, Jon" sort="Lloyd, Jon" uniqKey="Lloyd J" first="Jon" last="Lloyd">Jon Lloyd</name><affiliation><nlm:aff id="af34"><addr-line>Department of Life Sciences</addr-line>,<institution>Imperial College London</institution>,<addr-line>Silwood Park Campus, Buckhurst, Road, Ascot, Berkshire SL5 7PY</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Lovejoy, Thomas E" sort="Lovejoy, Thomas E" uniqKey="Lovejoy T" first="Thomas E." last="Lovejoy">Thomas E. Lovejoy</name><affiliation><nlm:aff id="af35"><addr-line>Environmental Science and Policy, and the Department of Public and International Affairs</addr-line>,<institution>George Mason University (GMU)</institution>,<addr-line>Washington, DC</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Malhi, Yadvinder" sort="Malhi, Yadvinder" uniqKey="Malhi Y" first="Yadvinder" last="Malhi">Yadvinder Malhi</name><affiliation><nlm:aff id="af36"><addr-line>Environmental Change Institute, School of Geography and the Environment</addr-line>,<institution>University of Oxford</institution>,<addr-line>Oxford</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Maracahipes, Leandro" sort="Maracahipes, Leandro" uniqKey="Maracahipes L" first="Leandro" last="Maracahipes">Leandro Maracahipes</name><affiliation><nlm:aff id="af37"><addr-line>Programa de Pós-graduação em Ecologia e Evolução</addr-line>,<institution>Universidade Federal de Goias</institution>,<addr-line>Goiânia, Goias</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Marimon, Beatriz S" sort="Marimon, Beatriz S" uniqKey="Marimon B" first="Beatriz S." last="Marimon">Beatriz S. Marimon</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Marimon Junior, Ben H" sort="Marimon Junior, Ben H" uniqKey="Marimon Junior B" first="Ben H." last="Marimon-Junior">Ben H. Marimon-Junior</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Mendoza, Casimiro" sort="Mendoza, Casimiro" uniqKey="Mendoza C" first="Casimiro" last="Mendoza">Casimiro Mendoza</name><affiliation><nlm:aff id="af38"><addr-line>Escuela de Ciencias Forestales, Unidad Académica del Trópico</addr-line>,<institution>Universidad Mayor de San Simón</institution>,<addr-line>Sacta</addr-line>,<country>Bolivia</country></nlm:aff></affiliation></author><author><name sortKey="Morandi, Paulo" sort="Morandi, Paulo" uniqKey="Morandi P" first="Paulo" last="Morandi">Paulo Morandi</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Neill, David A" sort="Neill, David A" uniqKey="Neill D" first="David A." last="Neill">David A. Neill</name><affiliation><nlm:aff id="af39"><addr-line>Universidad Estatal Amazónica, Puyo, Pastaza</addr-line>,<country>Ecuador</country></nlm:aff></affiliation></author><author><name sortKey="Vargas, Percy Nu Ez" sort="Vargas, Percy Nu Ez" uniqKey="Vargas P" first="Percy Nú Ez" last="Vargas">Percy Nú Ez Vargas</name><affiliation><nlm:aff id="af40"><institution>Universidad Nacional de San Antonio Abad del Cusco</institution>,<addr-line>Av. de la Cultura N° 733, Cusco</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="Oliveira, Edmar A" sort="Oliveira, Edmar A" uniqKey="Oliveira E" first="Edmar A." last="Oliveira">Edmar A. Oliveira</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Lenza, Eddie" sort="Lenza, Eddie" uniqKey="Lenza E" first="Eddie" last="Lenza">Eddie Lenza</name><affiliation><nlm:aff id="af24"><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Palacios, Walter A" sort="Palacios, Walter A" uniqKey="Palacios W" first="Walter A." last="Palacios">Walter A. Palacios</name><affiliation><nlm:aff id="af41"><institution>Universidad Técnica del Norte and Herbario Nacional del Ecuador</institution>,<addr-line>Casilla 17-21-1787, Av. Río Coca E6-115, Quito</addr-line>,<country>Ecuador</country></nlm:aff></affiliation></author><author><name sortKey="Pe Uela Mora, Maria C" sort="Pe Uela Mora, Maria C" uniqKey="Pe Uela Mora M" first="Maria C." last="Pe Uela-Mora">Maria C. Pe Uela-Mora</name><affiliation><nlm:aff id="af42"><institution>Universidad Regional Amazónica IKIAM</institution>,<addr-line>Tena</addr-line>,<country>Ecuador</country></nlm:aff></affiliation></author><author><name sortKey="Pipoly, John J" sort="Pipoly, John J" uniqKey="Pipoly J" first="John J." last="Pipoly">John J. Pipoly</name><affiliation><nlm:aff id="af43"><addr-line>Broward County Parks and Recreation Division, 950 NW 38th St., Oakland Park, FL 33309</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Pitman, Nigel C A" sort="Pitman, Nigel C A" uniqKey="Pitman N" first="Nigel C. A." last="Pitman">Nigel C. A. Pitman</name><affiliation><nlm:aff id="af44"><addr-line>Center for Tropical Conservation</addr-line>,<institution>Duke University</institution>,<addr-line>PO Box 90381, Durham, NC 27708</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Prieto, Adriana" sort="Prieto, Adriana" uniqKey="Prieto A" first="Adriana" last="Prieto">Adriana Prieto</name><affiliation><nlm:aff id="af45"><addr-line>Doctorado Instituto de Ciencias Naturales</addr-line>,<institution>Universidad ciol de Colombia</institution>,<country>Colombia</country></nlm:aff></affiliation></author><author><name sortKey="Quesada, Carlos A" sort="Quesada, Carlos A" uniqKey="Quesada C" first="Carlos A." last="Quesada">Carlos A. Quesada</name><affiliation><nlm:aff id="af30"><institution>INPA</institution>,<addr-line>Av. André Araújo, 2.936 - Petrópolis – 69.067-375, Manaus, Amazonas</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Ramirez Angulo, Hirma" sort="Ramirez Angulo, Hirma" uniqKey="Ramirez Angulo H" first="Hirma" last="Ramirez-Angulo">Hirma Ramirez-Angulo</name><affiliation><nlm:aff id="af46"><addr-line>Instituto de Investigaciones para el Desarrollo Forestal (INDEFOR), Facultad de Ciencias Forestales y Ambientales</addr-line>,<institution>Universidad de Los Andes</institution>,<addr-line>Conjunto Forestal, C.P. 5101, Mérida</addr-line>,<country>Venezuela</country></nlm:aff></affiliation></author><author><name sortKey="Rudas, Agustin" sort="Rudas, Agustin" uniqKey="Rudas A" first="Agustin" last="Rudas">Agustin Rudas</name><affiliation><nlm:aff id="af45"><addr-line>Doctorado Instituto de Ciencias Naturales</addr-line>,<institution>Universidad ciol de Colombia</institution>,<country>Colombia</country></nlm:aff></affiliation></author><author><name sortKey="Ruokolainen, Kalle" sort="Ruokolainen, Kalle" uniqKey="Ruokolainen K" first="Kalle" last="Ruokolainen">Kalle Ruokolainen</name><affiliation><nlm:aff id="af47"><addr-line>Department of Geography and Geology</addr-line>,<institution>University of Turku, 20014 Turku</institution>,<country>Finland</country></nlm:aff></affiliation></author><author><name sortKey="Salomao, Rafael P" sort="Salomao, Rafael P" uniqKey="Salomao R" first="Rafael P." last="Salomão">Rafael P. Salomão</name><affiliation><nlm:aff id="af26"><addr-line>Museu Paraense Emilio Goeldi, C.P. 399, 66.040-170, Belém, Pará</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Silveira, Marcos" sort="Silveira, Marcos" uniqKey="Silveira M" first="Marcos" last="Silveira">Marcos Silveira</name><affiliation><nlm:aff id="af48"><institution>Museu Universitário, Universidade Federal do Acre</institution>,<addr-line>Rio Branco, AC 69910-900</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Stropp, Juliana" sort="Stropp, Juliana" uniqKey="Stropp J" first="Juliana" last="Stropp">Juliana Stropp</name><affiliation><nlm:aff id="af49"><addr-line>Institute of Biological and Health Sciences (ICBS)</addr-line>,<institution>Federal University of Alagoas</institution>,<addr-line>Maceió, AL</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Ter Steege, Hans" sort="Ter Steege, Hans" uniqKey="Ter Steege H" first="Hans" last="Ter Steege">Hans Ter Steege</name><affiliation><nlm:aff id="af50"><institution>Naturalis Biodiversity Center</institution>,<addr-line>Vondellaan 55, Postbus 9517, Leiden 2300 RA</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Thomas Caesar, Raquel" sort="Thomas Caesar, Raquel" uniqKey="Thomas Caesar R" first="Raquel" last="Thomas-Caesar">Raquel Thomas-Caesar</name><affiliation><nlm:aff id="af51"><institution>Iwokrama Intertiol Centre for Rainforest Conservation and Development</institution>,<addr-line>77 High Street Kingston, Georgetown</addr-line>,<country>Guyana</country></nlm:aff></affiliation></author><author><name sortKey="Van Der Hout, Peter" sort="Van Der Hout, Peter" uniqKey="Van Der Hout P" first="Peter" last="Van Der Hout">Peter Van Der Hout</name><affiliation><nlm:aff id="af52"><addr-line>Van der Hout Forestry Consulting, Jan Trooststraat 6, Rotterdam 3078 HP</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Van Der Heijden, Geertje M F" sort="Van Der Heijden, Geertje M F" uniqKey="Van Der Heijden G" first="Geertje M. F." last="Van Der Heijden">Geertje M. F. Van Der Heijden</name><affiliation><nlm:aff id="af53"><addr-line>School of Geography</addr-line>,<institution>University of Nottingham</institution>,<addr-line>University Park, Nottingham NG7 2RD</addr-line>,<country>UK</country></nlm:aff></affiliation></author><author><name sortKey="Van Der Meer, Peter J" sort="Van Der Meer, Peter J" uniqKey="Van Der Meer P" first="Peter J." last="Van Der Meer">Peter J. Van Der Meer</name><affiliation><nlm:aff id="af54"><institution>Van Hall Larenstein University of Applied Sciences</institution>,<addr-line>PO Box 9001, 6880 GB Velp</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Vasquez, Rodolfo V" sort="Vasquez, Rodolfo V" uniqKey="Vasquez R" first="Rodolfo V." last="Vasquez">Rodolfo V. Vasquez</name><affiliation><nlm:aff id="af5"><addr-line>Jardín Botánico de Missouri</addr-line>,<institution>Prolongacion Bolognesi Mz. E</institution>,<addr-line>Lote 6, Oxapampa, Pasco</addr-line>,<country>Peru</country></nlm:aff></affiliation></author><author><name sortKey="Vieira, Simone A" sort="Vieira, Simone A" uniqKey="Vieira S" first="Simone A." last="Vieira">Simone A. Vieira</name><affiliation><nlm:aff id="af55"><addr-line>Universidade Estadual de Campinas</addr-line>,<institution>Núcleo de Estudos e Pesquisas Ambientais – NEPAM</institution>,<addr-line>Campinas, São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Vilanova, Emilio" sort="Vilanova, Emilio" uniqKey="Vilanova E" first="Emilio" last="Vilanova">Emilio Vilanova</name><affiliation><nlm:aff id="af56"><addr-line>Facultad de Ciencias Forestales y Ambientales</addr-line>,<institution>Universidad de Los Andes</institution>,<addr-line>Mérida</addr-line>,<country>Venezuela</country></nlm:aff></affiliation></author><author><name sortKey="Vos, Vincent A" sort="Vos, Vincent A" uniqKey="Vos V" first="Vincent A." last="Vos">Vincent A. Vos</name><affiliation><nlm:aff id="af57"><institution>Centro de Investigación y Promoción del Campesinado - regional Norte Amazónico</institution>,<addr-line>C/ Nicanor Gonzalo Salvatierra N° 362, Casilla 16, Riberalta</addr-line>,<country>Bolivia</country></nlm:aff></affiliation><affiliation><nlm:aff id="af58"><institution>Universidad Autónoma del Beni</institution>,<addr-line>Campus Universitario, Riberalta</addr-line>,<country>Bolivia</country></nlm:aff></affiliation></author><author><name sortKey="Wang, Ophelia" sort="Wang, Ophelia" uniqKey="Wang O" first="Ophelia" last="Wang">Ophelia Wang</name><affiliation><nlm:aff id="af59"><institution>Northern Arizona University</institution>,<addr-line>Flagstaff, AZ 86011</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Young, Kenneth R" sort="Young, Kenneth R" uniqKey="Young K" first="Kenneth R." last="Young">Kenneth R. Young</name><affiliation><nlm:aff id="af60"><addr-line>Department of Geography and the Environment</addr-line>,<institution>University of Texas at Austin</institution>,<addr-line>Austin, TX 78712</addr-line>,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Zagt, Roderick J" sort="Zagt, Roderick J" uniqKey="Zagt R" first="Roderick J." last="Zagt">Roderick J. Zagt</name><affiliation><nlm:aff id="af18"><addr-line>Tropenbos International, PO Box 232, Wageningen 6700 AE</addr-line>,<country>The Netherlands</country></nlm:aff></affiliation></author><author><name sortKey="Baker, Timothy R" sort="Baker, Timothy R" uniqKey="Baker T" first="Timothy R." last="Baker">Timothy R. Baker</name><affiliation><nlm:aff id="af1"><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></nlm:aff></affiliation></author></analytic><series><title level="j">Proceedings of the Royal Society B: Biological Sciences</title><idno type="ISSN">0962-8452</idno><idno type="eISSN">1471-2954</idno><imprint><date when="2016">2016</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Blomberg, Sp" uniqKey="Blomberg S">SP Blomberg</name></author><author><name sortKey="Garland, T" uniqKey="Garland T">T Garland</name></author><author><name sortKey="Ives, Ar" uniqKey="Ives A">AR Ives</name></author><author><name sortKey="Crespi, B" uniqKey="Crespi B">B Crespi</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Losos, Jb" uniqKey="Losos J">JB Losos</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Crisp, Md" uniqKey="Crisp M">MD Crisp</name></author><author><name sortKey="Cook, Lg" uniqKey="Cook L">LG Cook</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Revell, Lj" uniqKey="Revell L">LJ Revell</name></author><author><name sortKey="Harmon, Lj" uniqKey="Harmon L">LJ Harmon</name></author><author><name sortKey="Collar, Dc" uniqKey="Collar D">DC Collar</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Webb, Co" uniqKey="Webb C">CO Webb</name></author><author><name sortKey="Ackerly, Dd" uniqKey="Ackerly D">DD Ackerly</name></author><author><name sortKey="Mcpeek, Ma" uniqKey="Mcpeek M">MA McPeek</name></author><author><name sortKey="Donoghue, Mj" uniqKey="Donoghue M">MJ Donoghue</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Baraloto, C" uniqKey="Baraloto C">C Baraloto</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Yang, J" uniqKey="Yang J">J Yang</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Cadotte, Mw" uniqKey="Cadotte M">MW Cadotte</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Cadotte, Mw" uniqKey="Cadotte M">MW Cadotte</name></author><author><name sortKey="Cardinale, Bj" uniqKey="Cardinale B">BJ Cardinale</name></author><author><name sortKey="Oakley, Th" uniqKey="Oakley T">TH Oakley</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Uriarte, M" uniqKey="Uriarte M">M Uriarte</name></author><author><name sortKey="Swenson, Ng" uniqKey="Swenson N">NG Swenson</name></author><author><name sortKey="Chazdon, Rl" uniqKey="Chazdon R">RL Chazdon</name></author><author><name sortKey="Comita, Ls" uniqKey="Comita L">LS Comita</name></author><author><name sortKey="John Kress, W" uniqKey="John Kress W">W John Kress</name></author><author><name sortKey="Erickson, D" uniqKey="Erickson D">D Erickson</name></author><author><name sortKey="Forero Montana, J" uniqKey="Forero Montana J">J Forero-Montana</name></author><author><name sortKey="Zimmerman, Jk" uniqKey="Zimmerman J">JK Zimmerman</name></author><author><name sortKey="Thompson, J" uniqKey="Thompson J">J Thompson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Fan, Z X" uniqKey="Fan Z">Z-X Fan</name></author><author><name sortKey="Zhang, S B" uniqKey="Zhang S">S-B Zhang</name></author><author><name sortKey="Hao, G Y" uniqKey="Hao G">G-Y Hao</name></author><author><name sortKey="Ferry Slik, Jw" uniqKey="Ferry Slik J">JW Ferry Slik</name></author><author><name sortKey="Cao, K F" uniqKey="Cao K">K-F Cao</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Swenson, Ng" uniqKey="Swenson N">NG Swenson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Dexter, K" uniqKey="Dexter K">K Dexter</name></author><author><name sortKey="Chave, J" uniqKey="Chave J">J 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F.</given-names></name><xref ref-type="aff" rid="af53">53</xref></contrib><contrib contrib-type="author"><name><surname>van der Meer</surname><given-names>Peter J.</given-names></name><xref ref-type="aff" rid="af54">54</xref></contrib><contrib contrib-type="author"><name><surname>Vasquez</surname><given-names>Rodolfo V.</given-names></name><xref ref-type="aff" rid="af5">5</xref></contrib><contrib contrib-type="author"><name><surname>Vieira</surname><given-names>Simone A.</given-names></name><xref ref-type="aff" rid="af55">55</xref></contrib><contrib contrib-type="author"><name><surname>Vilanova</surname><given-names>Emilio</given-names></name><xref ref-type="aff" rid="af56">56</xref></contrib><contrib contrib-type="author"><name><surname>Vos</surname><given-names>Vincent A.</given-names></name><xref ref-type="aff" rid="af57">57</xref><xref ref-type="aff" rid="af58">58</xref></contrib><contrib contrib-type="author"><name><surname>Wang</surname><given-names>Ophelia</given-names></name><xref ref-type="aff" rid="af59">59</xref></contrib><contrib contrib-type="author"><name><surname>Young</surname><given-names>Kenneth R.</given-names></name><xref ref-type="aff" rid="af60">60</xref></contrib><contrib contrib-type="author"><name><surname>Zagt</surname><given-names>Roderick J.</given-names></name><xref ref-type="aff" rid="af18">18</xref></contrib><contrib contrib-type="author"><name><surname>Baker</surname><given-names>Timothy R.</given-names></name><xref ref-type="aff" rid="af1">1</xref></contrib></contrib-group><aff id="af1"><label>1</label><addr-line>School of Geography</addr-line>,<institution>University of Leeds</institution>,<addr-line>Leeds LS2 9JT</addr-line>,<country>UK</country></aff><aff id="af2"><label>2</label><addr-line>School of Geosciences</addr-line>,<institution>University of Edinburgh</institution>,<addr-line>201 Crew Building, King's Buildings, Edinburgh EH9 3FF</addr-line>,<country>UK</country></aff><aff id="af3"><label>3</label><addr-line>Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR</addr-line>,<country>UK</country></aff><aff id="af4"><label>4</label><institution>Université Paul Sabatier CNRS, UMR 5174 Evolution et Diversité Biologique</institution>,<addr-line>bâtiment 4R1, Toulouse 31062</addr-line>,<country>France</country></aff><aff id="af5"><label>5</label><addr-line>Jardín Botánico de Missouri</addr-line>,<institution>Prolongacion Bolognesi Mz. E</institution>,<addr-line>Lote 6, Oxapampa, Pasco</addr-line>,<country>Peru</country></aff><aff id="af6"><label>6</label><addr-line>Forest Ecology and Forest Management Group</addr-line>,<institution>Wageningen University and Research</institution>,<addr-line>PO Box 47, 6700 AA Wageningen</addr-line>,<country>The Netherlands</country></aff><aff id="af7"><label>7</label><addr-line>School of Anthropology and Conservation</addr-line>,<institution>University of Kent</institution>,<addr-line>Marlowe Building, Canterbury, Kent CT2 7NR</addr-line>,<country>UK</country></aff><aff id="af8"><label>8</label><institution>Fundación Con Vida</institution>,<addr-line>Medellin Cra 48: 20-114</addr-line>,<country>Colombia</country></aff><aff id="af9"><label>9</label><addr-line>Biological Dynamics of Forest Fragment Project (INPA & STRI), C.P. 478, Manaus, Amazonas 69.011-970</addr-line>,<country>Brazil</country></aff><aff id="af10"><label>10</label><addr-line>Geography, College of Life and Environmental Sciences</addr-line>,<institution>University of Exeter, Drive, Exeter</institution>,<addr-line>Rennes EX4 4RJ</addr-line>,<country>UK</country></aff><aff id="af11"><label>11</label><institution>National Institute for Space Research (INPE)</institution>,<addr-line>São José dos Campos, São Paulo</addr-line>,<country>Brazil</country></aff><aff id="af12"><label>12</label><addr-line>Museo de Historia Natural Noel Kempff Mercado</addr-line>,<institution>Universidad Autonoma Gabriel Rene Moreno</institution>,<addr-line>Casilla 2489, Av. Irala 565, Santa Cruz</addr-line>,<country>Bolivia</country></aff><aff id="af13"><label>13</label><addr-line>Alterra</addr-line>,<institution>Wageningen University and Research Centre</institution>,<addr-line>PO Box 47, Wageningen 6700 AA</addr-line>,<country>The Netherlands</country></aff><aff id="af14"><label>14</label><addr-line>UNELLEZ-Guanare, Programa del Agro y del Mar</addr-line>,<institution>Herbario Universitario (PORT)</institution>,<addr-line>Mesa de Cavacas, Estado Portuguesa 3350</addr-line>,<country>Venezuela</country></aff><aff id="af15"><label>15</label><addr-line>International Center for Tropical Botany, Department of Biological Sciences</addr-line>,<institution>Florida International University</institution>,<addr-line>Miami, FL 33199</addr-line>,<country>USA</country></aff><aff id="af16"><label>16</label><institution>Universidade Federal do Acre, Campus de Cruzeiro do Sul</institution>,<country>Acre, Brazil</country></aff><aff id="af17"><label>17</label><institution>INRA, UMR 1137 ‘Ecologie et Ecophysiologie Forestiere’</institution>,<addr-line>Champenoux 54280</addr-line>,<country>France</country></aff><aff id="af18"><label>18</label><addr-line>Tropenbos International, PO Box 232, Wageningen 6700 AE</addr-line>,<country>The Netherlands</country></aff><aff id="af19"><label>19</label><addr-line>National Park Service, 120 Chatham Lane, Fredericksburg, VA 22405</addr-line>,<country>USA</country></aff><aff id="af20"><label>20</label><institution>Smithsonian Institution</institution>,<addr-line>1100 Jefferson Dr, SW, Washington, DC 20560</addr-line>,<country>USA</country></aff><aff id="af21"><label>21</label><addr-line>Proyecto Castaña, Madre de Dios</addr-line>,<country>Peru</country></aff><aff id="af22"><label>22</label><addr-line>Centro de Energia Nuclear na Agricultura</addr-line>,<institution>Universidade de São Paulo</institution>,<addr-line>São Paulo, Sao Paulo</addr-line>,<country>Brazil</country></aff><aff id="af23"><label>23</label><addr-line>Department of Anthropology</addr-line>,<institution>University of Texas at Austin</institution>,<addr-line>SAC Room 5.150, 2201 Speedway Stop C3200, Austin, TX 78712</addr-line>,<country>USA</country></aff><aff id="af24"><label>24</label><institution>Universidade do Estado de Mato Grosso</institution>,<addr-line>Campus de Nova Xavantina, Caixa Postal 08, 78.690-000, Nova Xavantina, Mato Grosso</addr-line>,<country>Brazil</country></aff><aff id="af25"><label>25</label><addr-line>Department of Entomology</addr-line>,<institution>Smithsonian Institution</institution>,<addr-line>PO Box 37012</addr-line>, MRC 187,<addr-line>Washington, DC 20013-7012</addr-line>,<country>USA</country></aff><aff id="af26"><label>26</label><addr-line>Museu Paraense Emilio Goeldi, C.P. 399, 66.040-170, Belém, Pará</addr-line>,<country>Brazil</country></aff><aff id="af27"><label>27</label><institution>Cirad, UMR EcoFoG (AgroParisTech, CNRS, Inra, U Antilles, U Guyane)</institution>,<addr-line>Campus Agronomique, Kourou 97310</addr-line>,<country>French Guiana</country></aff><aff id="af28"><label>28</label><institution>Centro de Ecología IVIC</institution>,<addr-line>Caracas</addr-line>,<country>Venezuela</country></aff><aff id="af29"><label>29</label><addr-line>Institut für Geographie und Regionalforschung, University of Vienna, Wien</addr-line>,<country>Austria</country></aff><aff id="af30"><label>30</label><institution>INPA</institution>,<addr-line>Av. André Araújo, 2.936 - Petrópolis – 69.067-375, Manaus, Amazonas</addr-line>,<country>Brazil</country></aff><aff id="af31"><label>31</label><institution>Instituto de Investigaciones de la Amazonia Peruana</institution>,<addr-line>Apartado 784, Iquitos</addr-line>,<country>Peru</country></aff><aff id="af32"><label>32</label><institution>AGTECA - Amazónica</institution>,<addr-line>Santa Cruz</addr-line>,<country>Bolivia</country></aff><aff id="af33"><label>33</label><addr-line>Centre for Tropical Environmental and Sustainability Science (TESS) and College of Science and Engineering</addr-line>,<institution>James Cook University</institution>,<addr-line>Cairns, Queensland 4878</addr-line>,<country>Australia</country></aff><aff id="af34"><label>34</label><addr-line>Department of Life Sciences</addr-line>,<institution>Imperial College London</institution>,<addr-line>Silwood Park Campus, Buckhurst, Road, Ascot, Berkshire SL5 7PY</addr-line>,<country>UK</country></aff><aff id="af35"><label>35</label><addr-line>Environmental Science and Policy, and the Department of Public and International Affairs</addr-line>,<institution>George Mason University (GMU)</institution>,<addr-line>Washington, DC</addr-line>,<country>USA</country></aff><aff id="af36"><label>36</label><addr-line>Environmental Change Institute, School of Geography and the Environment</addr-line>,<institution>University of Oxford</institution>,<addr-line>Oxford</addr-line>,<country>UK</country></aff><aff id="af37"><label>37</label><addr-line>Programa de Pós-graduação em Ecologia e Evolução</addr-line>,<institution>Universidade Federal de Goias</institution>,<addr-line>Goiânia, Goias</addr-line>,<country>Brazil</country></aff><aff id="af38"><label>38</label><addr-line>Escuela de Ciencias Forestales, Unidad Académica del Trópico</addr-line>,<institution>Universidad Mayor de San Simón</institution>,<addr-line>Sacta</addr-line>,<country>Bolivia</country></aff><aff id="af39"><label>39</label><addr-line>Universidad Estatal Amazónica, Puyo, Pastaza</addr-line>,<country>Ecuador</country></aff><aff id="af40"><label>40</label><institution>Universidad Nacional de San Antonio Abad del Cusco</institution>,<addr-line>Av. de la Cultura N° 733, Cusco</addr-line>,<country>Peru</country></aff><aff id="af41"><label>41</label><institution>Universidad Técnica del Norte and Herbario Nacional del Ecuador</institution>,<addr-line>Casilla 17-21-1787, Av. Río Coca E6-115, Quito</addr-line>,<country>Ecuador</country></aff><aff id="af42"><label>42</label><institution>Universidad Regional Amazónica IKIAM</institution>,<addr-line>Tena</addr-line>,<country>Ecuador</country></aff><aff id="af43"><label>43</label><addr-line>Broward County Parks and Recreation Division, 950 NW 38th St., Oakland Park, FL 33309</addr-line>,<country>USA</country></aff><aff id="af44"><label>44</label><addr-line>Center for Tropical Conservation</addr-line>,<institution>Duke University</institution>,<addr-line>PO Box 90381, Durham, NC 27708</addr-line>,<country>USA</country></aff><aff id="af45"><label>45</label><addr-line>Doctorado Instituto de Ciencias Naturales</addr-line>,<institution>Universidad ciol de Colombia</institution>,<country>Colombia</country></aff><aff id="af46"><label>46</label><addr-line>Instituto de Investigaciones para el Desarrollo Forestal (INDEFOR), Facultad de Ciencias Forestales y Ambientales</addr-line>,<institution>Universidad de Los Andes</institution>,<addr-line>Conjunto Forestal, C.P. 5101, Mérida</addr-line>,<country>Venezuela</country></aff><aff id="af47"><label>47</label><addr-line>Department of Geography and Geology</addr-line>,<institution>University of Turku, 20014 Turku</institution>,<country>Finland</country></aff><aff id="af48"><label>48</label><institution>Museu Universitário, Universidade Federal do Acre</institution>,<addr-line>Rio Branco, AC 69910-900</addr-line>,<country>Brazil</country></aff><aff id="af49"><label>49</label><addr-line>Institute of Biological and Health Sciences (ICBS)</addr-line>,<institution>Federal University of Alagoas</institution>,<addr-line>Maceió, AL</addr-line>,<country>Brazil</country></aff><aff id="af50"><label>50</label><institution>Naturalis Biodiversity Center</institution>,<addr-line>Vondellaan 55, Postbus 9517, Leiden 2300 RA</addr-line>,<country>The Netherlands</country></aff><aff id="af51"><label>51</label><institution>Iwokrama Intertiol Centre for Rainforest Conservation and Development</institution>,<addr-line>77 High Street Kingston, Georgetown</addr-line>,<country>Guyana</country></aff><aff id="af52"><label>52</label><addr-line>Van der Hout Forestry Consulting, Jan Trooststraat 6, Rotterdam 3078 HP</addr-line>,<country>The Netherlands</country></aff><aff id="af53"><label>53</label><addr-line>School of Geography</addr-line>,<institution>University of Nottingham</institution>,<addr-line>University Park, Nottingham NG7 2RD</addr-line>,<country>UK</country></aff><aff id="af54"><label>54</label><institution>Van Hall Larenstein University of Applied Sciences</institution>,<addr-line>PO Box 9001, 6880 GB Velp</addr-line>,<country>The Netherlands</country></aff><aff id="af55"><label>55</label><addr-line>Universidade Estadual de Campinas</addr-line>,<institution>Núcleo de Estudos e Pesquisas Ambientais – NEPAM</institution>,<addr-line>Campinas, São Paulo</addr-line>,<country>Brazil</country></aff><aff id="af56"><label>56</label><addr-line>Facultad de Ciencias Forestales y Ambientales</addr-line>,<institution>Universidad de Los Andes</institution>,<addr-line>Mérida</addr-line>,<country>Venezuela</country></aff><aff id="af57"><label>57</label><institution>Centro de Investigación y Promoción del Campesinado - regional Norte Amazónico</institution>,<addr-line>C/ Nicanor Gonzalo Salvatierra N° 362, Casilla 16, Riberalta</addr-line>,<country>Bolivia</country></aff><aff id="af58"><label>58</label><institution>Universidad Autónoma del Beni</institution>,<addr-line>Campus Universitario, Riberalta</addr-line>,<country>Bolivia</country></aff><aff id="af59"><label>59</label><institution>Northern Arizona University</institution>,<addr-line>Flagstaff, AZ 86011</addr-line>,<country>USA</country></aff><aff id="af60"><label>60</label><addr-line>Department of Geography and the Environment</addr-line>,<institution>University of Texas at Austin</institution>,<addr-line>Austin, TX 78712</addr-line>,<country>USA</country></aff><author-notes><corresp id="cor1">e-mail: <email>fecoelhos@gmail.com</email></corresp><fn fn-type="other"><p>Electronic supplementary material is available online at <uri xlink:href="https://dx.doi.org/10.6084/m9.figshare.c.3582146">https://dx.doi.org/10.6084/m9.figshare.c.3582146</uri>.</p></fn></author-notes><pub-date pub-type="ppub"><day>14</day><month>12</month><year>2016</year></pub-date><pub-date pub-type="pmc-release"><day>14</day><month>12</month><year>2016</year></pub-date><pmc-comment> PMC Release delay is 0 months and 0 days and was based on the
<volume>283</volume><issue>1844</issue><elocation-id>20161587</elocation-id><history><date date-type="received"><day>21</day><month>7</month><year>2016</year></date><date date-type="accepted"><day>3</day><month>11</month><year>2016</year></date></history><permissions><copyright-statement>© 2016 The Authors.</copyright-statement><copyright-year>2016</copyright-year><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/4.0/"><license-p>Published by the Royal Society under the terms of the Creative Commons Attribution License <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">http://creativecommons.org/licenses/by/4.0/</ext-link>, which permits unrestricted use, provided the original author and source are credited.</license-p></license></permissions><self-uri content-type="pdf" xlink:href="rspb20161587.pdf"></self-uri><abstract><p>Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change.</p></abstract><kwd-group><kwd>tropical tree</kwd><kwd>trait</kwd><kwd>convergent evolution</kwd><kwd>divergent selection</kwd><kwd>phylogenetic signal</kwd></kwd-group><funding-group specific-use="FundRef"><award-group><funding-source><institution-wrap><institution>Coordenação de Aperfeiçoamento de Pessoal de Nível Superior</institution><institution-id>http://dx.doi.org/10.13039/501100002322</institution-id></institution-wrap></funding-source><award-id>117913-6</award-id></award-group></funding-group><custom-meta-group><custom-meta><meta-name>cover-date</meta-name><meta-value>December 14, 2016</meta-value></custom-meta></custom-meta-group></article-meta></front><body><sec id="s1"><label>1.</label><title>Introduction</title><p>Evolutionary heritage may act as a major constraint on the ecological roles that species in a lineage can occupy. Even under a random model of trait evolution where functional traits drift in state over time (e.g. a Brownian motion model), we would expect closely related species to have similar functional trait values and similar ecologies due to their shared common ancestry [<xref rid="RSPB20161587C1" ref-type="bibr">1</xref>,<xref rid="RSPB20161587C2" ref-type="bibr">2</xref>]. However, both divergent selection and convergent evolution lead to weaker relationships between species relatedness and their ecological similarity [<xref rid="RSPB20161587C1" ref-type="bibr">1</xref>,<xref rid="RSPB20161587C3" ref-type="bibr">3</xref>,<xref rid="RSPB20161587C4" ref-type="bibr">4</xref>]. Hence, although it is often assumed that close relatives are more similar because they retain the ecological characteristics of their ancestors, in many clades the ancestral character state may not be conserved. Thus, rather than being simply assumed, the tendency of closely related species to have similar ecological characteristics needs to be tested.</p><p>The strength of the link between trait variation and phylogenetic relatedness has a wide range of implications for understanding ecological and evolutionary processes and can be measured by the magnitude of phylogenetic signal (PS) [<xref rid="RSPB20161587C1" ref-type="bibr">1</xref>,<xref rid="RSPB20161587C2" ref-type="bibr">2</xref>]. For example, if a selected trait has significant PS, the relatedness of species can help us to understand the underlying mechanisms that drive community structure [<xref rid="RSPB20161587C5" ref-type="bibr">5</xref>–<xref rid="RSPB20161587C7" ref-type="bibr">7</xref>]. The presence of significant PS also suggests that the sum of phylogenetic distances among species that occur within a community (i.e. phylogenetic diversity) is a useful proxy for functional diversity and that, in turn, phylogenies of tree taxa may contribute to understanding ecosystem function [<xref rid="RSPB20161587C8" ref-type="bibr">8</xref>,<xref rid="RSPB20161587C9" ref-type="bibr">9</xref>]. In addition, if trait values are more similar than expected by chance among closely related lineages, we can predict the trait values for species where trait data are not available.</p><p>Understanding the relative importance of evolutionary heritage versus selection in determining trait variation requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate these processes. Previous studies of the degree of PS among traits of tropical trees, such as seed mass, leaf structure and chemistry, trunk characteristics and range size, have shown variable results [<xref rid="RSPB20161587C6" ref-type="bibr">6</xref>,<xref rid="RSPB20161587C7" ref-type="bibr">7</xref>,<xref rid="RSPB20161587C10" ref-type="bibr">10</xref>–<xref rid="RSPB20161587C13" ref-type="bibr">13</xref>]. For example, some studies show significant PS [<xref rid="RSPB20161587C6" ref-type="bibr">6</xref>,<xref rid="RSPB20161587C7" ref-type="bibr">7</xref>,<xref rid="RSPB20161587C13" ref-type="bibr">13</xref>], while for the same traits other studies have failed to detect any PS, with closely related species exhibiting rather different trait values [<xref rid="RSPB20161587C10" ref-type="bibr">10</xref>,<xref rid="RSPB20161587C11" ref-type="bibr">11</xref>]. A key limitation of many of these studies is the limited spatial and phylogenetic scale of study, as well as the resolution of the phylogeny that they have used [<xref rid="RSPB20161587C14" ref-type="bibr">14</xref>]. Here, we explore patterns of PS at large spatial and phylogenetic scales using a sequence-based phylogeny to test whether there are significant levels of PS for four key traits related to the major axes of life-history variation among tropical trees: tree growth and mortality rates, wood density and potential tree size. These traits are related to resource acquisition and allocation, defence and dispersal ability [<xref rid="RSPB20161587C15" ref-type="bibr">15</xref>,<xref rid="RSPB20161587C16" ref-type="bibr">16</xref>] and represent important axes of functional variation which drive variation in plant performance and function in many ecosystems [<xref rid="RSPB20161587C17" ref-type="bibr">17</xref>]. Moreover, those traits are strongly related to differences in carbon fluxes and storage among species [<xref rid="RSPB20161587C18" ref-type="bibr">18</xref>]. As a result, understanding PS in these traits may help to understand and model ecosystem processes in such highly diverse tropical forests such as Amazonia, which may harbour more than 16 000 tree species [<xref rid="RSPB20161587C19" ref-type="bibr">19</xref>].</p><p>Studying PS at large spatial scales is important because the scale of study affects the strength of PS. At small scales, patterns of PS can be obscured because co-occurring species represent just a small fraction of the species richness of clades [<xref rid="RSPB20161587C20" ref-type="bibr">20</xref>,<xref rid="RSPB20161587C21" ref-type="bibr">21</xref>]. Small spatial scales encompass limited environmental variation, so the species pool is limited to representatives of different lineages that may have similar ecological traits and environmental requirements: this pattern results in a smaller range in traits and low PS. The strength of this effect depends on how environmental variability changes with spatial scale, on the degree of habitat specialization by species and the proportion of clades that are sampled in small-scale studies [<xref rid="RSPB20161587C6" ref-type="bibr">6</xref>,<xref rid="RSPB20161587C7" ref-type="bibr">7</xref>]. However, in general, larger spatial scales incorporate greater environmental heterogeneity and encompass a larger number of lineages with a wider range of trait values. Inferring patterns of PS that are more representative of evolutionary trends therefore typically requires measurement across large spatial scales, including a wide range of environmental conditions and taxa from a broad array of clades [<xref rid="RSPB20161587C22" ref-type="bibr">22</xref>].</p><p>The patterns of PS also depend on traits under investigation and their specific evolutionary history. Some traits may exhibit phylogenetic conservatism where traits in specific lineages are constrained to certain trait values. For example, complex traits, such as growth and mortality, may depend in complex ways on multiple, interacting gene loci [<xref rid="RSPB20161587C23" ref-type="bibr">23</xref>,<xref rid="RSPB20161587C24" ref-type="bibr">24</xref>] which impose strong constraints on trait variation. Alternatively, traits may show no PS because they are under strong selective pressure and/or because they show phenotypic plasticity in response to environmental conditions [<xref rid="RSPB20161587C20" ref-type="bibr">20</xref>,<xref rid="RSPB20161587C25" ref-type="bibr">25</xref>].</p><p>Here, we use a large dataset of several hundred permanent forest plots that occur across a wide range of the environmental conditions from all nine Amazonian countries [<xref rid="RSPB20161587C26" ref-type="bibr">26</xref>], to quantify key demographic traits of more than 300 lineages of tropical trees and explore the PS of these traits using recently published molecular genus- [<xref rid="RSPB20161587C13" ref-type="bibr">13</xref>] and species-level phylogenies ([<xref rid="RSPB20161587C27" ref-type="bibr">27</xref>], KG Dexter & RT Pennington 2013, unpublished data). By exploring how traits are correlated and the strength of PS, our goal is to address the fundamental question of whether repeated convergent and divergent evolution of life-history strategies has erased PS for life-history-related traits in tropical trees, or whether phylogenetic information can be used to understand ecosystem function in the world's most diverse and ecologically important forest.</p></sec><sec sec-type="methods" id="s2"><label>2.</label><title>Material and methods</title><sec id="s2a"><label>(a)</label><title>Plot data</title><p>This study used inventory data from all trees and palms greater than or equal to 10 cm diameter (DBH) in 577 forest plots from the RAINFOR forest plot network (<xref ref-type="fig" rid="RSPB20161587F1">figure 1</xref>; electronic supplementary material, S1) across lowland closed-canopy South American tropical forests. This network is centred on Amazonia and includes plots in forests on the Guiana Shield, in the Choco and northern South America; however, hereafter for simplicity we refer to this sampling region as ‘Amazonia’. Plots are located in old growth, unlogged forests and range in size from 0.04 to 25 ha (most being 1 ha). They span a precipitation gradient from 1300 to 7436 mm yr<sup>−1</sup> [<xref rid="RSPB20161587C28" ref-type="bibr">28</xref>], a broad range of soil types [<xref rid="RSPB20161587C29" ref-type="bibr">29</xref>], and are found below 500 m in elevation. Data were extracted from the ForestPlots.net database which curates tree-by-tree records from RAINFOR and other plot networks [<xref rid="RSPB20161587C26" ref-type="bibr">26</xref>,<xref rid="RSPB20161587C30" ref-type="bibr">30</xref>].
<fig id="RSPB20161587F1" orientation="portrait" position="float"><label>Figure 1.</label><caption><p>Map of location of 577 selected plots in lowland tropical South America over a backcloth of the precipitation gradient (annual precipitation, from the WorldClim dataset [<xref rid="RSPB20161587C28" ref-type="bibr">28</xref>]). The map shows plots, with annual precipitation greater than 1300 mm yr<sup>−1</sup> and altitude less than 500 m. Yellow circles: single census, plots used exclusively for estimating wood density and potential tree size; red circles: multi censuses, plots used for estimating wood density, potential tree size, growth and mortality rates.</p></caption><graphic xlink:href="rspb20161587-g1"></graphic></fig></p><p>For productivity and mortality analyses, we used a subset of 257 repeated census plots with a minimum monitoring period of 2 years from 1962 to 2014. Mean census interval length is 4.4 years and plot mean total monitoring period is 9.9 years. During each census, all surviving trees and palms were measured, dead trees were documented and new trees with greater than or equal to 10 cm DBH were recorded. More detailed measurement methods and plot characteristics have been previously published (e.g. [<xref rid="RSPB20161587C31" ref-type="bibr">31</xref>,<xref rid="RSPB20161587C32" ref-type="bibr">32</xref>]). All recorded species and genus names were checked and standardized using the Taxonomic Name Resolution Service [<xref rid="RSPB20161587C33" ref-type="bibr">33</xref>]. We excluded all trees and palms not identified to genus level (7.9% of stems).</p></sec><sec id="s2b"><label>(b)</label><title>Trait data</title><p>Trait mean values of potential tree size, mean and maximum growth rates, mortality rates and wood density were calculated at both the genus and species levels. Our main analyses were performed at the genus level and covered genera present in a recently published genus-level phylogeny for Amazonian trees [<xref rid="RSPB20161587C13" ref-type="bibr">13</xref>]. Species-level trait data for those clades where we had species-level phylogenies with sufficient sampling of species in our dataset (more than 20 species): Burseraceae [<xref rid="RSPB20161587C27" ref-type="bibr">27</xref>] and <italic>Inga</italic> (KG Dexter & RT Pennington 2013, unpublished data) were used to investigate whether patterns of PS at the genus level were consistent with species-level patterns. Species-level trait data were also used to account for intrageneric variation in the genus-level analyses of PS: the species-level data were used to calculate the standard error of each trait within each genus, and these values were incorporated into the calculations of PS (described below) [<xref rid="RSPB20161587C34" ref-type="bibr">34</xref>]. In the methods below, all the details are given for trait values calculated at the genus level; similar calculations and methods were used at the species level.</p><p>Potential tree size, mean and maximum growth rates were all calculated in terms of tree diameter, basal area and biomass for each genus with at least 20 individuals across multiple censuses.</p><p>Potential tree size was estimated as the 95th percentile of the size distribution of all trees within each genus. For trees with multiple measurements, we selected the maximum size across different censuses to define these distributions. Tree above-ground biomass per stem was calculated using the pan-tropical, three parameter allometric equation (diameter, wood density and <italic>E</italic>) of Chave, Rejou-Mechain [<xref rid="RSPB20161587C35" ref-type="bibr">35</xref>], which assumes that tree diameter–height relationships depend linearly on bioclimatic variables (<italic>E</italic>), where <italic>E</italic> is a measurement of environmental stress based on measures of temperature seasonality and precipitation seasonality derived from the WorldClim dataset [<xref rid="RSPB20161587C28" ref-type="bibr">28</xref>] and a measure of Climatic Water Deficit extracted from a global gridded dataset [<xref rid="RSPB20161587C35" ref-type="bibr">35</xref>]. Palm biomass was estimated using a palm-specific allometric equation based on diameter [<xref rid="RSPB20161587C36" ref-type="bibr">36</xref>].</p><p>For each genus, we computed both mean growth rate and the 95th percentile of growth rates, to represent maximum growth rates within each genus, across all stems. To calculate these parameters, mean stem-level growth rate was first estimated as the mean growth per year across multiple censuses and maximum stem-level growth as the maximum growth rate per year calculated across multiple censuses. Trees with mean negative growth rates (0.9% of stems) were excluded in order to normalize the data (similar to [<xref rid="RSPB20161587C37" ref-type="bibr">37</xref>]). We also excluded palms, which do not have secondary growth, nine trees exhibiting diameter growth greater than 80 mm yr<sup>−1</sup> which may represent recording errors and stems where diameter measurements were not made using a tape measure (0.12% of all stems). If a change in the point of measurement (POM) was made during the measurement record of any given tree, we calculated growth rates using the arithmetic mean of the diameter measured at the original POM and the diameter at the new POM [<xref rid="RSPB20161587C38" ref-type="bibr">38</xref>].</p><p>Mortality rates were estimated for all genera with a minimum of 100 individuals in the plot data, based on the number of individuals found alive in the initial and final censuses of each plot. To estimate average mortality rates within each genus, the survival probability of individual trees within each clade was modelled as an exponentially declining function of the monitoring period while accounting for variation in tree size [<xref rid="RSPB20161587C39" ref-type="bibr">39</xref>,<xref rid="RSPB20161587C40" ref-type="bibr">40</xref>].</p><p>To account for the wide range of environmental conditions across plots [<xref rid="RSPB20161587C29" ref-type="bibr">29</xref>], we used mixed models to calculate genus-level values of potential tree size, mean and maximum growth rates and mortality rates while accounting for systematic variation in these parameters among plots [<xref rid="RSPB20161587C40" ref-type="bibr">40</xref>] (see the electronic supplementary material, S2).</p><p>Wood density data were extracted from the Global Wood Density database [<xref rid="RSPB20161587C41" ref-type="bibr">41</xref>,<xref rid="RSPB20161587C42" ref-type="bibr">42</xref>] and average values calculated for each genus in the phylogeny [<xref rid="RSPB20161587C43" ref-type="bibr">43</xref>].</p></sec><sec id="s2c"><label>(c)</label><title>Trait correlations</title><p>To identify relationships among genus-level traits, we conducted a phylogenetic principal component analysis (PPCA) [<xref rid="RSPB20161587C44" ref-type="bibr">44</xref>] including genera where we have a complete set of trait data. PPCA incorporates the expected correlation among traits due to their shared evolutionary history into the principal component analyses [<xref rid="RSPB20161587C45" ref-type="bibr">45</xref>]. We standardized trait values to a mean of zero and unit variance to ensure that each trait contributed equally to the PPCA.</p></sec><sec id="s2d"><label>(d)</label><title>Phylogenetic signal</title><p>In order to estimate PS for traits, we used Blomberg's <italic>K</italic> [<xref rid="RSPB20161587C1" ref-type="bibr">1</xref>]. This metric quantifies the amount of variance in an observed trait in relation to the expected trait variance under a Brownian motion model of evolution [<xref rid="RSPB20161587C1" ref-type="bibr">1</xref>,<xref rid="RSPB20161587C4" ref-type="bibr">4</xref>]. Under this model of evolution, trait values drift randomly over time, with small changes being more likely than large changes within a given unit of time (trait values at <italic>t</italic><sub>1</sub> are chosen from a normal distribution centred on the trait value at <italic>t</italic><sub>0</sub>). This model generates trait data where the covariance among trait values for taxa is proportional to the duration of their shared evolutionary history [<xref rid="RSPB20161587C4" ref-type="bibr">4</xref>]. Values of <italic>K</italic> equal to 0 indicate that there is no PS, while <italic>K</italic> equal to 1 indicates high PS and is the expected value under a Brownian motion model of evolution. Intermediate values (0 < <italic>K</italic> < 1) indicate intermediate levels of PS. To assess significance in <italic>K</italic>, we recalculated <italic>K</italic> on the tree with randomized tips a thousand times, and compared the simulated values with the observed value of <italic>K</italic>. If the observed value fell outside the range given by 2.5–97.5 percentiles of the simulated values, this value was considered significant.</p><p>We accounted for intrageneric trait variation in the calculation of <italic>K</italic> by measuring the standard error for each genus, treating individual genera as species and intrageneric variation as intraspecific variation <italic>sensu</italic> [<xref rid="RSPB20161587C34" ref-type="bibr">34</xref>]. For genera where the standard error could not be computed, we assigned the mean value of the standard error for all genera with estimates for multiple species [<xref rid="RSPB20161587C34" ref-type="bibr">34</xref>]. Including this within-genus variation allows us to account for uncertainty in trait estimation (e.g. population variation and measurement error), improve parameter estimation and reduce bias in the calculation of PS [<xref rid="RSPB20161587C1" ref-type="bibr">1</xref>,<xref rid="RSPB20161587C34" ref-type="bibr">34</xref>].</p><p>We also calculated PS using Pagel's <italic>λ</italic> [<xref rid="RSPB20161587C46" ref-type="bibr">46</xref>] in order to explore whether our results were dependent on the particular method used to calculate PS (see the electronic supplementary material, S3).</p></sec><sec id="s2e"><label>(e)</label><title>Sensitivity analysis</title><p>To investigate whether our results were affected by the spatial scale of our study, we repeated our analyses using 26 plots within 55 km of each other near Manaus. Similarly, to verify whether our results were affected by our use of genus-level data, we conducted the same analyses at the species level for the genus <italic>Inga</italic> and the Protieae (Burseraceae). Likewise, to investigate whether the number of lineages included in the analyses affected the extent of PS, we repeated the calculations of PS with just the genera with a complete set of trait values (214).</p><p>Statistical analyses were performed in the R v. 3.1.1 program [<xref rid="RSPB20161587C47" ref-type="bibr">47</xref>], using ape [<xref rid="RSPB20161587C48" ref-type="bibr">48</xref>], phytools [<xref rid="RSPB20161587C44" ref-type="bibr">44</xref>] and data.table [<xref rid="RSPB20161587C49" ref-type="bibr">49</xref>] packages.</p></sec></sec><sec id="s3"><label>3.</label><title>Results</title><sec id="s3a"><label>(a)</label><title>Trait data</title><p>All traits measured varied substantially among genera (<xref ref-type="table" rid="RSPB20161587TB1">table 1</xref> and <xref ref-type="fig" rid="RSPB20161587F2">figure 2</xref>): wood density varied eightfold, potential size in tree diameter 12-fold, potential size in biomass 814-fold, maximum growth rates in tree diameter 23-fold, mean diameter growth rates 35-fold and mortality rates 275-fold. Overall, the trait values after correcting for environmental variation and those estimated directly from the database without accounting for variation among plots were highly correlated with each other (<italic>p</italic> < 0.001 in all cases and <italic>τ</italic> ranging from 0.59 to 0.79).
<fig id="RSPB20161587F2" orientation="portrait" position="float"><label>Figure 2.</label><caption><p>Phylogeny (based on rbcL and matK plastid gene) of 497 Amazonian tree and palm genera. Number of genera varied in the different phylogenies according to the selection criterion for each trait (see Material and methods). Branches are coloured according to (<italic>a</italic>) wood density (wd g.cm<sup>3</sup>), (<italic>b</italic>) potential tree size in diameter (Max.D cm), (<italic>c</italic>) maximum tree growth in diameter (Max.gr cm yr<sup>−1</sup>) and (<italic>d</italic>) mortality rates (% yr<sup>−1</sup>). Continuous traits were coloured using a continuous colour gradient, with colour codes indicate the wide range of trait values, from blue to red, indicating higher and lower trait values, respectively. Phylogenies for each trait with all tips labelled are available in the electronic supplementary material (S6).</p></caption><graphic xlink:href="rspb20161587-g2"></graphic></fig><table-wrap id="RSPB20161587TB1" orientation="portrait" position="float"><label>Table 1.</label><caption><p>Summary of trait data, including number of genera per trait, number of species and number of individuals used for selection criterion, minimum, maximum and mean trait values per genera. In addition, PS for absolute trait values, accounting for intrageneric variation, environmental variation and both environmental and intrageneric variation. PS measured using Blomberg's <italic>K</italic>. Statistical significance in probability tests are indicated by asterisks.</p></caption><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col></colgroup><thead valign="bottom"><tr><th rowspan="5" align="left" colspan="1"></th><th rowspan="5" align="left" colspan="1">traits</th><th rowspan="5" align="left" colspan="1">units</th><th rowspan="5" align="left" colspan="1">no. ind</th><th rowspan="5" align="left" colspan="1">no. genera</th><th rowspan="5" align="left" colspan="1">no. species</th><th rowspan="5" align="left" colspan="1">range</th><th rowspan="5" align="left" colspan="1">mean</th><th colspan="4" align="left" rowspan="1">PS (<italic>K</italic>)<hr></hr></th></tr><tr><th colspan="4" align="left" rowspan="1">intrageneric variation<hr></hr></th></tr><tr><th align="left" rowspan="1" colspan="1">no<hr></hr></th><th align="left" rowspan="1" colspan="1">yes<hr></hr></th><th align="left" rowspan="1" colspan="1">no<hr></hr></th><th align="left" rowspan="1" colspan="1">yes<hr></hr></th></tr><tr><th colspan="4" align="left" rowspan="1">environmental variation<hr></hr></th></tr><tr><th align="left" rowspan="1" colspan="1">no</th><th align="left" rowspan="1" colspan="1">no</th><th align="left" rowspan="1" colspan="1">yes</th><th align="left" rowspan="1" colspan="1">yes</th></tr></thead><tbody><tr><td rowspan="1" colspan="1">wood density</td><td rowspan="1" colspan="1">wd</td><td rowspan="1" colspan="1">g.cm<sup>3</sup></td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">497</td><td rowspan="1" colspan="1">1324</td><td rowspan="1" colspan="1">0.15–1.21</td><td rowspan="1" colspan="1">0.61</td><td rowspan="1" colspan="1">0.26***</td><td rowspan="1" colspan="1">0.30***</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td></tr><tr><td rowspan="5" colspan="1">potential size</td><td rowspan="1" colspan="1">maximum diameter</td><td rowspan="1" colspan="1">cm</td><td rowspan="1" colspan="1">244 362</td><td rowspan="1" colspan="1">383</td><td rowspan="1" colspan="1">1412</td><td rowspan="1" colspan="1">14.5–171.1</td><td rowspan="1" colspan="1">459.4</td><td rowspan="1" colspan="1">0.23***</td><td rowspan="1" colspan="1">0.31***</td><td rowspan="1" colspan="1">0.20***</td><td rowspan="1" colspan="1">0.29***</td></tr><tr><td rowspan="1" colspan="1">maximum diameter × wd</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">244 362</td><td rowspan="1" colspan="1">383</td><td rowspan="1" colspan="1">1412</td><td rowspan="1" colspan="1">4.94–154.69</td><td rowspan="1" colspan="1">28.08</td><td rowspan="1" colspan="1">0.27***</td><td rowspan="1" colspan="1">0.34***</td><td rowspan="1" colspan="1">0.25***</td><td rowspan="1" colspan="1">0.32***</td></tr><tr><td rowspan="1" colspan="1">maximum basal area</td><td rowspan="1" colspan="1">m<sup>2</sup></td><td rowspan="1" colspan="1">244 362</td><td rowspan="1" colspan="1">383</td><td rowspan="1" colspan="1">1412</td><td rowspan="1" colspan="1">0.02–2.3</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.23***</td><td rowspan="1" colspan="1">0.31***</td><td rowspan="1" colspan="1">0.21**</td><td rowspan="1" colspan="1">0.26***</td></tr><tr><td rowspan="1" colspan="1">maximum basal area × wd</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">244 362</td><td rowspan="1" colspan="1">383</td><td rowspan="1" colspan="1">1412</td><td rowspan="1" colspan="1">0.01–0.13</td><td rowspan="1" colspan="1">0.13</td><td rowspan="1" colspan="1">0.26***</td><td rowspan="1" colspan="1">0.32***</td><td rowspan="1" colspan="1">0.23***</td><td rowspan="1" colspan="1">0.29***</td></tr><tr><td rowspan="1" colspan="1">maximum biomass</td><td rowspan="1" colspan="1">kg</td><td rowspan="1" colspan="1">244 362</td><td rowspan="1" colspan="1">383</td><td rowspan="1" colspan="1">1412</td><td rowspan="1" colspan="1">54.63–44443.1</td><td rowspan="1" colspan="1">2760.6</td><td rowspan="1" colspan="1">0.25***</td><td rowspan="1" colspan="1">0.28***</td><td rowspan="1" colspan="1">0.22***</td><td rowspan="1" colspan="1">0.28***</td></tr><tr><td rowspan="6" colspan="1">growth rates</td><td rowspan="1" colspan="1">maximum growth in diameter</td><td rowspan="1" colspan="1">cm yr<sup>−1</sup></td><td rowspan="1" colspan="1">134 303</td><td rowspan="1" colspan="1">329</td><td rowspan="1" colspan="1">1024</td><td rowspan="1" colspan="1">0.19–4.38</td><td rowspan="1" colspan="1">0.93</td><td rowspan="1" colspan="1">0.19***</td><td rowspan="1" colspan="1">0.25***</td><td rowspan="1" colspan="1">0.18***</td><td rowspan="1" colspan="1">0.25***</td></tr><tr><td rowspan="1" colspan="1">maximum growth in basal area</td><td rowspan="1" colspan="1">m<sup>2</sup> yr<sup>−1</sup></td><td rowspan="1" colspan="1">134 303</td><td rowspan="1" colspan="1">329</td><td rowspan="1" colspan="1">1024</td><td rowspan="1" colspan="1">0.003–0.03</td><td rowspan="1" colspan="1">0.005</td><td rowspan="1" colspan="1">0.22***</td><td rowspan="1" colspan="1">0.32***</td><td rowspan="1" colspan="1">0.21***</td><td rowspan="1" colspan="1">0.29***</td></tr><tr><td rowspan="1" colspan="1">maximum growth in biomass</td><td rowspan="1" colspan="1">kg yr<sup>−1</sup></td><td rowspan="1" colspan="1">134 303</td><td rowspan="1" colspan="1">329</td><td rowspan="1" colspan="1">1024</td><td rowspan="1" colspan="1">0.21–95.23</td><td rowspan="1" colspan="1">6.17</td><td rowspan="1" colspan="1">0.25***</td><td rowspan="1" colspan="1">0.39***</td><td rowspan="1" colspan="1">0.23***</td><td rowspan="1" colspan="1">0.33***</td></tr><tr><td rowspan="1" colspan="1">mean growth in diameter</td><td rowspan="1" colspan="1">cm yr<sup>−1</sup></td><td rowspan="1" colspan="1">133 656</td><td rowspan="1" colspan="1">327</td><td rowspan="1" colspan="1">1000</td><td rowspan="1" colspan="1">0.05–1.74</td><td rowspan="1" colspan="1">0.26</td><td rowspan="1" colspan="1">0.18***</td><td rowspan="1" colspan="1">0.25***</td><td rowspan="1" colspan="1">0.19***</td><td rowspan="1" colspan="1">0.29***</td></tr><tr><td rowspan="1" colspan="1">mean growth in basal area</td><td rowspan="1" colspan="1"> m<sup>2</sup> yr<sup>−1</sup></td><td rowspan="1" colspan="1">133 656</td><td rowspan="1" colspan="1">327</td><td rowspan="1" colspan="1">1000</td><td rowspan="1" colspan="1">0–0.01</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0.20***</td><td rowspan="1" colspan="1">0.27***</td><td rowspan="1" colspan="1">0.19***</td><td rowspan="1" colspan="1">0.29***</td></tr><tr><td rowspan="1" colspan="1">mean growth in biomass</td><td rowspan="1" colspan="1">kg yr<sup>−1</sup></td><td rowspan="1" colspan="1">133 656</td><td rowspan="1" colspan="1">327</td><td rowspan="1" colspan="1">1000</td><td rowspan="1" colspan="1">0.15–21.76</td><td rowspan="1" colspan="1">1.67</td><td rowspan="1" colspan="1">0.23***</td><td rowspan="1" colspan="1">0.30***</td><td rowspan="1" colspan="1">0.19***</td><td rowspan="1" colspan="1">0.25***</td></tr><tr><td rowspan="1" colspan="1">mortality</td><td rowspan="1" colspan="1">mean stem mortality</td><td rowspan="1" colspan="1">% yr<sup>−1</sup></td><td rowspan="1" colspan="1">156 495</td><td rowspan="1" colspan="1">221</td><td rowspan="1" colspan="1">306</td><td rowspan="1" colspan="1">0.04–10.98</td><td rowspan="1" colspan="1">1.08</td><td rowspan="1" colspan="1">0.17**</td><td rowspan="1" colspan="1">0.25**</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td></tr><tr><td rowspan="1" colspan="1">PPCA1</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">214</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">0.18**</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td></tr><tr><td rowspan="1" colspan="1">PPCA2</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">214</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">0.21***</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td><td rowspan="1" colspan="1">—</td></tr></tbody></table><table-wrap-foot><fn><p>***<italic>p</italic> < 0.001, **<italic>p</italic> < 0.05, *<italic>p</italic> < 0.1.</p></fn></table-wrap-foot></table-wrap></p></sec><sec id="s3b"><label>(b)</label><title>Trait relationships</title><p>Trait associations among lineages were analysed with a PPCA: 83% of the variation in the four-dimensional space was accounted for by the first two axes (<xref ref-type="fig" rid="RSPB20161587F3">figure 3</xref>). The first axis (PPCA1) explained 52.8% of the variation and shows strong positive loadings for mortality and maximum growth rates, while wood density was negatively associated with this axis (electronic supplementary material, S4). PPCA1 thus represents a continuum from pioneer and light demanding lineages with low wood density and fast demographic traits (e.g. high mortality and growth rates) to non-pioneer lineages with high wood density and slow demographic rates. The second axis (PPCA2) explained 30.5% of the variation and was associated more closely with potential tree size, and reflects the variation from individuals of understory genera, to individuals of canopy and emergent lineages (<xref ref-type="fig" rid="RSPB20161587F3">figure 3</xref>).
<fig id="RSPB20161587F3" orientation="portrait" position="float"><label>Figure 3.</label><caption><p>PPCAs for the first two principal components with PC loadings for the four traits studied here: wood density, potential tree size in terms of diameter (Max. diameter), potential growth rates in terms of diameter (Maxgr. growth rate) and annual mortality rates (Mortality rate). Points represent 221 genera of trees; position of 22 key genera marked in bold and named.</p></caption><graphic xlink:href="rspb20161587-g3"></graphic></fig></p></sec><sec id="s3c"><label>(c)</label><title>Phylogenetic signal</title><p>All traits and the first two PPCA axes exhibited significant PS, with closely related genera being more similar than expected by chance, using either Bloomberg's <italic>K</italic> (<xref ref-type="table" rid="RSPB20161587TB1">table 1</xref>) or Pagel's <italic>λ</italic> (electronic supplementary material, S3). Because estimates of Pagel's <italic>λ</italic> and Blomberg's <italic>K</italic> are strongly correlated and most studies of PS in tropical trees have focused on the <italic>K</italic> metric rather than <italic>λ</italic>, we focus our results and discussion on the calculations using Blomberg's <italic>K-</italic>value.</p><p>Traits showed significant and similar values for <italic>K</italic>, varying from 0.25 to 0.39 and from 0.17 to 0.27, with and without accounting for intrageneric variation, respectively. These <italic>K</italic>-values indicate that evolutionarily related genera tend to be more similar to each other, but less than expected under a BM model of evolution (<xref ref-type="table" rid="RSPB20161587TB1">table 1</xref>). Finally, removing the environmental contribution to trait variation did not substantially alter the magnitude of PS (<xref ref-type="table" rid="RSPB20161587TB1">table 1</xref>).</p></sec><sec id="s3d"><label>(d)</label><title>Sensitivity analyses</title><p>Although using just the Manaus plot data significantly reduced the number of genera, species and individual trees included in the analyses, PS at smaller spatial scales showed similar patterns to PS calculated using the whole dataset (electronic supplementary material, S5). Similarly, reducing the number of lineages to genera we had all trait values showed congruent patterns of PS (electronic supplementary material, S5). In addition, all traits showed similar or slightly higher Blomberg's <italic>K</italic> values for just <italic>Inga</italic> or Protieae than for all taxa together (electronic supplementary material, S5).</p></sec></sec><sec id="s4"><label>4.</label><title>Discussion</title><p>This is the first study, to our knowledge, to investigate the extent of PS for traits that quantify the main axes of life-history variation in survival and growth of trees at such a large phylogenetic and spatial scale. Our results demonstrate that for Amazonian forests, closely related genera have similar life-history strategies, with all traits showing similar levels of PS (<xref ref-type="table" rid="RSPB20161587TB1">table 1</xref> and <xref ref-type="fig" rid="RSPB20161587F2">figure 2</xref>; electronic supplementary material, S6). The similar level of PS found across all the different, correlated traits suggests that the main axes of life-history variation among lineages of Amazonian trees may represent the result of repeated evolution of a suite of coordinated functional characteristics.</p><sec id="s4a"><label>(a)</label><title>Relationships among traits</title><p>Strong correlations among traits were represented by two major axes of variation, which are likely to be associated with adaptations to horizontal and vertical light gradients. Ecological differences among species adapted to gaps versus the shaded understory or to the understory versus the canopy are well established as the principal axes of functional variation among tropical forest tree species [<xref rid="RSPB20161587C50" ref-type="bibr">50</xref>,<xref rid="RSPB20161587C51" ref-type="bibr">51</xref>]. The first axis runs from pioneer and light demanding genera with low wood density and fast demographic traits (e.g. high mortality and high growth rates) to shade tolerant genera with dense wood and slow demographic traits. The second axis represents variation in tree size and contrasts understorey genera, from lineages of canopy trees. For example, these axes distinguish <italic>Cecropia</italic> and <italic>Croton</italic>, classic pioneers with low wood density and fast demographic traits, from <italic>Hirtella</italic>—a typically dense-wooded and slow-growing understory genus of trees. Lineages of emergent trees which all achieve very large potential tree sizes (e.g. <italic>Bertholletia</italic>, <italic>Ceiba</italic>, <italic>Hura, Dipteryx</italic>), are also distinguished in this analysis by their different wood densities and growth rates (<xref ref-type="fig" rid="RSPB20161587F3">figure 3</xref>).</p></sec><sec id="s4b"><label>(b)</label><title>Phylogenetic signal</title><p>Our results demonstrate significant levels of PS among demographic and structural traits of tropical trees, with Blomberg's <italic>K</italic> ranging from 0.25 to 0.39. This pattern suggests that evolutionary relationships provide useful information about the ecological similarity of these lineages. However, while our analyses of PS shows that evolutionarily related lineages have more similar traits than expected by chance, their values are lower than expected under a pure BM model of evolution (<xref ref-type="table" rid="RSPB20161587TB1">table 1</xref> and <xref ref-type="fig" rid="RSPB20161587F2">figure 2</xref>) under which <italic>K</italic>-values would be close to 1. PS can be lower than expected under BM if there is convergent evolution across distantly related lineages and/or divergent selection among closely related groups [<xref rid="RSPB20161587C3" ref-type="bibr">3</xref>,<xref rid="RSPB20161587C4" ref-type="bibr">4</xref>]. This result therefore suggests that there has been repeated convergent evolution and/or divergent selection, along the two main axes of variation identified by the PPCA analysis (<xref ref-type="fig" rid="RSPB20161587F3">figure 3</xref>). This finding suggests that adaptations to light gaps, or understorey and canopy light environments, have repeatedly evolved within multiple lineages of tropical trees as shown by the different pioneer and shade tolerant genera within a series of unrelated families (e.g. <italic>Cecropia</italic> versus <italic>Brosimum</italic> (Urticaceae/Moraceae), <italic>Vismia</italic> versus <italic>Calophyllum</italic> (Clusiaceae) and <italic>Inga</italic> versus <italic>Dipteryx</italic>/<italic>Parkia</italic> (Fabaceae); <xref ref-type="fig" rid="RSPB20161587F2">figure 2</xref>).</p></sec><sec id="s4c"><label>(c)</label><title>Sensitivity analyses</title><p>The PS found here for trees across lowland closed-canopy South American forests is generally stronger than previously reported in the literature for tropical forests in smaller scale analyses (electronic supplementary material, S7). In previous studies, some traits showed low but significant PS [<xref rid="RSPB20161587C6" ref-type="bibr">6</xref>,<xref rid="RSPB20161587C7" ref-type="bibr">7</xref>,<xref rid="RSPB20161587C13" ref-type="bibr">13</xref>], while others have even found that traits are randomly dispersed over the phylogeny [<xref rid="RSPB20161587C10" ref-type="bibr">10</xref>,<xref rid="RSPB20161587C11" ref-type="bibr">11</xref>]. However, although <italic>K</italic>-values are standardized to allow comparison between traits and phylogenetic trees [<xref rid="RSPB20161587C1" ref-type="bibr">1</xref>,<xref rid="RSPB20161587C4" ref-type="bibr">4</xref>], direct comparisons of PS are affected by differences in the spatial and taxonomic scale of the studies, the number of lineages and the use of different kinds of phylogenies.</p><p>A first issue for comparing the extent of PS among studies is variation in spatial scales. However, here we show that the higher PS in this study is unlikely to be an artefact of our larger spatial scale: restricting our analyses to 26 plots around Manaus shows consistent patterns, with similar levels of PS for all traits compared to analyses for the whole Amazon (electronic supplementary material, S5).</p><p>Secondly, different numbers of lineages in different studies may play a role in determining variation in the extent of PS. Although Blomberg's <italic>K</italic> is efficient at detecting the strength of similarity among closely related lineages for sample sizes greater than 20 [<xref rid="RSPB20161587C1" ref-type="bibr">1</xref>], the ability to detect different levels of PS may increase with larger sample sizes [<xref rid="RSPB20161587C52" ref-type="bibr">52</xref>]. To address this issue, we conducted a set of analyses restricted to genera for which we had all trait values (214 genera). As estimates of <italic>K</italic> are highly consistent when we include fewer genera (electronic supplementary material, S5), it appears that the number of lineages is unlikely to have caused the observed trends of high levels of PS for our traits.</p><p>Thirdly, most of previous studies [<xref rid="RSPB20161587C6" ref-type="bibr">6</xref>,<xref rid="RSPB20161587C7" ref-type="bibr">7</xref>,<xref rid="RSPB20161587C10" ref-type="bibr">10</xref>–<xref rid="RSPB20161587C12" ref-type="bibr">12</xref>] were conducted at the species level, and taxonomic scale can also affect the degree of PS. PS in any trait may vary at different taxonomic scales; a single trait can have high similarity at one level (e.g. genus level) but this pattern can break down at higher or lower taxonomic levels [<xref rid="RSPB20161587C52" ref-type="bibr">52</xref>]. Here, the PS of these traits at the species level within the Protieae and <italic>Inga</italic> were similar or slightly greater than for the genus-level results (electronic supplementary material, S5), suggesting that our results are consistent at finer taxonomic levels. However, as our analyses at low taxonomic levels were limited to two lineages it remains to be fully tested whether the result indeed holds within all clades of Neotropical trees.</p><p>Finally, the use of different kinds of phylogenies is likely to affect the extent of similarity among related species that is reported in different studies (electronic supplementary material, S7). Much previous work was carried out using community-level phylogenies, restricted to locally co-occurring species [<xref rid="RSPB20161587C6" ref-type="bibr">6</xref>,<xref rid="RSPB20161587C12" ref-type="bibr">12</xref>] and in many cases using unresolved phylogenies with relationships represented as polytomies [<xref rid="RSPB20161587C11" ref-type="bibr">11</xref>]. Such community-level phylogenies may lack sister lineages for many clades that may be critical to effectively measure PS. In addition, the use of trees with many polytomies, e.g. those which add genera and species as polytomies onto backbone family-level trees [<xref rid="RSPB20161587C53" ref-type="bibr">53</xref>], leads to uncertainty in PS estimates [<xref rid="RSPB20161587C14" ref-type="bibr">14</xref>]. More importantly, phylogenetic sampling may play a major role in determining the extent of PS. Although the genus-level phylogeny used here is far from complete, our analyses do encompass a far wider range of lineages than previous studies, including the major angiosperm lineages present in the Amazon basin.</p><p>Our results demonstrate that there is significant PS for key demographic and structural traits in tropical forests. This finding opens the way for clearer predictions of how evolutionary diversity relates to ecosystem structure and function, and how different drivers will, in turn, affect the evolutionary diversity of Amazonian forests. For example, this study suggests that community-level measures of evolutionary relatedness among species are likely to be good predictors of the structure and functioning of these ecosystems [<xref rid="RSPB20161587C8" ref-type="bibr">8</xref>,<xref rid="RSPB20161587C9" ref-type="bibr">9</xref>]. These results also indicate that changes in environmental conditions or disturbance regimes that favour particular life-history strategies will ultimately erode evolutionary diversity [<xref rid="RSPB20161587C54" ref-type="bibr">54</xref>,<xref rid="RSPB20161587C55" ref-type="bibr">55</xref>], although the presence of some convergent evolution across lineages may prevent significant loss of phylogenetic diversity over some scales of anthropogenic disturbance [<xref rid="RSPB20161587C56" ref-type="bibr">56</xref>]. Our results may therefore help to resolve why different studies of the effect of disturbance on phylogenetic diversity have obtained contrasting results [<xref rid="RSPB20161587C54" ref-type="bibr">54</xref>–<xref rid="RSPB20161587C56" ref-type="bibr">56</xref>]: in particular, this study suggests that investigating the PS of traits that influence species ability to persist after disturbance within the species pool of interest will be critical to understand how disturbance will alter phylogenetic diversity. Finally, our results also suggest that any long-term changes in the evolutionary diversity of intact Amazonian forests may indicate functional shifts in these diverse ecosystems. Overall, the phylogenetic structure of life-history strategies within Amazon tree communities described in this study helps to provide a predictive framework to understand how such complex systems will respond to global change and anthropogenic disturbance.</p></sec></sec><sec sec-type="supplementary-material"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="SD1"><caption><title>List of plot data used in the present study</title></caption><media mimetype="application" mime-subtype="pdf" xlink:href="rspb20161587supp1.pdf" orientation="portrait" id="d35e2694" position="anchor"></media></supplementary-material></sec><sec sec-type="supplementary-material"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="SD2"><caption><title>Methods for calculating trait intrinsic value</title></caption><media mimetype="application" mime-subtype="pdf" xlink:href="rspb20161587supp2.pdf" orientation="portrait" id="d35e2702" position="anchor"></media></supplementary-material></sec><sec sec-type="supplementary-material"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="SD3"><caption><title>Supporting reults</title></caption><media mimetype="application" mime-subtype="pdf" xlink:href="rspb20161587supp3.pdf" orientation="portrait" id="d35e2710" position="anchor"></media></supplementary-material></sec><sec sec-type="supplementary-material"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="SD4"><caption><title>Phylogeny of 497 Amazonian tree genera</title></caption><media mimetype="application" mime-subtype="pdf" xlink:href="rspb20161587supp4.pdf" orientation="portrait" id="d35e2718" position="anchor"></media></supplementary-material></sec><sec sec-type="supplementary-material"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="SD5"><caption><title>Comparison between published values of phylogenetic signal and values found in the present study</title></caption><media mimetype="application" mime-subtype="pdf" xlink:href="rspb20161587supp5.pdf" orientation="portrait" id="d35e2726" position="anchor"></media></supplementary-material></sec></body><back><ack><title>Acknowledgements</title><p>We thank Ademir Ruschel, Alvaro Cogollo, Anand Roopsind, Antonio Lola da Costa, Antonio S. Lima, Armando Torres-Lezama, Atila Oliveira, Carlos Céron, Claudinei Santos, Eder das Neves, Eliana Jimenez-Rojas, Foster Brown, Freddy Ramirez Arevalo, Georgia Pickvance, Gloria Galeano, Guido Pardo, Ieda Amaral, Ima Vieira, James Singh, John Terborgh, Juan Licona, Luzmilla Arroyo, Marielos Peña-Claros, Marisol Toledo, Natalino Silva, Patricia Loayza and Wenderson Castro for allowing use of their data and the many colleagues and field assistants who have contributed to the development of the RAINFOR network. This paper is 705 in the Technical Series of the Biological Dynamics Fragments Project (BDFFP-INPA/STRI).</p></ack><sec id="s5" sec-type="dataaccess"><title>Data accessibility</title><p>The permanently archived data package of the genus-level trait dataset (wood density, potential tree size, maximum and mean growth rates and mortality rates) are available from <uri xlink:href="https://doi.org/10.5521/FORESTPLOTS.NET/2016_4">https://doi.org/10.5521/FORESTPLOTS.NET/2016_4</uri>.</p></sec><sec id="s6" sec-type="aucontribute"><title>Authors' contributions</title><p>F.C.S. and T.R.B. conceived the study; F.C.S., T.R.B. and K.G.D. designed the study; F.C.S. analysed data and wrote the paper; all co-authors collected field data or data management and analyses tools. O.L.P., Y.M. and J.L. conceived the RAINFOR forest census plot network. K.G.D., J.C. and R.T.P. contributed phylogenetic data used in this study. All co-authors commented and/or approved the manuscript.</p></sec><sec id="s7" sec-type="competing"><title>Competing Interests</title><p>We declare we have no competing interests.</p></sec><sec id="s8" sec-type="funding"><title>Funding</title><p>The field data used in this study have been generated by the RAINFOR network, which has been supported by a Gordon and Betty Moore Foundation grant, the European Union's Seventh Framework Programme projects 283080, ‘GEOCARBON’; and 282664, ‘AMAZALERT’; ERC grant ‘Tropical Forests in the Changing Earth System’), and Natural Environment Research Council (NERC) Urgency, Consortium and Standard Grants ‘AMAZONICA’ (NE/F005806/1), ‘TROBIT’ (NE/D005590/1) and ‘Niche Evolution of South American Trees’ (NE/I028122/1). Additional data were included from the Tropical Ecology Assessment and Monitoring (TEAM) Network — a collaboration between Conservation International, the Missouri Botanical Garden, the Smithsonian Institution and the Wildlife Conservation Society, and partly funded by these institutions, the Gordon and Betty Moore Foundation, and other donors. Fieldwork was also partially supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico of Brazil (CNPq), project Programa de Pesquisas Ecológicas de Longa Duração (PELD-403725/2012-7). F.C.S. is supported by a PhD scholarship from Coordination for the Improvement of Higher Education Personnel - Brasil (CAPES) (117913-6). 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