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In a field study conducted at 38 sites in two regions, we measured the abundance of alien invasive species, species richness of the plant community, total and soil extractable pools of P and N, soil phosphatase activity and the root phosphatase activity of nine common plant species. An analysis of the data using structural equation modelling (SEM) revealed no significant relationship between soil extractable-P concentrations and the abundance of alien plants (Fig. 1), despite the fact that previous studies in Cerrado have found P fertilization to promote the invasion of alien species31. However, the SEM did find abundance of alien invasive plants to be influenced by native species richness in two contrasting ways. One way was a direct negative relationship between species richness and the abundance of invasive species, which is consistent with the stochastic niche hypothesis and with results of some previous studies14,15,16. This pattern was also observed in a direct regression between the two variables (Suppl. Figure 2). The other way was an indirect and positive effect of species richness that was mediated via phosphatase, suggesting that invasive plants may benefit from organic P released through phosphatase produced by soil microbes and/or plant roots.
Figure 1

Structural equation model (SEM) showing direct (blue arrow) and indirect (orange arrow) connections between plant species richness and the abundance of alien plants in the Cerrado. The possible connection between species richness and soil phosphatase activity (PME) follows results obtained in the Jena Biodiversity Experiment29. Also connections between the total soil P and soil extractable P (Mehlich) pools on soil phosphatase activity, as well as a direct connection between soil extractable P and abundance of alien plants are included in the SEM. Plant variables are recorded on 334-m2 plots using the Braun–Blanquet scale, soil parameters are from the top 10-cm soil. Numbers associated with paths between variables are path coefficients presented as standardized values (scaled by the standard deviations of the variables). Solid arrows show significant connections (*p  More

Computational Research Division, Lawrence Berkeley National Laboratory, Berkeley, CA, 94720, USA
Gilberto Pastorello, Danielle Christianson, You-Wei Cheah, Abdelrahman Elbashandy, Catharine van Ingen & Deb Agarwal

DIBAF, University of Tuscia, Viterbo, 01100, Italy
Carlo Trotta, Eleonora Canfora, Diego Polidori, Alessio Ribeca, Alessio Collalti, Claudia Consalvo, Giacomo Nicolini, Simone Sabbatini, Michele Tomassucci, Riccardo Valentini, Domenico Vitale & Dario Papale

Euro-Mediterranean Centre on Climate Change Foundation (CMCC), Lecce, 73100, Italy
Eleonora Canfora, Diego Polidori, Alessio Ribeca, Serena Marras, Giacomo Nicolini, Costantino Sirca, Donatella Spano, Riccardo Valentini, Domenico Vitale & Dario Papale

Climate & Ecosystem Sciences Division, Lawrence Berkeley National Laboratory, Berkeley, CA, 94720, USA
Housen Chu, Thomas Powell, Naama Raz-Yaseef, Sebastien Biraud & Margaret Torn

Department of Civil Engineering, California State University, Sacramento, CA, 95819, USA
Cristina Poindexter

Department of Geography, Environment, and Spatial Sciences, Michigan State University, East Lansing, MI, 48823, USA
Jiquan Chen

Department of Computer Science, University of Virginia, Charlottesville, VA, 22904, USA
Marty Humphrey

TERN Ecosystrem Processes, Menzies Creek, VIC3159, Australia
Peter Isaac

Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing, 100101, China
Leiming Zhang, Xiaoqin Dai, Yongtao He, Peili Shi & Huimin Wang

Department of Soil Science, University of Manitoba, Winnipeg, MB, R3T2N2, Canada
Brian Amiro

Department of Agroecology and Environment, Agroscope Research Institute, Zürich, 8046, Switzerland
Christof Ammann

School of Geography and Earth Sciences, McMaster University, L8S4K1, Hamilton, ON, Canada
M. Altaf Arain, Eric Beamesderfer & Myroslava Khomik

Department of Physical Geography and Ecosystem Science, Lund University, Lund, 22362, Sweden
Jonas Ardö, Marcin Jackowicz-Korczynski, Frans-Jan Parmentier, Norbert Pirk & Torbern Tagesson

Department of Agronomy and Horticulture, University of Nebraska-Lincoln, Lincoln, NE, 68583, USA
Timothy Arkebauer

School of Ecosystem and Forest Sciences, The University of Melbourne, Richmond, VIC3121, Australia
Stefan K. Arndt, Anne Griebel & Ian McHugh

Department of Biology, Research Group PLECO, University of Antwerp, Antwerp, 2610, Belgium
Nicola Arriga, Bert Gielen & Marilyn Roland

Joint Research Centre, European Commission, Ispra, 21027, Italy
Nicola Arriga, Ignacio Goded, Carsten Gruening & Giovanni Manca

TERRA Teaching and Research Center, University of Liege, Gembloux, B-5030, Belgium
Marc Aubinet, Anne De Ligne, Bernard Heinesch & Christine Moureaux

Finnish Meteorological Institute, Helsinki, 00560, Finland
Mika Aurela, Juha Hatakka, Tuomas Laurila, Annalea Lohila & Juha-Pekka Tuovinen

ESPM, University of California Berkeley, Berkeley, CA, 94720, USA
Dennis Baldocchi, Allen H. Goldstein, Siyan Ma, Joseph Verfaillie & Robin Weber

Global Institute for Water Security, University of Saskatchewan, Saskatoon, SK, S7N3H5, Canada
Alan Barr

Climate Research Division, Environment and Climate Change Canada, Saskatoon, SK, S7N3H5, Canada
Alan Barr

Department of Sustainable Agro-ecosystems and Bioresources, Research and Innovation Centre, Fondazione Edmund Mach, San Michele All’adige, 38010, Italy
Luca Belelli Marchesini, Mauro Cavagna, Isaac Chini, Damiano Gianelle, Barbara Marcolla & Roberto Zampedri

Department of Landscape Design and Sustainable Ecosystems, Agrarian‐Technological Institute, RUDN University, Moscow, 117198, Russia
Luca Belelli Marchesini

Direction du marché du carbone, Ministère du Développement durable de l’Environnement et de la Lutte contre les changements climatiques, Québec, QC, G1R5V7, Canada
Onil Bergeron

School of Agriculture and Environment, University of Western Australia, Crawley, 6009, Australia
Jason Beringer & Richard Silberstein

Institute of Hydrology and Meteorology, Technische Universität Dresden, Tharandt, 01737, Germany
Christian Bernhofer, Uwe Eichelmann, Thomas Grünwald, Markus Hehn, Uta Moderow & Heiko Prasse

Université Paris-Saclay, CNRS, AgroParisTech, Ecologie Systématique et Evolution, Orsay, 91405, France
Daniel Berveiller, Nicolas Delpierre & Eric Dufrêne

Biological Systems Engineering, University of Nebraska-Lincoln, Lincoln, NE, 68583, USA
Dave Billesbach & Adam J. Liska

Faculty of Land and Food Systems, University of British Columbia, Vancouver, BC, V6T1Z4, Canada
Thomas Andrew Black, Rachhpal Jassal & Zoran Nesic

Department of Geography, University of Colorado, Boulder, CO, 80309, USA
Peter D. Blanken & Sean P. Burns

Department of Civil, Environmental & Geodetic Engineering, Ohio State University, Columbus, OH, 43210, USA
Gil Bohrer

Alfred Wegener Institute Helmholtz Centre for Polar and Marine Research, Potsdam, 14482, Germany
Julia Boike

Geography Department, Humboldt-Universität zu Berlin, Berlin, Germany
Julia Boike

Forest Resources, University of Minnesota, St Paul, MN, 55108, USA
Paul V. Bolstad

Université de Lorraine, AgroParisTech, INRAE, UMR Silva, Nancy, 54000, France
Damien Bonal

ISPA, Bordeaux Sciences Agro, INRAE, Villenave d’Ornon, 33140, France
Jean-Marc Bonnefond, Denis Loustau & Virginie Moreaux

School of Biological Sciences, University of Utah, Salt Lake City, UT, 84112, USA
David R. Bowling

School of Forest Resources and Conservation, University of Florida, Gainesville, FL, 32611, USA
Rosvel Bracho

McMaster University Library, McMaster University, Hamilton, ON, L8S4L6, Canada
Jason Brodeur

Thünen Institute of Climate-Smart Agriculture, Federal Research Institute of Rural Areas, Forestry and Fisheries, Braunschweig, 38116, Germany
Christian Brümmer, Antje Lucas-Moffat & Frederik Schrader

Department of Environmental Systems Science, ETH Zurich, Zurich, 8092, Switzerland
Nina Buchmann, Werner Eugster, Iris Feigenwinter, Mana Gharun, Lukas Hörtnagl, Regine Maier & Sebastian Wolf

INRAE UMR ECOFOG, Kourou, 97387, French Guiana
Benoit Burban

Mesoscale and Microscale Meteorology Laboratory, National Center for Atmospheric Research, Boulder, CO, 80301, USA
Sean P. Burns

Université Paris-Saclay, INRAE, AgroParisTech, UMR ECOSYS, Thiverval-Grignon, 78850, France
Pauline Buysse, Pierre Cellier & Benjamin Loubet

Australian Landscape Trust, Renmark, SA5341, Australia
Peter Cale

State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing, 100093, China
Shiping Chen & Xingguo Han

Department of Bioscience, Arctic Research Center, Aarhus University, Roskilde, 4000, Denmark
Torben R. Christensen, Marcin Jackowicz-Korczynski, Efrén López-Blanco & Mikhail Mastepanov

School of Life Sciences, University of Technology Sydney, Sydney, 2007, Australia
James Cleverly & Derek Eamus

Terrestrial Ecosystem Research Network TERN, University of Technology, Sydney, 2007, Australia
James Cleverly

Institute for Agricultural and Forestry Systems in the Mediterranean, National Research Council of Italy, Ercolano, 80056, Italy
Alessio Collalti, Ettore D’Andrea, Paul di Tommasi, Daniela Famulari & Vincenzo Magliulo

Research Institute on Terrestrial Ecosystems, National Research Council of Italy, Porano, 05010, Italy
Claudia Consalvo

Biospheric Sciences Laboratory, NASA Goddard Space Flight Center, Greenbelt, MD, 20771, USA
Bruce D. Cook

Environmental Science Division, Argonne National Laboratory, Lemont, IL, 60439, USA
David Cook & Roser Matamala

Canadian Forest Service, Natural Resources Canada, Québec, QC, G1V4C7, Canada
Carole Coursolle

Centre d’étude de la forêt, Faculté de foresterie, de géographie et de géomatique, Université Laval, Québec, QC, G1V0A6, Canada
Carole Coursolle & Hank A. Margolis

Climate Change Unit, Environmental Protection Agency of Aosta Valley, Saint Christophe, 11020, Italy
Edoardo Cremonese, Gianluca Filippa & Marta Galvagno

Department of Evolution, Ecology, and Organismal Biology, Ohio State University, Columbus, OH, 43210, USA
Peter S. Curtis

Instituto de Astronomia, Geofísica e Ciências Atmosféricas, Universidade de São Paulo, São Paulo, SP, 01000-000, Brazil
Humberto da Rocha

Department of Meteorology and Atmospheric Science, The Pennsylvania State University, University Park, PA, 16802, USA
Kenneth J. Davis

Institute of Research on Terrestrial Ecosystems, National Research Council of Italy, Montelibretti, 00010, Italy
Bruno De Cinti

UMR Eco&Sols, CIRAD, Montpellier, 34060, France
Agnes de Grandcourt & Yann Nouvellon

Pedology, Embrapa Amazonia Oriental, Belém, PA, 68020640, Brazil
Raimundo C. De Oliveira

Atmospheric and Oceanic Sciences, University of Wisconsin-Madison, Madison, WI, 53706, USA
Ankur R. Desai

Departamento de Métodos Cuantitativos y Sistemas de Información, Facultad de Agronomía, UBA, Buenos Aires, 1417, Argentina
Carlos Marcelo Di Bella

Department of Earth Sciences, Vrije Universiteit Amsterdam, Amsterdam, 1081 HV, The Netherlands
Han Dolman & Ko van Huissteden

Desertification and Geoecology Department, Experimental Station of Arid Zones, CSIC, Almería, 04120, Spain
Francisco Domingo

School of Life Science, Shanxi University, Taiyuan, 030006, China
Gang Dong

HydroFocus, Davis, CA, 95618, USA
Sabina Dore

Institute of BioEconomy, National Research Council of Italy, Sassari, 07100, Italy
Pierpaolo Duce

Department of Earth, Environment, and Physics, Worcester State University, Worcester, MA, 01602, USA
Allison Dunn

Department of Matter and Energy Fluxes, Global Change Research Institute of the Czech Academy of Sciences, Brno, 60300, Czech Republic
Jiří Dušek, Dalibor Janouš, Marian Pavelka, Pavel Sedlák & Ladislav Šigut

ElObeid Research Station, Agricultural Research Corporation, ElObeid, 51111, Sudan
Hatim Abdalla M. ElKhidir

Airborne Research Australia, TERN Ecosystem Processes Central Node, Parafield, 5106, Australia
Cacilia M. Ewenz

Department of Botany, Program in Ecology, University of Wyoming, 1000 E. Univ. Ave, Laramie, WY, 82071, USA
Brent Ewers

Institute of BioEconomy, National Research Council of Italy, Rome, 00100, Italy
Silvano Fares

Research Centre for Forestry and Wood, Council for Agricultural Research and Economics, Rome, 00166, Italy
Silvano Fares

Geoscience Australia, Canberra, 2601, Australia
Andrew Feitz

Department of Geosciences and Natural Resource Management, University of Copenhagen, Copenhagen, 1350, Denmark
Rasmus Fensholt, Birger Ulf Hansen & Torbern Tagesson

Energy Analysis & Environmental Impacts Division, Lawrence Berkeley National Laboratory, Berkeley, CA, 94720, USA
Marc Fischer

USDA Forest Service, Rocky Mountain Research Station, Fort Collins, CO, 80526, USA
John Frank & William Massman

Institute of BioEconomy, National Research Council of Italy, Firenze, 50145, Italy
Beniamino Gioli

School of Natural Resources, University of Nebraska-Lincoln, Lincoln, NE, 68583, USA
Anatoly Gitelson, Andy Suyker & Elizabeth Walter-Shea

Max Planck Institute for Biogeochemistry, Jena, 03641, Germany
Mathias Goeckede & Olaf Kolle

Department of Biology, Virginia Commonwealth University, Richmond, VA, 23284, USA
Christopher M. Gough

Department of Earth System Science, University of California, Irvine, CA, 92697, USA
Michael L. Goulden

Agrosphere (IBG3), Forschungszentrum Jülich, Jülich, 52428, Germany
Alexander Graf & Marius Schmidt

Department of Ecology, University of Innsbruck, Innsbruck, 6020, Austria
Albin Hammerle & Georg Wohlfahrt

International Joint Research Laboratory for Global Change Ecology, School of Life Sciences, Henan University, Kaifeng, 450000, China
Shijie Han

Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang, 110016, China
Shijie Han & Junhui Zhang

Department of Forest Ecosystems and Society, Oregon State University, Corvallis, OR, 97333, USA
Chad Hanson, Hyojung Kwon & Bev Law

College of Resources and Environment, University of Chinese Academy of Sciences, Beijing, 100190, China
Yongtao He

School of Ecosystem and Forest Sciences, The University of Melbourne, Creswick, VIC3363, Australia
Nina Hinko-Najera

Research Institute for the Environment and Livelihoods, Charles Darwin University, Darwin, 0909, Australia
Lindsay Hutley

Department of Environmental Engineering, Technical University of Denmark (DTU), Kongens Lyngby, 2800, Denmark
Andreas Ibrom & Kim Pilegaard

Institute for Agro-Environmental Sciences, National Agriculture and Food Research Organization, Tsukuba, 305-8604, Japan
Hiroki Ikawa

Wageningen Environmental Research, Wageningen University and Research, Wageningen, 6708PB, The Netherlands
Wilma Jans

Key Laboratory of Vegetation Ecology, Ministry of Education, Northeast Normal University, Changchun, 130024, China
Shicheng Jiang

Research Faculty of Agriculture, Hokkaido University, Sapporo, 060-8589, Japan
Tomomichi Kato

GI-Core, Hokkaido University, Sapporo, 060-0808, Japan
Tomomichi Kato

Geography and Environmental Management, Waterloo, ON, N2L3G1, Canada
Myroslava Khomik

Institute of Meteorology and Climate Research, Karlsruhe Institute of Technology, Garmisch-Partenkirchen, 82467, Germany
Janina Klatt, Hans Peter Schmid & Rainer Steinbrecher

Bioclimatology, University of Goettingen, Goettingen, 37077, Germany
Alexander Knohl, Lukas Siebicke & Frank Tiedemann

Centre of Biodiversity and Sustainable Land Use (CBL), University of Goettingen, Goettingen, 37077, Germany
Alexander Knohl

Department of Geography, The University of British Columbia, Vancouver, BC, V6T1Z2, Canada
Sara Knox

Research Institute for Global Change, Institute of Arctic Climate and Environment Research, Japan Agency for Marine-Earth Science and Technology, Yokoama, 236-0001, Japan
Hideki Kobayashi

Biological Sciences, University of Adelaide, Adelaide, SA5064, Australia
Georgia Koerber & Wayne Meyer

Graduate School of Agriculture, Kyoto University, Kyoto, 606-8502, Japan
Yoshiko Kosugi

Graduate School of Bioagricultural Sciences, Nagoya University, Nagoya, 4648601, Japan
Ayumi Kotani

Department of Applied Physics, University of Granada, Granada, 18071, Spain
Andrew Kowalski & Enrique P. Sánchez-Cañete

Water systems and Global Change group, Wageningen University, Wageningen, 6500, The Netherlands
Bart Kruijt

A.N. Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow, 119071, Russia
Julia Kurbatova, Ivan Shironya, Andrej Varlagin & Natalia Vygodskaya

Head Office, Integrated Carbon Observation System (ICOS ERIC), Helsinki, 00560, Finland
Werner L. Kutsch

Natural Resources Institute Finland, Helsinki, 00790, Finland
Samuli Launiainen

Key Laboratory of Adaptation and Evolution of Plateau Biota, Northwest Institute of Plateau Biology, Chinese Academy of Sciences, Xining, 810008, China
Yingnian Li

Centre for Tropical Environmental Sustainability Studies, James Cook University, Cairns, 4878, Australia
Michael Liddell

CEFE, CNRS, Univ Montpellier, Montpellier, 34293, France
Jean-Marc Limousin, Jean-Marc Ourcival & Serge Rambal

Forestry and Environment Division, Forest Research Institute Malaysia (FRIM), Kepong, 52109, Malaysia
Marryanna Lion

Institute for Atmosphere and Earth System Research/Physics, University of Helsinki, Helsinki, 00560, Finland
Annalea Lohila, Ivan Mammarella, Üllar Rannik & Timo Vesala

Department of Botany, School of Natural Sciences, Trinity College Dublin, Dublin, D02PN40, Ireland
Ana López-Ballesteros

German Meteorological Service (DWD), Centre for Agrometeorological Research, Braunschweig, 38116, Germany
Antje Lucas-Moffat

Micrometeorology, University of Bayreuth, Bayreuth, 95440, Germany
Johannes Lüers

Bayreuth Center of Ecology and Environmental Research, 95448, Bayreuth, Germany
Johannes Lüers

CSIRO Land and Water, Floreat, 6014, Australia
Craig Macfarlane

Department of Agriculture, University of Sassari, Sassari, 07100, Italy
Serena Marras, Costantino Sirca & Donatella Spano

Oulanka research station, University of Oulu, Kuusamo, 93900, Finland
Mikhail Mastepanov

Dept. Biological Sciences, Wellesley College, Wellesley, MA, 02481, USA
Jaclyn Hatala Matthes

Research Institute on Terrestrial Ecosystems, National Research Council of Italy, Monterotondo Scalo, 00015, Italy
Francesco Mazzenga

Department of Geography and Planning, Queen’s University, Kingston, ON, K7L3N6, Canada
Harry McCaughey

Environmental Analytics NZ, Ltd. Raumati South, Paraparaumu, 5032, New Zealand
Andrew M. S. McMillan

Mazingira Centre, International Livestock Research Institute (ILRI), Nairobi, 00100, Kenya
Lutz Merbold

NOAA/OAR/Air Resources Laboratory, 325 Broadway, Boulder, CO, 80303, USA
Tilden Meyers

Atmospheric Sciences Research Center, State University of New York at Albany, Albany, NY, 12203, USA
Scott D. Miller

Forest Department of South Tyrol, Bolzano, 39100, Italy
Stefano Minerbi & Leonardo Montagnani

Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ, 85721, USA
Russell K. Monson, Natalia Restrepo-Coupe & Scott R. Saleska

Faculty of Science and Technology, Free University of Bolzano, Bolzano, 39100, Italy
Leonardo Montagnani

Department of Plant Biology, University of Illinois at Urbana-Champaign, Urbana, IL, 61801, USA
Caitlin E. Moore

IHE Delft, Delft, 2611, The Netherlands
Eddy Moors

Faculty of Science, VU Amsterdam, Amsterdam, 1081, The Netherlands
Eddy Moors

University Grenoble Alpes, IRD, CNRS, IGE, Grenoble, 38000, France
Virginie Moreaux

School of Engineering and Applied Sciences, Harvard University, Cambridge, MA, 02138, USA
J. William Munger & Steven Wofsy

Department of Earth and Planetary Sciences, Harvard University, Cambridge, MA, 02138, USA
J. William Munger & Steven Wofsy

School of Forestry and Resource Conservation, National Taiwan University, Taipei, 0617, Taiwan
Taro Nakai

International Arctic Research Center, University of Alaska Fairbanks, Fairbanks, AK, 99775, USA
Taro Nakai

Environment and Climate, Research Institute for Nature and Forest, Geraardsbergen, 9500, Belgium
Johan Neirynck

Department of Ecosystem Science and Management, Texas A&M University, College Station, TX, 77843, USA
Asko Noormets

Research Institute for the Environment and Livelihoods, Charles Darwin University, Casuarina, 0810, Australia
Matthew Northwood

Grupo de Estudios Ambientales, Instituto de Matemática Aplicada San Luis (UNSL & CONICET), San Luis, D5700HHW, Argentina
Marcelo Nosetto

Facultad de Ciencias Agropecuarias (UNER), Oro Verde, 3100, Argentina
Marcelo Nosetto

Eco&Sols, Univ Montpellier-CIRAD-INRA-IRD-Montpellier SupAgro, Montpellier, 34060, France
Yann Nouvellon

O’Neill School of Public and Environmental Affairs, Indiana University Bloomington, Bloomington, IN, 47405, USA
Kimberly Novick

Global Change Research Group, Dept. Biology, San Diego State University, San Diego, CA, 92182, USA
Walter Oechel & Donatella Zona

Department of Geography, College of Life and Environmental Sciences, University of Exeter, Exeter, EX44RJ, United Kingdom
Walter Oechel

Department of Agroecology, Aarhus University, Tjele, 8830, Denmark
Jørgen Eivind Olesen

iCLIMATE, Aarhus University, Tjele, 8830, Denmark
Jørgen Eivind Olesen

Department of Geology, Wayne State University, Detroit, MI, 48202, USA
Shirley A. Papuga

Department of Geosciences, University of Oslo, Oslo, 0315, Norway
Frans-Jan Parmentier

Department of Geography, University of Zurich, Zurich, 8057, Switzerland
Eugenie Paul-Limoges

Department of Forest Ecology and Management, Swedish University of Agricultural Sciences, Umeå, 90183, Sweden
Matthias Peichl

Hawkesbury Institute for the Environment, Western Sydney University, Penrith, 2751, Australia
Elise Pendall & Victor Resco de Dios

Department of Biology, Indiana University Bloomington, Bloomington, IN, 47401, USA
Richard P. Phillips

Instituto de Clima y Agua, Instituto Nacional de Tecnologia Agropecuaria (INTA), Buenos Aires, 1686, Argentina
Gabriela Posse & Sebastian Westermann

CSIRO Land and Water, Wembley, 6913, Australia
Norbert Pirk & Suzanne M. Prober

Center for Global Change & Earth Observations, Michigan State University, East Lansing, MI, 48823, USA
David Reed

School of Life Science and Engineering, Southwest University of Science and Technology, Mianyang, 621010, China
Victor Resco de Dios

Departamento de Química e Física, Universidade Federal da Paraiba, Areia, PB, 58397-000, Brazil
Borja R. Reverter

Remote Sensing and Geoinformatics, GFZ German Research Centre for Geosciences, Potsdam, 14473, Germany
Torsten Sachs & Christian Wille

Andalusian Institute for Earth System Research (CEAMA-IISTA), Granada, 18006, Spain
Enrique P. Sánchez-Cañete & Penélope Serrano-Ortíz

Ciencias del Agua y Medioambiente, Instituto Tecnológico de Sonora, Ciudad Obregón, 85000, Mexico
Zulia M. Sanchez-Mejia

Geographical Institute, University of Cologne, Cologne, 50923, Germany
Karl Schneider

Department of Industry, Innovation and Science, Geoscience Australia, Canberra, 2609, Australia
Ivan Schroder

Southwest Watershed Research Center, USDA-ARS, Tucson, AZ, 85719, USA
Russell L. Scott

Institute of Atmospheric Physics of the Czech Academy of Sciences, Prague, 14100, Czech Republic
Pavel Sedlák

Department of Ecology, University of Granada, Granada, 18071, Spain
Penélope Serrano-Ortíz

National Hulunber Grassland Ecosystem Observation and Research Station & Institute of Agricultural Resources and Regional Planning, Chinese Academy of Agricultural Sciences, Beijing, 100081, China
Changliang Shao

School of Science, Edith Cowan University, Joondalup, 6027, Australia
Richard Silberstein

Sentek Pty Ltd, Stepney, SA5069, Australia
Robert M. Stevens

National Ecological Observatory Network Program, Boulder, CO, 80301, USA
Cove Sturtevant

Kansai Research Center, Forestry and Forest Products Research Institute, Kyoto, 612-0855, Japan
Satoru Takanashi

College of Urban and Environmental Sciences, Peking University, Beijing, 100871, China
Yanhong Tang

School of Earth, Atmosphere and Environment, Monash University, Clayton, 3800, Australia
Nigel Tapper

Space Science and Engineering Center, University of Wisconsin-Madison, Madison, WI, 53706, USA
Jonathan Thom

Terrasystem srl, Viterbo, 01100, Italy
Michele Tomassucci

USDA Forest Service, Rocky Mountain Research Station, Missoula, MT, 59808, USA
Shawn Urbanski

Meteorology and Air Quality group, Wageningen University, 6500, Wageningen, The Netherlands
Michiel van der Molen

Fenner School of Environment and Society, Australian National University Canberra, Canberra, ACT 2600, Australia
Eva van Gorsel

Department of Civil Engineering, Monash University, Clayton, 3800, Australia
Jeffrey P. Walker

CSIRO Land and Water, Canberra, 2601, Australia
William Woodgate

South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, 510650, China
Caroline Vincke & Yuelin Li

College of Applied Meteorology, Nanjing University of Information Science & Technology, Nanjing, 210044, China
Guoyi Zhou

Department of Animal and Plant Sciences, University of Sheffield, Sheffield, S102TN, United Kingdom
Donatella Zona

Dario Papale, Gilberto Pastorello, Margaret Torn, and Deb Agarwal conceived of and organized the FLUXNET2015 dataset. Eleonora Canfora, You-Wei Cheah, Danielle Christianson, Housen Chu, Dario Papale, Gilberto Pastorello, Diego Polidori, Cristina Poindexter, and Carlo Trotta were responsible for quality checking, post-processing, and creating data products. You-Wei Cheah, Danielle Christianson, Abdelrahman Elbashandy, Marty Humphrey, Gilberto Pastorello, Diego Polidori, Alessio Ribeca, Carlo Trotta, and Catharine van Ingen were responsible for code and software preparation and implementation (processing pipeline and data distribution platform). Eleonora Canfora, You-Wei Cheah, Jiquan Chen, Danielle Christianson, Housen Chu, Peter Isaac, Dario Papale and Leiming Zhang managed the flux data and metadata collections. Gilberto Pastorello, Dario Papale, Deb Agarwal, Sebastien Biraud, You-Wei Cheah, Danielle Christianson, Housen Chu, and Margaret Torn conceived of the paper and prepared the first draft that was reviewed and commented on by all the coauthors. All the coauthors collected, processed and contributed the data, also participated in the quality assessment and correction of errors. The link between sites and coauthors is provided in Supplementary Table SM9. More

The design and basic descriptive epidemiology of the IDEAL project is fully described in Bronsvoort et al.3. To give the IDEAL database a context, we briefly describe the study in relation to the samples taken.
The study population
Between 2007–2009, 548 free-grazing indigenous East African Shorthorn Zebu calves in Western Kenya were recruited. The project was based in Busia, a town in the west of Kenya on the border with Uganda. The town had a veterinary lab that was able to be used and developed by the project.
Calves were recruited using a stratified two-stage random cluster study design (Fig. 1). In the first stage, a weighted random sample of 20 sublocations was selected from across the four agro-ecological zones represented in the project area. This area was roughly 45 × 90 km2. This area was chosen because each area would be drivable from Busia and back to the lab and all visits completed within a day.
In the second stage, approximately 28 3–7 day-old calves were randomly recruited from each sublocation. The sublocations were visited on a rolling 5-week cycle to ensure there was an even distribution of calves recruited across space and season over the study period. Only one calf per dam was recruited and a farmer could only have one calf at a time in the study. Recruited calves were followed for their first year (51 weeks) of life. A calf was selected at random from all those available and eligible on the day of the recruitment visit.
As described in Bronsvoort et al.3, each calf had to meet the following inclusion criteria to be enrolled in the study: (1) it was between 3 and 7 days old at recruitment; (2) it was not born as a result of artificial insemination of the dam; (3) the dam was not managed under zero-grazing conditions (as this is likely to reflect cattle associated with dairy production and potential exotic genetics and thus not be representative of the traditional small holder farming system); (4) it did not have any congenital deformities (the project was interested in infectious causes of disease and the association with host genetics, rather than direct genetic disease). Recruitment was conditional on the farmer allowing access to the calf, willingness to report clinical episodes to the project team, and agreeing not to self-treat their calves. Farmers were compensated at a rate agreed with the staff of the District Veterinary Office; this comprised the estimated cost of raising the calf for one year as calves were nominally owned by the project for that year. Owners were free to refuse to participate3 and consent was obtained in a language in which the participant was confident.
There were two periods during the IDEAL project when sampling and recruitment was suspended. In the first, field work was suspended for 6 weeks in 2008 due to political unrest. This resulted in a small number of calves missing one or two 5 weekly visits. The second was due to an extended holiday period that resulted in staffing problems in 2009/2010.
Data collection
Below, we briefly describe how the IDEAL data was collected. See Bronsvoort et al.3 for more details.
Routine clinical examination of calves
At the recruitment visit, a household questionnaire was completed by interview with the calf owner or head of the household. This questionnaire collected information about the farmer and the farm, such as the type of livestock kept, and animal management practices. Calves were raised according to the farmer’s practices.
Calves received a routine systematic physical examination from a veterinary surgeon at the recruitment visit and these examinations where repeated every 5 weeks until the calf was 51 weeks old. Some routine measurements were taken during the exam (body weight and lymph node width). All abnormalities were noted and all areas were noted as checked. Ectoparasites found were identified. During each visit a standard set of biological samples were taken for further laboratory analysis. In addition, a questionnaire was carried out at each visit to update the IDEAL project about other activities on the farm such as illness or treatment of the other livestock and animal movements. At the final visit, physical phenotype was recorded following a standardised format. The study design, sampling procedure, clinical examinations and sample collection are summarised in Figs. 2 and 3, and can also be read in detail in the supporting documentation online (http://data.ctlgh.org/ideal/).
Clinical episodes and post-mortem examinations of calves
If a calf became unwell, the farmer was asked to contact the IDEAL project veterinary surgeon. The calf was visited and a full clinical examination and history was taken. Both routine and additional appropriate specific samples were collected based on the syndrome observed3. If a calf was deemed to be suffering and if that suffering would be alleviated with treatment, that treatment was given and the calf was no longer visited and was censored from the study from that point. If calves were suffering and that could only be alleviated by euthanasia, this was carried out and the calf was examined post-mortem following the standard protocol. Following the death or euthanasia of a calf, a full gross post-mortem examination was carried out using standard veterinary procedures and clinically appropriate samples collected for further analysis. Cause of death was attributed using a panel of experts with access to all available diagnostic results and the necropsy report. This was carried out for all calves that died at a single timepoint after the close of the project.
Examination of the dams
A limited clinical examination of the dam was performed at recruitment and in subsequent routine visits until the calf was weaned. At these visits the girth was measured and body condition was scored and the udder was examined for evidence of lesion or mastitis that could affect calf nutritional intake. Phenotypic measurements of coat colour and pattern, horn length and shape, ear shape, size of hump and dewlap was recorded at recruitment3.
Laboratory analysis
Blood samples collected into EDTA tubes were used for differential blood cell counts, performed using the pocH-100iV Diff (Sysmex®, Europe GMBH). The haematological parameters investigated are listed in Online-only Table 1. In addition to the automated blood analysis, EDTA unclotted samples were used to make thin blood smears for manual differential cell counts. These smears were transported to the University of Pretoria, South Africa, where blood smears were stained with Diff Quick (Kyron, South Africa) for differential counts. Packed cell volume (PCV) was measured manually using a Hawksley micro-hematocrit reader4. Total serum protein (TSP) was measured from 100 μL serum using a refractometer (model RHC-200ATC, Westover Scientific).
Peripheral ear vein blood smears were collected and examined for haemoparasites by microscopy. Thin smears were fixed using methanol and stained using Giemsa. Thick smears were directly stained. One hundred fields were examined under an oil immersion lens. Haemoparasites present were identified to genus level.
Reverse line blot (RLB) hybridization assay was performed as previously described5 to detect tick-borne haemoparasites in the blood (Theileria, Anaplasma, Ehrlichia, and Babesia (Online-only Table 1))5.
The p104 nested PCR was carried out on calves where ECF was suspected on clinical grounds to specifically identify T. parva6.
Whole blood collected in EDTA was mixed in sodium EDTA tubes in a 1:1 ratio with ‘magic buffer’ (which acted as an anti-coagulant, anti-fungus, anti-bacterial and preservative; Biogen Diagnostica, Villaviciosa De Odon, Spain) at the recruitment visit in readiness for genomic analysis. DNA was extracted from these samples using the Nucleon Genomic DNA extraction kit (TepnelnLife Sciences, Manchester, UK). The Illumina® BovineSNP50 v. 1 BeadChip (Illumina Inc., San Diego, CA, USA) used to genotype the cattle. Genotyping of the 548 calves was carried out at the USDA-ARS bovine functional (Beltsville, MD, USA) and GeneSeek (https://genomics.neogen.com) laboratories using the genome assembly v3.0. In addition, due to the cost of sequencing at the time, a subset of 114 cattle were genotyped using the Illumina® BovineHD Genotyping BeadChip.
Faecal samples were also collected during the study and these where routinely screened using the standard Baermann and McMasters protocols7. The number of strongyle eggs per gram of faeces was evaluated using the McMasters counting technique. These could be read to the nearest 50 eggs per gram. Sedimentation was carried out for detection of fluke eggs and larval cultures were used to speciate strongyle eggs7. Species where reported at the highest level; strongyle eggs/strongyloides/coccidia/nematodirus. See Online-only Table 1 for more details on the parasites detected.
Serum samples were collected from blood collected into plain vacutainer tubes. These were stored in duplicate for serological analyses. Species-specific antibody response enzyme-linked immunosorbent assays (ELISAs) were performed and analysed according to the manufacturer’s instructions.
The full list of pathogens and viruses for which the cattle have so far been screened can be found in Online-only Table 1. The prevalence of pathogens currently identified across the whole study period and at each visit is presented in Figs. 4 and 5.
Since the database is linked to a biobank, the list of pathogens tested and screened for is continuously updated as new tests are performed or new tools are developed. More

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