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Coastal and Estuarine Studies Series Editors: Malcolm J. Bowman Christopher N.K. Coastal and Estuarine Studies 55 ThomasC . Malone,A lenkaM alej, LawrenceW . HardingJ, r., NenadS modlaka, and R. EugeneT urner( Eds.) Ecosystems at the Land-Sea Margin Drainage Basin to Coastal Sea American GeophysicalU nion Washington, Series Editors Malcolm J. Bowman Marine Sciences Research Center State Universityo f New York Stony Brook, NY 11794, USA Christopher N. K. Mooers Divisiono f AppliedM arine Physics RSMAS Universityo f Miami 4600 RickenbackerC swy. Miami, FL 33149-1098, USA Editors Thomas C. Malone Nenad Smodlaka Horn Point EnvironmentalL aboratory Director, Ruder Bo(cid:127),kovie Institute Universityo f Maryland Centerf or Marine ResearchR ovinj Cambridge, MD 21613 IRB, Zagreb G. Paliaga 5 Alenka Malej 52210 Rovinj National Instituteo f Biology Croatia Marine Station Piran Forance 41 R. Eugene Turner 66330 Piran Departmento f Oceanography Slovenia LouisianaS tate University Baton Rouge, LA 70803, USA Lawrence W. Harding, Jr. Horn Point EnvironmentalL aboratory University of Maryland Cambridge, MD 21613 Library of Congress Cataloging-in-Publication Data Ecosystems at the land-sea margin: drainage basin to coastal sea / Thomas C. Malone...[et al.], (eds.) p. cm -- (Coastal and estuarine studies, ISSN 0733-9569; 55) Includes bibliographicarl eferences. ISBN 0-87590-269-3 1. Coastal ecology. 2. Estuarinee cology. 3. Nutrientc ycles. I. Malone, Thomas C. II. Series QH541.5.C65E26 1999 577.5'1--dc21 98-49007 ClP ISSN 0733-9569 ISBN 0-87590-269-3 Copyright1 999 by the AmericanG eophysicalU nion, 2000 FloridaA ve., NW, Washington,D C 20009, USA. Figures,t ables, and shorte xcerptsm ay be reprintedi n scientificb ooksa nd journalsi f the source is properly cited. Authorizationto photocopyi tems for internalo r personalu se, or the internalo r personalu se of specific clients, is granted by the American Geophysical Union for librariesa nd other users registeredw ith the CopyrightC learanceC enter (CCC) TransactionaRl eportingS ervice, provided that the base fee of $1.50 per copy plus$ 0.35 per page is paidd irectlyt o CCC, 222 RosewoodD r., Danvers, MA 01923. 0733-9569/99/$01.50+0.35. This consent does not extend to other kinds of copying,s uch as copyingf or creating new collectivew orks or for resale. The reproductiono f mutiple copies and the use of full articles or the use of extracts, includingf igures and tables, for commercialp urposesr equiresp ermissionf rom AGU. Printed in the United States of CONTENTS Preface Thomas C. Malone ......................................... ix Introduction Nenad Smodlaka,T homasC . Malone,/tlenka Malej, and Lawrence(cid:127) . Harding, dr. ................................. I . The ChesapeakeB ay and Northern Adriatic Sea Drainage Basins: Land-Cover and Nutrient Export Steven1 4/S. eagle,R omanoP agnotta,a nd Ford A. Cross ............ 7 . Shallow Water and Shoreline Ecosystemso f the ChesapeakeB ay Compared to the Northern Adriatic Sea: Transformation of Habitat at the Land-Sea Margin ,]. Court Stevenson,(cid:127) lanez 1. Marusi(cid:127) Bartolo Ozreti(cid:127) Anna Marson, Giovanni Cecconii,a nd Michael S. Kearney ..................... 29 The Inland Sea: Circulation of ChesapeakeB ay and the Northern Adriatic W. C. Boicourt,M . Kuzmi(cid:127) and T. S. Hopkins. ................... 81 Production and Fate of Phytoplankton:A nnual Cyclesa nd Interannual Variability LawrenceW . Harding,(cid:127) lr., Danilo Degobbis, and Robert Precali ........................................ 131 . Dinofiagellate Bloomsa nd Mucilage Production Kevin G. Sellnet and Serena Fonda-Umani ..................... 173 Distributions and Trophic Implicationso f Microzooplankton D. WayneC oatsa nd Noelia Revelante. ........................ 207 Potential Links of Jellyfish to Eutrophication and Fisheries JenniferE . Purcell,A lenkaM alej, and/tdam Benovid. ............ 241 Macrobenthic Communities and Eutrophication Roger1 . E. Newella ndd OrgA . Ott ............................ 265 10. Pelagic-BenthicC ouplinga nd Nutrient Cycling W. M. Kemp,(cid:127) I. Faganeli,S . PuskaricE, . M. Smith, and 14/R. . Boynton ........................................ 295 11. Fisheries' Trends in Catches, Abundance, and Management E. D. Houde, S. (cid:127)lukid-Peladi(cid:127) S. B. Brandt, and S. D. Leach ....... 341 12. Summary and Conclusions AlenkaM alej, LawrenceW . Harding,J r., NenadS modlaka, and Thomas C. Malone ..................................... 367 List of Contributors ...................................................... iSb ooiks d edicattoeN do elRiae velaan pteio, neienrt hee coloogfy icrozoopad laendkicwtoaiafnteen,m dd otahneadrni, n spirtoaut sio n PREFACE Nutrient and contaminanitn putst o estuariesa nd coastals eas,t he exploitationo f living resources,t ranslocationo f nonindigenouss pecies, and habitat loss or modificationa rea mongt hem osts ignificanat nds ustaineadn thropogeniacl terationso f coastael cosystemsA.l thought he chaptersth atf ollow toucho n all of thesei ssuest,h e causesa ndc onsequenceosf nutriente nrichmenat re emphasizedN. utriente nrichment of estuariesa nd coastals eash as increasedd ramaticallyin recentd ecadesl,a rgelya s a consequencoe f the combustiono f fossil fuels and land-usep atternsi n coastal watershedsr elated to increasesin populationd ensity and agriculture.H ow these changesa nd the compoundinge ffectso f meteorologicael ventsa nd local-regional expressionosf globalc limatec hangew ill play out in termso f ecosystemdy namicsa re importantq uestiontsh at will driver esearcha nd monitoringin the coastazl onef or decades to come. Thish istoryo f thisv olumeb eginsw ith a workshopo rganizedin 1995t o conduct a comparativea nalysiso f ecosystemosf the land-seam arginf rom drainageb asint o coastals ea. The goals of the workshop", Trendsi n Land-Use,W ater Quality and FisheriesA: Comparisoonf theN orthernA driaticS eaa ndt heC hesapeakBea y",w ere to assesso ur currentu nderstandinogf nutrienti nputsa nd their effects;t o identify prioritiesf or future researcha nd monitoringt;o facilitatet he developmenot f cooperativree searchp rogramasn dt het ransfeor f informationa ndt echnologya;n dt o establishc loserl inkagesb etweene nvironmentaslc iencea ndm anagement. Thew orkshople dt o a collaboratioanm ongs cientistfsr omA ustria,C roatia,I taly, Sloveniaa ndt he United Statest o write this book. In the processt,h reeq uestionsw ere addressed(1:) How arel and-uspe atternrse flectedin thep athwayasn dr ateso f nitrogen andp hosphoruesx portf rom landt o water?( 2) How dot hep hysicacl haracteristicosf coastael cosystemtsh,e irs ize,s hapea ndc irculationre gimesi,n fluenceth e effectso f nutrienet nrichment(?3 ) To what extentc anc hangesin fisheriesb e relatedt o changes in waterq uality?A lthoughc omparisoonf then orthernA driatica ndC hesapeakBea y hasy ieldedin terestinign sighttso theseq uestionsit, h asa lsoi llustratedth e inadequacy of currenkt nowledgea sa basisfo r understandinagn dp redictingth ee ffectso f land-use practiceos n coastael cosystemasn dt her esourcetsh eys upportW. e believet hatt his volume will stimulater esearcha nd monitoringp rogramsw hich will enable us to understantdh e effectso f land-usep racticeosn coastael stuarinaen dm arinee cosystems. Thisu nderstandinwgi ll enablet he formulationa ndi mplementatioonf moree ffective policiesfo r integratemd anagemeonft thee nvironmenatn dn aturarle sources. Thee ditorsg ratefullya cknowledgteh es ponsorshoipf thew orkshopb y the U.S.- Croatiana ndU .S.-SlovenianJ ointB oardsw ith fundingf rom the Republico f Croatia, StateD irectoratefo r EnvironmentM; inistryf or Environmentaal ndP hysicalP lanning of theR epublico f Sloveniaa;n db y theU .S.N ationaOl ceanographaincd A tmospheric Administration,N ational ScienceF oundation,L and-Margin EcosystemR esearch Programa,n dE nvironmentaPlr otectioAn gencyM, ultiscaleE xperimentaElc osystem ResearchC entero f the Universityo f MarylandC enterf or EnvironmentaSl cience. Thomas C. Malone Introduction NenadS modlakaT, homasC . Malone,A lenkaM alej and LawrenceW . Harding,J r. The coastalz one from drainageb asin to coastalo cean is a mosaico f complex, interactinge cosystemtsh at includeu plandt errestrial,w etland,f reshwatere, stuarinea nd marineh abitatsI.t is a regiont hatp rovidesa diversityo f goodsa nds ervicesfr om desirable environmentfso r peoplet o live andr ecreatet,o portsa ndw aterwaysfo r shippinga ndn atural resourcefso r food andf uel. Preciselyfo r theser easonsh, umanp opulationsa re becoming increasinglyc oncentrateadl ongc ontinentaml arginsw herep rojectionsin dicatet hat over 75% of the world'sp opulationw ill live by the year2 020. A recenta nalysiso f the valueo f ecosystemse rvicess uggesttsh atc oastawl atersc, omprisingo nly 6% of thew orld'ss urface area,a ccounfto r nearly4 0% of goodsa nd services[C ostanzae t al., 1997]. Althought his analysisis controversiali,t underscoreasn importantr eality: as humanp opulationsa nd activitiesin creasein coastawl atershedsth, ec ombinede ffectso f globalc limatec hangea nd humana lterationso f the environmenat re likely to be especiallyp ronouncedin coastal aquatice cosystemwsh ereg oodsa nds ervicesa rec oncentrateadn di nputso f materialsa nd energyf rom land,s ea,a ir, andp eoplec onverge[ Malonea ndN emazie,1 996]. Nutrient enrichment[ Cole et al., 1993; Vitousek et al. 1997], anoxice vents[ Rabalais et al., 1991; Malone, 1992; Justide t al., 1993], harmful algal blooms [Smayda, 1990; Hallegraeff,1 993], fish kills [Granelie t al., 1989], habitatl oss [Mitch et al., 1994], the exploitationo f living resources[ Houde and Rutherford, 1993], and translocationo f nonindigenoussp ecies[C arlton,1 996] area mongt he moreo bviouse xampleso f suchh uman alterationsE. pisodicm eteorologicaelv entsa ndl ongert ermc limatec hangew ill continuet o compoundth e environmentaelf fectso f local andr egionalh umana ctivitiesI.n additiont o their profounde ffectso n the habitatsb, iodiversitya ndp roductivityo f coastael cosystems, environmentaclh angess ucha s thesew ill make coastale cosystemms ore susceptibleto naturalh azardsm, orec ostlyt o live in, ando f lessv aluet o society[ USNRC, 1994;W atson et al., 1998]. A majors cientificc hallengea risingf rom humane xploitationo f coastael cosystemiss thed evelopmenotf a predictiveu nderstandinogf ther elationshipbse tweenla nd-usep attems in coastald rainageb asinsa nd changesin the water quality and living resourceso f their receivingw atersM. any coastaal quatice cosystemisn,c ludingth e northernA driaticS ea( NA) Ecosystemast the Land-SeaM argin:D rainageB asint o CoastalS ea Coastala nd EstuarineS tudiesV olume 55, Pages1 -6 Copyright1 999 by the AmericanG eophysical 2 Introduction andC hesapeaBkea y( CB)( Fig.1 ),h aveb eens ubjectetdo increaseins n utrienint putsth at reflectc hangeisn land-usep atternsa s humanp opulationd ensitieisn coastawl atersheds havei ncrease[dS eaglee t al., thisv olume]T. hec haptertsh atf ollowa ret hep roducot f a workshotph atw asa firsta ttemptto conducat comparativaen alysiosf ecosystemofs t he land-sema arginf,r omd rainagbea sinto coastasle aa, sa meanosf developinag m orer obust understandionfg th er elationshibpest weecnh anginlga nd-uspea tternasn dt hes tructuraen d functiono f coastaml arinee cosystem[sM alonee t al., 1996]. Then ortherAnd riaticS ea( NA) andC hesapeaBkea y( CB)s ystemesx hibiitm portant similaritieasn dc ontrastths atm otivatetdh ec omparativaen alysirse porteidn thisb ook.F or purposoesf c omparisothne,S usquehanRnivae rd rainagbea sina ndt hem ainsteomf t heB ay, rathetrh anth eC B systemas a wholea ndi tsd rainagaer eaa, reo ftenu sedin thec haptetrhsa t followT. hisi s largelyb ecausneu trienitn putsto t ributarieost herth ant heS usquehananrae taken up within their estuariesa nd becauseu nknown but small amountso f dissolved inorgannicu trientasr ee xportetdo t hem ainsteomf CB (alsore ferretdo ast heC Bp roper). TheS usquehanRnaiv era ccountfso r ~70%o f totaln itrogenlo adsto theC B asa wholea nd moret han8 0% of thei nputo f dissolveidn organinci trogento them ainstemof theB ay [Schubealn dP ritchard1, 986].T otali nputi s usedf or theN A becausae significanbt,u t unknowfnra ctiono,f theP oR iverin puits advecterda pidlyo uto f theN A whilei nputsfr om the smallerr iversa ret akenu p withint heb oundarieosf theN A. Riverd ischargaec countfso r mostn utrienitn putst o boths ystemsa,n di nputso f freshwatearn dn utrientws ithine acha red ominatebdy a singlefi ver,t heP o in theN A and theS usquehaninna C B (Table1 ). On theo therh and,t hep opulatiodne nsityo f the NA drainagbea sina reai s moret hant wot imesth ato f theC B systeman dn earlyfo urt imesth at of theS usquehandnraa inagbea sin(T able1 ).D ifferenceins populatiodne nsitayr er eflected in nutrienetx portws ithf ivefinefl uxesb eingm oret hant wot imesh igheirn theN A thani n CB (Table1 ).E xpectedn itrogenfl uxesb asedo n populatiodne nsity[C olee t al., 1993],a re 31,35x0 10a m oyl -if ort heN A( 211p eopklem '2 f ort hee ntirNe Ab asina)n d1 3,57x0 1 0a moyl -(cid:127)f orC B( 57p eopklem '2 i nt heS usquehaRnivneaBr asinT)h. ea nalysoifsC olee ta l. [1993]w asd onef or 39 coastawl atershedresp resentindgra inagbea sinsw ithp opulation densitiersa ngingo vern earlyfo uro rderso f magnitudaen de ncompassitnhgep opulation densitieosf theN A andC B drainagbe asinsT. he estimateosf Boyntone t al. [ 1995]a nd Degobbiasn dG ilmartin[ 1 990]g ivenin Table1 arel owert hant hesep redictionbsu, tt hey arew ithint he9 5% confidencleim itso f ther egressiolnin ed erivedb y Colee t al. [ 1993]. Highern utrienlto adsa ndt heh ighr atioo f waterv olumeto freshwateflro w (freshwater fill time)o f theN A (~ 10y ) comparetdo theC B (~ 1 y) couldb ei nterpretetdo meant hat ther esidencteim eo f nutrientisn theN A is muchlo ngerth ani n theC B, resultinign more efficientr ecyclinga ndh igherp rimaryp roductioinn the NA. However,a s shownin Tables 1 and 2: CB hasa muchla rgedr rainagbea sina reap eru nitw aterv olume(2 ,250k m- (cid:127)) thant heN A (186k m'(cid:127)); nutrienlto adings caledto waterv olumeis aboufto urt imesh igherin CB than in the N A; thes urfacea reat o volumera tioo f CB is five timesh igherth ant hato f theN A. Theser elationshipnso t onlys uggestht atC B mayb e mores ensitivteo anthropogenic disturbanicne i tsd rainagbea sintsh ani st heN A, theys uggestht atc, ontrartyo the Smodlakae t al. 3 12* 13= 14* 46 ø 44" 7'7* 76* Figure1 . Planavr iewso f then ortherAn driatica ndC hesapeake 4 Introduction TABLE 1. Drainageb asins tatistics[B oyntone t al. 1995; Schubeal nd Pritchard1 986;D egobbisa nd Gilmartin1 990;S eaglee t al., thisv olume]. NorthernA driatic ChesapeakBe ay Po R. Total SusquehannRa. Total Areak, m2 71,000 118,000 70,000 166,000 Population 16,000,000 25,000,000 4,000,000 16,000,000 MeanR iverF low,m 3 s' (cid:127) 1,500 2,900 1,100 2,300 TN-Expor1t,0 6m oly '(cid:127) 11,575 23,610 7,570 10,860 TP-Expor1t0,6 m oyl -! 495 910 200 350 above (highern utrienti nputs and longer freshwaterf ill time for the NA then the CB), primaryp roductions houldb e highera ndn utrientc yclingm oree fficienti n CB. For the mostp an, responseosf the two systemsto increaseisn nutriente nrichmenat re expectedto be governedb y how nutrientin putsa red istributedb y circulationa ndm ixinga nd by their physicald imensionsT. he shape,l argerv olume,a nd openc onnectiono f the NA to theA driatica sa wholer esulti n morev ariable( lessp redictablec)i rculationp attemsth ani n CB wherew aterm otionsa rem oret opographicallcyo nstraine[dB oicourte t al., thisv olume]. CB is long,n arrowa nd shallowa ndm ostf reshwatera ndn utrienti nputso ccuri ts at north end3 20 km fromt heo ceanT. hisr esultsin strongn orth-soutghr adientsin salinitya ndi n the concentrationosf nutrientsa nd phytoplanktobni omass[ Hardinge t al., this volume].I n contrastt, he NA is deepera nd broadert han CB, and the Po River dischargeg enerally spreadsso utha longt he Italianc oastr esultingin rapide xportf rom theN A ands trongw est- eastg radientisn salinityn, utrientsa, ndp hytoplanktobnio massD. urings ummerl,o w salinity water tendst o spreadf arther eastwardd ue to seasonatl hermal stratificationa nd to the developmenotf a cyclonice ddyi n theN A [Boicouret t al., thisv olume].T hee ffectso f these differenceosn n utriendt istributionasre e xacerbatebdy thel ocationo f them ajorf ivefine inputsto thet wos ystemis.e, .,r apida dvectioonf theP od ischargteo thes outho uto f theN A comparetdo ther etentioonf theS usquehanRnaiv erd ischargweh osein flowi s at theh ead of CB. In additiont,h ef atea nde ffectso f nutrienitn putsd ependto a greate xtento n the timingo f peakisn freshwatfelor w( anda ssociatneudt rienfltu x)r elativteo t hea nnuacly cle of thermasl tratificatioann dt hed evelopmeonft then orthemc ycloniecd dyi n theN A and to the annualc ycleo f surfacete mperaturien CB. Differenceins thed istributioonf benthicar eaw ithd eptha rea lsoe xpectetdo i nfluence ther esponsoef aquatiecc osystemtos n utrienitn putsV. erticallyC, B is characterizbeyd broads, hallowfl anksw itha deepc hanne(1l 0-50m ) runningd ownt hea xiso f theB ay's mainstemw,h ilet heN A hasn arrowfl anksw itha broadf,l at bottomth ats lowlys hoaltso then orthT. heh ypsographcuicr vefso rt het wob asinasr es ubstantiadlliyff erenet,. g.,o nly 10%o f benthics urfacea reai n CB is deepetrh an1 5 m comparetdo moret han9 0% in the NA [Boicourett al.,t hisv olumeK; empe t al.,t hisv olume]T. hus,a lthougthh ec hannel

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