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NITROGEN CYCLING IN THE AMERICAS:NATURAL AND ANTHROPOGENIC INFLUENCES AND CONTROLS NITROGEN CYCLING IN THE AMERICAS: NATURAL AND ANTHROPOGENIC INFLUENCES AND CONTROLS Edited by LUIZ A. MARTINELLI Cena-University of São Paulo, Brazil & ROBERT W. HOWARTH Cornell University, U.S.A. Reprinted fromBiogeochemistry Volume 79, Nos. 1–2, 2006 123 A C.I.P. catalogue record for this book is available from the Library of Congress. ISBN 1-4020-4717-7 Published by Kluwer Academic Publishers, P.O. Box 17, 3300 AADordrecht, The Netherlands. Sold and distributed in the U.S.A. and Canada by Kluwer Academic Publishers 101 Philip Drive, Norwell, MA02061, U.S.A. In all other countries, sold and distributed by Kluwer Academic Publishers, P.O. Box 322, 3300 AH Dordrecht, The Netherlands. Printed on acid-free paper. All Rights Reserved © 2006 Springer No part of the material protected by this copyright notice may be reproduced or utlized in any form or by any means, electronic or mechanical, including photo- copying, recording or by any information storage and retrieval system, without written permission from the copyright owner. Printed in the Netherlands. Table of Contents LUIZ A. MARTINELLI & ROBERT W. HOWARTH / Preface 1–2 LUIZ A. MARTINELLI, ROBERT W. HOWARTH, ELVIRA CUEVAS, SOLANGE FILOSO, AMY T. AUSTIN, LORETA DONOSO, VERA HUSZAR, DENNIS KEENEY, LUCIENE L. LARA, CARLOS LLERENA, GEORGE McISSAC, ERNESTO MEDINA, JORGE ORTIZ-ZAYAS, DONALD SCAVIA, DAVID W. SCHINDLER, DORIS SOTO & ALAN TOWNSEND / Sources of reactive nitrogen affecting ecosystems in Latin America and the Caribbean: current trends and future perspectives 3–24 DAVID W. SCHINDLER, PETER J. DILLON & HANS SCHREIER / A review of anthropogenic sources of nitrogen and their effects on Canadian aquatic ecosystems 25–44 AMY T. AUSTIN, GERVASIO PIÑEIRO & MARINA GONZALEZ-POLO / More is less: agricultural impacts on the N cycle in Argentina 45–60 SOLANGE FILOSO, LUIZ ANTONIO MARTINELLI, ROBERT W. HOWARTH, ELIZABETH W. BOYER & FRANK DENTENER / Human activities changing the nitrogen cycle in Brazil 61–89 JULIO A. BAISRE / Assessment of nitrogen flows into the Cuban landscape 91–108 JORGE R. ORTIZ-ZAYAS, ELVIRA CUEVAS, OLGA L. MAYOL-BRACERO, LORETO DONOSO, IVONNE TREBS, DEBORA FIGUEROA-NIEVES & WILLIAM H. MCDOWELL / Urban influences on the nitrogen cycle in Puerto Rico 109–133 MERCY J. BORBOR-CORDOVA, ELIZABETH W. BOYER, WILLIAM H. MCDOWELL & CHARLES A. HALL / Nitrogen and phosphorus budgets for a tropical watershed impacted by agricultural land use: Guayas, Ecuador 135–161 R.W. HOWARTH, D.P. SWANEY, E.W. BOYER, R. MARINO, N. JAWORSKI & C. GOODALE / The influence of climate on average nitrogen export from large watersheds in the Northeastern United States 163–186 DONALD SCAVIA & SUZANNE B. BRICKER / Coastal eutrophication assessment in the United States 187–208 M.M.C. BUSTAMANTE, E. MEDINA, G.P. ASNER, G.B. NARDOTO & D.C. GARCIA-MONTIEL / Nitrogen cycling in tropical and temperate savannas 209–237 VERA L.M. HUSZAR, NINA F. CARACO, FABIO ROLAND & JONATHAN COLE / Nutrient–chlorophyll relationships in tropical–subtropical lakes: do temperate models fit? 239–250 JEAN P. H. B. OMETTO, JAMES R. EHLERINGER, TOMAS F. DOMINGUES, JOSEPH A. BERRY, FRANÇOISE Y. ISHIDA, EDMAR MAZZI, NIRO HIGUCHI, LAWRENCE B. FLANAGAN, GABRIELA B. NARDOTO & LUIZ A. MARTINELLI / The stable carbon and nitrogen isotopic composition of vegetation in tropical forests of the Amazon Basin, Brazil 251–274 Biogeochemistry(2006)79:1–2 (cid:2)Springer2006 DOI10.1007/s10533-006-9012-z Preface This special issue presents the products of three workshops designed to compare differences in nitrogen cycling between tropical and temperate eco- systems across the Americas, as well as differences in how human societies shave transformed the nitrogen cycle according to economic, social and cul- tural characteristics of the diverse countries of the Americas. The first work- shopon‘‘Nfluxesandprocessesintropicalandtemperatesystems’’washeldin the city of Ubatuba, State of Sa˜o Paulo, Brazil, in March 2003. From this workshop,astrong consensus developedtoforman ongoing‘‘InterAmerican NitrogenNetwork.’’Towardsthisend,twomoreworkshopswereheld,onein San Juan, Puerto Rico in May 2004, and one in May of 2005 in the city of Brası´lia,Brazil. Thisefforttobuild theInterAmericanNitrogenNetwork isa joint effort of the Latin American Nitrogen Center and the North American Nitrogen Center, two of the five continental-scale regional centers of the International Nitrogen Initiative (INI) created in 2003 by SCOPE and the IGBP. The INI’s main objective is ‘‘to optimize nitrogen’s beneficial role in sustainable food production and minimize nitrogen’s negative effects on human health and the environment resulting from food and energy produc- tion.’’WehopethereaderswillagreewithusthatthisissueofBiogeochemistry isanexcellentstartfortheInterAmericanNitrogenNetworkandtheeffortsof the INI in the Americas. ThefirstpaperpresentedinthisissuebyMartinelliandotherparticipantsof the San Juan workshop is an overview of the nitrogen sources across the Americas and how they vary from one region to another. This paper is an introductory paper for a series of six following papers that detailed the nitrogen cycle in different countries or in important sub-regions of these countries. Among these papers, Schindler et al. discuss N-deposition related issues in Canada, followed by Austin et al. who present a nitrogen budget for Argentina, one of the most important regions of cereal production in the world. Filoso et al. for Brazil, Baisre for Cuba and Ortiz for Puerto Rico synthesize quantitative nitrogen assessments for these whole countries, while Borbor et al. restrict their analyses for the most developed watershed in Equador. After these country-scale assessments, Howarth et al. and Scavia & Bricker illustrate aspects of the nitrogen cycle in important regions of the USA. Howarth et al. illustrate how climate and human activity interact to regulatethenitrogenexportfromthemajor16watershedsoftheNortheastern USAregion,oneofthemostpollutedregionsofthecountry.Scavia&Bricker present a comprehensive approach to problems that the coastal regions of the USA are facing due to the large amount of nitrogen that these key areas receive. The last three papers in the issue are mostly a product of the first 2 workshop held in Ubatuba and deal more with nitrogen as a key element in natural systems. Bustamante et al. present nitrogen as one of the most important limiting element in savanna ecosystems throughout the world. Huszar et al. analyze nitrogen limitation in tropical lakes in comparison to temperate lakes. Finally, Ometto et al. present a detailed description of the variability of nitrogen and carbon stable isotopes in tropical forests of the Amazon region. We believe this volume represents a significant advancement inourabilitytounderstandnaturalprocessesandhumanperturbationsofthe nitrogen cycle in both temperate and tropical systems, as well as to recognize several major differences in nitrogen cycling across several countries and regions of the Americas. We are in debt with the many institutions and people who helped us to organize our workshops and prepare the manuscripts. Elvira Cuevas and MercedesBustamanteandtheirstaffswerewonderfulhostessesinSanJuanand Brası´lia, respectively. The Ubatuba workshop had financial support from the Fundac¸a˜o de Amparo a` Pesquisa do Estado de Sa˜o Paulo (FAPESP) and the EssoBrasileiradePetro´leocompany.TheSanJuanworkshopwassupportedby theInterAmericanInstituteforGlobalChanges(IAI)andbytheUniversityof PuertoRicothroughagrantfromtheUSNationalScienceFoundationtoCrest (Center for Applied Tropical Ecology and Conservation). The Brası´lia work- shop was mainly supported by the Conselho Nacional de Desenvolvimento ePesquisas(CNPq),throughitsPro-Sulprogram,andalsobytheIAIandthe University of Brası´lia. We also thank Suzanne Mekking, the publisher of Biogeochemistry, forher flexibility and support inpreparing this issue. Luiz A. Martinelli CENA, Laboratory of Isotope Ecology University of Sa˜o Paulo Piracicaba, SP, Brazil e-mail: [email protected] Robert W. Howarth Department of Ecology Evolutionary Biology Cornell University Ithaca, USA Biogeochemistry(2006)79:3–24 (cid:2)Springer2006 DOI10.1007/s10533-006-9000-3 Sources of reactive nitrogen affecting ecosystems in Latin America and the Caribbean: current trends and future perspectives LUIZ A. MARTINELLI1,*, ROBERT W. HOWARTH2, ELVIRA CUEVAS3, SOLANGE FILOSO2, AMY T. AUSTIN4, LORETA DONOSO5, VERA HUSZAR6, DENNIS KEENEY7,8, LUCIENE L. LARA1, CARLOS LLERENA9, GEORGE MCISSAC10, ERNESTO MEDINA11, JORGE ORTIZ-ZAYAS12, DONALD SCAVIA13, DAVID W. SCHINDLER14, DORIS SOTO15 and ALAN TOWNSEND16 1CENA,Av.Centena´rio303,13416-000,Piracicaba,SP,Brazil;2Ecology&EnvironmentalBiology Department,CornellUniversity,E311CorsonHall,Cornell,Ithaca,NY14853,USA;3Departmentof Biology,CollegeofNaturalSciences,UniversityofPuertoRico,POBox23360SanJuan,PuertoRico 00931-3360;4FacultyofAgronomy,UniversityofBuenosAiresandIFEVA-CONICET,AvenidaSan Martı´n 4453, C1417DSE, Buenos Aires, Argentina; 5Instituto Venezuelano de Investigaciones Cientı´ficas – IVIC, Centro de Ecologı´a, IVIC, Aptdo. 21827, Caracas 1020-A, Venezuela; 6Departamento de Botaˆnica, Universidade Federal do Rio de Janeiro, Museo Nacional, Quinta da BoaVistas/n,Sa˜oCristo´va˜o–RiodeJaneiro,RJ20940040,Brasil;7AgricultureandBiosystems Engineering,IowaStateUniversity,3402EisenhowerAve.,Ames,Iowa50010,USA;8Institutefor Agriculture and Trade Policy, Minneapolis, Minnesota, USA9Universidad Nacional Agraria La Molina,Apartado456,Lima,Peru;10DepartmentofNaturalResourcesandEnvironmentalSciences, UniversityofIllinoisatUrbanaChampaign,w-503TurnerHall,1102SouthGoodwinAvenue,Urbana, IL61801,USA;11InstitutoVenezolanodeInvestigacionesCientı´ficas,LaboratoryPlantPhysiological Ecology, Centro de Ecologı´a, IVIC, Aptdo. 21827, Caracas, 1020-A, Venezuela; 12Institute for TropicalEcosystemsStudies,UniversityofPuertoRico,POBox23341SanJuan,PuertoRico00931- 3341;13SchoolofNaturalResourcesandEnvironment,UniversityofMichigan,G520Dana1115,Ann Arbor,MI,48109,USA;14EcologyDepartment,UniversityofAlberta,Z811,BiologicalSciences Bldg.,114St–89Ave,Edmonton,Alberta,T6G2E1,Canada;15UniversidadAustraldeChile,Casilla 1327,PuertoMontt,Chile;16INSTAARandEPO-Biology,UniversityofColorado,156030thSt., Boulder,CO80309,USA;*Authorforcorrespondence(e-mail:[email protected];phone:+55- 19-3429-4074;fax:+55-19-3434-9210) Keywords: Biomassburning,Caribbean,LatinAmerica,Nitrogen,Reactiveagriculture,Urban- ization Abstract. While the amount of reactive nitrogen circulating at the global level has increased markedlyinthelastcentury,theeffectsofthisincreasearelargelyseenattheregionalleveldueto interactingecologicalandsocio-economicfactors.Incontrastwithmostotherregionsoftheworld, LatinAmericaandtheCaribbean(LA-Ca)standoutduetothefactthatthemajorinputofreactive nitrogen(Nr)stilloccursnaturallyviabiologicalnitrogenfixation(BNF)innaturalecosystemsas opposedtoanthropogenicinputsofsyntheticfertilizer,fossilfuelcombustionandcroppingwith leguminousspecies.Largelyduetoeconomicreasons,theconsumptionoffertilizerNintheLA-Ca regionisstilllowincomparisonwiththeaverageconsumptionoftheworld.However,thefertilizer Nconsumptionisincreasingatamuchfasterratethanthatindevelopedregionsoftheworld,like 4 USAandCanada.TheNrproductionthroughBNFincultivatedplantsthatfixnitrogen(C-BNF)is 5timeslowerthanthatoccurringnaturallyinLatinAmerica,butisstillequivalentto16%ofthe worldC-BNF.Thecultivationofnitrogen-fixingcropspeciesintheLA-Caregionisalsoincreasing, almostentirelyduetotheexpansionofsoybeanfieldsinthecentralandnorthernregionsofBrazil andthePamparegionofArgentina.Otheranthropogenicactivitiesintheregionthatcontributeto an increase in the circulation of reactive nitrogen include the impact of biomass burning and urbanization.Inthelastdecade,anaverageof47,000km2peryearofforestswasburnedintheLA- Ca region. The environmental impact of urban centers in the LA-Ca region has become very important,sinceanintenseurbanizationprocessisoccurringinthisregion,atanintensitythatfar exceedsurbandevelopmentinthenorthernhemisphere.Theconsequencesofincreasedurbanization includeincreasedemissionsofNO totheatmosphereduetothefossilfuelcombustion,andthelack x ofsewagetreatmentfacilitiesinmostcitiesoftheLA-Caresultinalargevolumeofuntreatedsewage dischargedintosurfacewaters,creatingseriousenvironmentalproblems.Thecombinationofrapid urbanizationandagriculturalintensificationinthisregionsuggestthatconcerniswarrantedforthe potentialforincreaseinthecirculationofreactivenitrogenintheverynearfuture.Atthesametime, theopportunitystillexiststomitigatesomeoftheconsequencesofhumanimpactonthenitrogen cycleinaregionthatstillmaintainsalargefractionofitsnaturalecosystemsintact. Introduction The concentration of reactive nitrogen (Nr) on the Earth has dramatically increasedinthelastcenturyduemainlytotheproductionofsyntheticnitrogen fertilizer through the Haber–Bosch process, fossil fuel combustion, and the cultivation of plants that associate with bacteria that fix atmospheric N (Ho- warthet al.2002;Gallowayet al.2004).Ithasbeenestimatedthatin1860the productionofanthropogenicNrwas15 TgN yr)1,andthatbytheearly1990’s it increased by almost 10 times to approximately 140 Tg N yr)1 (Galloway et al. 2004). Although these estimates are important in providing a global pictureofthealterationoftheNcyclebyhumanactivity,theydonotallowus to propose actions that would help mitigate the deleterious effects of N in the environment and for human health occurring at regional or local scale. Much moredetailedspatialinformationontheproductionandfateofNrisneededto propose such actions. Galloway et al. (2004) investigated geopolitical N budgets and showed that Latin America stands out as the region with highest inputs of Nr by naturally occurring biological nitrogen fixation (BNF) – an amount equivalent to 25% oftheworld’sNrcreatedinterrestrialecosystems.TheNrproductionthrough BNF in cultivated plants that fix nitrogen (C-BNF) in 1995 was 5 times lower thanthatoccurringnaturallyinLatinAmerica,butwasstillequivalentto16% of the world C-BNF. On the other hand, Latin America was responsible in 1995foronly6%oftheworld’sNrcreatedthroughfossilfuelcombustion,and for only 4% of the Nr originated from synthetic N fertilizer (Galloway et al. 2004). In this paper, we provide an evaluation of the origin of reactive nitrogen in Latin America, and compare it to other areas of the world. The contrasts between impacts at the global and regional scale may help to suggest ways in 5 whichwecanmitigatethedeleteriouseffectsofhumanimpactontheNcyclein thecomingdecades.Assessingnotonlythecurrentstateofhumanimpact,but the velocity of change in this region highlights the importance of focusing on regionswherethepotentialhumanimpactinthecomingdecadeswillbelarge, and where preventative steps could be taken to minimize the negative impacts of human-induced changes in nitrogen cycling. The geography and socio-economic aspects of the Latin America and the Caribbean (LA-Ca) regions Geography Becausestatisticsaregenerallycategorizedbycountryorpoliticalunitinmost data bases, we define Latin America and the Caribbean, including their countries and territories, as shown in Table 1 (http://www.cia.gov/cia/publi- cations/factbook/geos/cj.html). The most typical definition of Latin America is the one that considers the countries of South America, Central America, the Caribbean, and southern North America that speak a Romance language, including Spanish, Portu- guese, and French (Table 1). The Caribbean is defined as a group of islands locatedintheCaribbeanSea,organizedin25distinctpoliticalunits,including statesandterritories(Table 1).Geographicaloreconomicalandsocialindexes are available for most independent countries and, in some cases, for political units as well. For example, Puerto Rico is a commonwealth of the United States (US), and as a result, its data are sometimes aggregated to that the US statistics, while in other cases, they are reported as an independent political unit.AsmostproductionanduseofNrislinked toeconomicindicators,such as use of fossil fuel, fertilizer use, and cultivation of commercially important plants that have N-fixing symbioses, it is useful to provide Latin American economic and social indicators for context. Social and economical indicators The Gross Domestic Product (GDP) index (Human Development Report 2003), (defined as the total value of goods and services produced by a nation and expressed on a per capita basis) varied from 0.48 to 0.83 for LA-Ca countries in 2003 (Figure 1). Approximately 40% of the LA-Ca countries had a GDP index higher than theoverallaveragefordevelopingcountries(0.70).BoliviaandNicaraguahad indexesonlyslightlyhigherthan0.5,whileHaitiwastheonlycountrywithan indexlowerthantheaverageoftheleastdevelopingcountries(0.5).(Figure 1). The highest indexes were observed in the Bahamas and Barbados. However, bothvaluesweresignificantlylowerthantheaverageoftherichestcountriesof

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