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P1:OTA/XYZ P2:ABC fm JWBK440/Bellinger March12,2010 22:8 PrinterName:YettoCome Freshwater Algae Freshwater Algae: Identification and Use as Bioindicators E dward G. Bellinger and David C. Sigee © 2010 John Wiley & Sons, Ltd. ISBN: 978-0-470-05814-5 i P1:OTA/XYZ P2:ABC fm JWBK440/Bellinger March12,2010 22:8 PrinterName:YettoCome Freshwater Algae Identification and Use as Bioindicators Edward G. Bellinger Environmental Science and Policy, Central European University, Hungary and David C. Sigee Manchester University, UK A John Wiley & Sons, Ltd, Publication iii P1:OTA/XYZ P2:ABC fm JWBK440/Bellinger March12,2010 22:8 PrinterName:YettoCome Thiseditionfirstpublished2010,(cid:1)C 2010byJohnWiley&Sons,Ltd Wiley-BlackwellisanimprintofJohnWiley&Sons,formedbythemergerofWiley’sglobalScientific,TechnicalandMedical businesswithBlackwellPublishing. RegisteredOffice: JohnWiley&Sons,Ltd,TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK OtherEditorialOffices: 9600GarsingtonRoad,Oxford,OX42DQ,UK 111RiverStreet,Hoboken,NJ07030-5774,USA Fordetailsofourglobaleditorialoffices,forcustomerservicesandforinformationabouthowtoapplyforpermissiontoreusethe copyrightmaterialinthisbookpleaseseeourwebsiteatwww.wiley.com/wiley-blackwell TherightoftheauthortobeidentifiedastheauthorofthisworkhasbeenassertedinaccordancewiththeCopyright,Designsand PatentsAct1988. Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmitted,inanyformorby anymeans,electronic,mechanical,photocopying,recordingorotherwise,exceptaspermittedbytheUKCopyright,Designsand PatentsAct1988,withoutthepriorpermissionofthepublisher. Wileyalsopublishesitsbooksinavarietyofelectronicformats.Somecontentthatappearsinprintmaynotbeavailablein electronicbooks. Designationsusedbycompaniestodistinguishtheirproductsareoftenclaimedastrademarks.Allbrandnamesandproduct namesusedinthisbookaretradenames,servicemarks,trademarksorregisteredtrademarksoftheirrespectiveowners.The publisherisnotassociatedwithanyproductorvendormentionedinthisbook.Thispublicationisdesignedtoprovideaccurate andauthoritativeinformationinregardtothesubjectmattercovered.Itissoldontheunderstandingthatthepublisherisnot engagedinrenderingprofessionalservices.Ifprofessionaladviceorotherexpertassistanceisrequired,theservicesofa competentprofessionalshouldbesought. LibraryofCongressCataloguing-in-PublicationData Bellinger,E.G. Freshwateralgae:identificationanduseasbioindicators/EdwardG.BellingerandDavidC.Sigee. p. cm. Includesbibliographicalreferencesandindex. ISBN978-0-470-05814-5(cloth) 1.Freshwateralgae. 2.Indicators(Biology). 3.Environmentalmonitoring. I.Sigee,D.C. II.Title. QK570.25.B452010 579.8–dc22 2009052145 AcataloguerecordforthisbookisavailablefromtheBritishLibrary. Setin10/12ptTimesbyAptaraInc.,NewDelhi,India PrintedinGreatBritainbyAntonyRowe,Ltd.Chippenham,Wilts Firstimpression2010 iv P1:OTA/XYZ P2:ABC fm JWBK440/Bellinger March12,2010 22:8 PrinterName:YettoCome Contents Preface ix CopyrightAcknowledgements xi 1 IntroductiontoFreshwaterAlgae 1 1.1 Generalintroduction 1 1.1.1 Algae–anoverview 1 1.1.2 Algaeasprimaryproducers 2 1.1.3 Freshwaterenvironments 3 1.1.4 Planktonicandbenthicalgae 3 1.1.5 Sizeandshape 4 1.2 Taxonomicvariation–themajorgroupsofalgae 5 1.2.1 Microscopicalappearance 8 1.2.2 Biochemistryandcellstructure 9 1.2.3 Molecularcharacterizationandidentification 9 1.3 Blue-greenalgae 12 1.3.1 Cytology 12 1.3.2 Morphologicalandtaxonomicdiversity 13 1.3.3 Ecology 15 1.3.4 Blue-greenalgaeasbio-indicators 17 1.4 Greenalgae 17 1.4.1 Cytology 17 1.4.2 Morphologicaldiversity 17 1.4.3 Ecology 18 1.4.4 Greenalgaeasbioindicators 21 1.5 Euglenoids 21 1.5.1 Cytology 22 1.5.2 Morphologicaldiversity 22 1.5.3 Ecology 23 1.5.4 Euglenoidsasbioindicators 23 1.6 Yellow-greenalgae 23 1.6.1 Cytology 23 1.6.2 Morphologicaldiversity 24 1.6.3 Ecology 24 1.6.4 Yellow-greenalgaeasbioindicators 24 v P1:OTA/XYZ P2:ABC fm JWBK440/Bellinger March12,2010 22:8 PrinterName:YettoCome vi CONTENTS 1.7 Dinoflagellates 25 1.7.1 Cytology 25 1.7.2 Morphologicaldiversity 26 1.7.3 Ecology 27 1.8 Cryptomonads 28 1.8.1 Cytology 28 1.8.2 Comparisonwitheuglenoidalgae 29 1.8.3 Biodiversity 29 1.8.4 Ecology 30 1.8.5 Cryptomonadsasbioindicators 30 1.9 Chrysophytes 31 1.9.1 Cytology 31 1.9.2 Morphologicaldiversity 31 1.9.3 Ecology 32 1.9.4 Chrysophytesasbioindicators 32 1.10 Diatoms 33 1.10.1 Cytology 33 1.10.2 Morphologicaldiversity 35 1.10.3 Ecology 38 1.10.4 Diatomsasbioindicators 39 1.11 Redalgae 40 1.12 Brownalgae 40 2 Sampling,BiomassEstimationandCountsofFreshwaterAlgae 41 A. PLANKTONICALGAE 41 2.1 Protocolforcollection 41 2.1.1 Standingwaterphytoplankton 42 2.1.2 Riverphytoplankton 46 2.2 Modeofcollection 46 2.2.1 Phytoplanktontrawlnet 46 2.2.2 Volumesamplers 48 2.2.3 Integratedsampling 49 2.2.4 Sedimenttraps 50 2.3 Phytoplanktonbiomass 51 2.3.1 Turbidity 51 2.3.2 Dryweightandash-freedryweight 53 2.3.3 Pigmentconcentrations 53 2.4 Flowcytometry:automatedanalysisofphytoplanktonpopulations 60 2.5 Microscopecountsofspeciespopulations 61 2.5.1 Samplepreservationandprocessing 61 2.5.2 Speciescounts 63 2.5.3 Conversionofspeciescountstobiovolumes 70 2.5.4 Chemicalcleaningofdiatoms 75 2.6 Diversitywithinsingle-speciespopulations 76 2.6.1 Molecularanalysis 76 2.6.2 Analyticalmicroscopicaltechniques 77 P1:OTA/XYZ P2:ABC fm JWBK440/Bellinger March12,2010 22:8 PrinterName:YettoCome CONTENTS vii B. NON-PLANKTONICALGAE 82 2.7 Deepwaterbenthicalgae 82 2.7.1 Benthic–pelagiccoupling 83 2.7.2 Benthicalgaeandsedimentstability 84 2.7.3 Invertebrategrazingofbenthicalgae 84 2.8 Shallowwatercommunities 84 2.8.1 Substrate 84 2.8.2 Algalcommunities 87 2.9 Algalbiofilms 89 2.9.1 Mucilaginousbiofilms 89 2.9.2 Biomass 89 2.9.3 Taxonomiccomposition 91 2.9.4 Matrixstructure 93 2.10 Periphyton–algalmats 93 2.10.1 Inorganicsubstratum 93 2.10.2 Plantsurfaces 95 3 AlgaeasBioindicators 99 3.1 Bioindicatorsandwaterquality 99 3.1.1 Biomarkersandbioindicators 99 3.1.2 Characteristicsofbioindicators 100 3.1.3 Biologicalmonitoringversuschemicalmeasurements 101 3.1.4 Monitoringwaterquality:objectives 102 3.2 Lakes 105 3.2.1 Contemporaryplanktonicandattachedalgaeasbioindicators 105 3.2.2 Fossilalgaeasbioindicators:lakesedimentanalysis 106 3.2.3 Waterqualityparameters:inorganicandorganicnutrients,acidityandheavymetals 110 3.3 Wetlands 119 3.4 Rivers 121 3.4.1 Theperiphytoncommunity 121 3.4.2 Riverdiatoms 121 3.4.3 Evaluationofthediatomcommunity 122 3.4.4 Humanimpactsanddiatomindices 123 3.4.5 Calculationofdiatomindices 126 3.4.6 Practicalapplicationsofdiatomindices 128 3.5 Estuaries 133 3.5.1 Ecosystemcomplexity 133 3.5.2 Algaeasestuarinebioindicators 134 4 AKeytotheMoreFrequentlyOccurringFreshwaterAlgae 137 4.1 Introductiontothekey 137 4.1.1 Usingthekey 137 4.1.2 Morphologicalgroupings 138 4.2 Keytothemaingeneraandspecies 138 P1:OTA/XYZ P2:ABC fm JWBK440/Bellinger March12,2010 22:8 PrinterName:YettoCome viii CONTENTS 4.3 Listofalgaeincludedandtheiroccurrenceinthekey 241 4.4 Algalidentification:bibliography 243 Glossary 245 References 255 Index 265 P1:OTA/XYZ P2:ABC fm JWBK440/Bellinger March12,2010 22:8 PrinterName:YettoCome Preface Almost any freshwater or brackish water site will human and natural impacts in aquatic systems be- containoneormanyspeciesofalgae.Althoughthey causeoftheiroftenrapidresponsetochangesinthe aremainlymicroscopicandthereforenotasvisually environment. Examples include nutrient enrichment apparentaslargeraquaticorganisms,suchashigher (eutrophication),industrialpollutionandchangesto plants or fish, algae play an equally important role the hydrological regime of the water body. Some intheecologyofthesewaterbodies.Theirpresence groupsofalgaepreservewellasfossilsingeological cansometimesbenoticedwhentheyoccurasdense deposits such as lake sediments, analysis of which populations, colouring the water and in some cases givesusinformationonpastenvironmentalchanges. formingmassivesurfacescums. This book comes at a time of increasing concern Freshwater algae constitute a very diverse group over the widespread effects of human activities on oforganisms.Theirrangeofshapesandbeautywhen the general environment of this planet. Monitoring viewed through a microscope have delighted biol- shiftsinalgalpopulationgivesusaninsightintothese ogists for more than a hundred years. They have an changes.Weneedtobeabletoassessthe‘health’of enormousrangeofsizefromlessthanonemicrometre aquaticsystemssuchaslakesandrivers,sincewateris toseveralcentimetres(forthestoneworts)–equalling vitaltobothhumanandgeneralecosystemsurvival. the size span (104) for higher plants seen in a trop- Knowledge of algal population dynamics can help icalrainforestAlgalmorphology isdiverse,ranging usdevelopeffectivemanagementstrategiesforthose from single cells to complex colonies and fila- systems.Includedinthisbookaresectionsonthegen- ments. Some species are capable of active move- eralfeaturesofthemainfreshwateralgalgroupswith ment.Theterm‘algae’embracesanumberofphyla notesontheirecology,methodsofsamplecollection (e.g. Cyanophyta, Bacillariophyta and Chlorophyta) andenumeration,usingalgaeasindictorsofenviron- of chlorophyll-containing organisms with different mentalconditionsand,finally,akeytotheidentifica- growth forms and cytologies. Algae are important tionofthemorefrequentlyoccurringgenera.Theau- primary producers in both freshwater and marine thorshavetriedtocombinedescriptivematerialwith systems. In many lakes and rivers they generate originalcolourphotographsandlinedrawings,where biomass which is the foundation of diverse food possible, to help the reader. We would also like to chains. Although algae have beneficial impacts on gratefullyacknowledgethehelpandencouragement aquaticecosystemstheycanalsohaveadverseeffects. ofcolleaguesandstudents,andparticularlyappreci- When present in very large numbers they can pro- ate the direct contributions of postdoctoral workers duce ‘blooms’ that, on decomposition, deoxygenate and research studentsmentioned inthelistof copy- the water – causing fish death and other ecological rightacknowledgements.Wewouldalsoliketothank problems.Somealgaeproducetoxinsthatarelethalto ourfamiliesfortheirunderstandingandpatiencedur- bothaquaticandterrestrialorganisms.Itisimportant ingthepreparationofthistext. tobeawareoftheseimpactsandtomonitorwatersfor We hope that all those using the book will find thepresenceofthesepotentiallyharmfulorganisms. it useful, and will enjoy the numerous colour pho- Algae can be used to flag up and assess a range of tographsoftheseverybeautifulorganisms. ix P1:OTA/XYZ P2:ABC fm JWBK440/Bellinger March12,2010 22:8 PrinterName:YettoCome Copyright Acknowledgements Weareverygratefultothefollowingindividualsfor material – Academic Press, American Health As- allowing us to publish previously unpublished mat- sociation, Cambridge University Press, Journal of ter: Andrew Dean (Tables 2.3, 2.4), Matt Capstick PlanktonResearch,McGraw-Hill,Phycologia,Pren- (Figs.4.10–4.12,4.42,4.62,4.64,4.66,4.68–4.70a, ticeHall. 4.73a)andHudaQari(Fig.2.8). Wealsothankthefollowingcopyrightholdersfor giving us permission to use previously published xi P1:OTA/XYZ P2:ABC c01 JWBK440/Bellinger March15,2010 11:55 PrinterName:YettoCome 1 Introduction to Freshwater Algae 1.1 General introduction tosynthetic–generatingcomplexcarboncompounds from carbon dioxide and light energy. Some algae Algaearewidelypresentinfreshwaterenvironments, have become secondarily heterotrophic, taking up such as lakes and rivers, where they are typically complexorganicmoleculesbyorganotrophy orhet- presentasmicro-organisms–visibleonlywiththeaid erotrophy (Tuchman, 1996), but still retaining fun- ofalightmicroscope.Althoughrelativelyinconspic- damentalgeneticaffinitieswiththeirphotosynthetic uous,theyhaveamajorimportanceinthefreshwater relatives(Pfandletal.,2009). environment, both in terms of fundamental ecology The term ‘algae’ (singular alga) is not strictly a andinrelationtohumanuseofnaturalresources. taxonomic term but is used as an inclusive label for Thisbookconsidersthediversityofalgaeinfresh- a number of different phyla that fit the broad de- waterenvironmentsandgivesageneraloverviewof scriptionnotedabove.Theseorganismsincludeboth the major groups of these organisms (Chapter 1), prokaryotes(Section1.3,cellslackingamembrane- methods of collection and enumeration (Chapter 2) boundnucleus)andeukaryotes(cellswithanucleus and keys to algal groups and major genera (Chap- plustypicalmembrane-boundorganelles). ter 4). Algae are considered as indicators of en- Humanshavelongmadeuseofalgalspecies,both vironmental conditions (bioindicators) in terms of living and dead. Fossil algal diatomite deposits, for individual species (Chapter 1) and as communities example, in the form of light but strong rocks, have (Chapter3). beenusedasbuildingmaterialsandfiltrationmediain waterpurificationandswimmingpools.Somefossil algae,suchasBotryococcus,cangiverisetooil-rich deposits.Certainspeciesofgreenalgaearecultivated 1.1.1 Algae – an overview for the purpose of extracting key biochemicals for use in medicine and cosmetics. Even blue-green al- Theword‘algae’originatesfromtheLatinwordfor gae,oftenregardedasnuisanceorganisms,mayhave seaweed and is now applied to a broad assemblage beneficialuses.ThisisparticularlythecaseforSpir- of organisms that can be defined both in terms of ulina,whichwasharvestedbytheAztecsofMexico morphologyandgeneralphysiology.Theyaresimple andisstillusedbythepeoplearoundLakeChadas organisms, without differentiation into roots, stems a dietary supplement. Spirulina tablets may still be and leaves – and their sexual organs are not en- obtained in some health food shops. Blue-green al- closedwithinprotectivecoverings.Intermsofphysi- gaeare,however,arebetterknowninthefreshwater ology,theyarefundamentallyautotrophic(obtaining environment as nuisance organisms, forming dense all their materials from inorganic sources) and pho- blooms having adverse effects on human activities Freshwater Algae: Identification and Use as Bioindicators E dward G. Bellinger and David C. Sigee © 2010 John Wiley & Sons, Ltd. ISBN: 978-0-470-05814-5

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Freshwater Algae: Identification and Use as Bioindicators provides a comprehensive guide to temperate freshwater algae, with additional information on key species in relation to environmental characteristics and implications for aquatic management. The book uniquely combines practical material on t
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