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ThorpandCovich’sFreshwaterInvertebrates

AGlobalSeriesofBooksontheIdentiﬁcation, Ecology,andGeneralBiologyofInlandWaterInvertebrates byExpertsfromAroundtheWorld

FourthEdition

SeriesEditor:JamesH.Thorp

VolumeI:EcologyandGeneralBiology

EditedbyJamesH.ThorpandD.ChristopherRogers Published2015

VolumeII:KeystoNearcticFauna

EditedbyJamesH.ThorpandD.ChristopherRogers Published2016

VolumeIII:KeystoNeotropicalHexapoda

EditedbyNeusaHamada,JamesH.Thorp,andD.ChristopherRogers Published2018

VolumeIV:KeystoPalaearcticFauna

EditedbyD.ChristopherRogersandJamesH.Thorp Published2019

VolumeV:KeystoNeotropicalandAntarcticFauna

EditedbyCristinaDamborenea,D.ChristopherRogersandJamesH.Thorp Published2020

VolumesinDevelopment

KeystoAustralasianFauna

KeystoAfrotropicalFauna

PlannedFutureVolumes

KeystoOrientalandOceanianFauna

KeystoOrientalandOceanianHexapoda

KeystoNearcticHexapoda

KeystoPalaearcticHexapoda

RelatedPublications

FieldGuidetoFreshwaterInvertebratesofNorthAmerica

JamesH.ThorpandD.ChristopherRogers FirstEdition(2011)

KeystoNeotropicaland AntarcticFauna

ThorpandCovich’sFreshwater Invertebrates e VolumeV

FourthEdition

Editedby

CristinaDamborenea

ConsejoNacionaldeInvestigacionesCientı´ﬁcasyTe ´ cnicas, MuseodeLaPlata,FacultadCienciasNaturalesyMuseo, UniversidadNacionaldeLaPlata,Argentina

D.ChristopherRogers

KansasBiologicalSurveyand,TheBiodiversityInstitute UniversityofKansas,Lawrence,KS,USA

JamesH.Thorp

KansasBiologicalSurveyand DepartmentofEcologyandEvolutionaryBiology UniversityofKansas,Lawrence,KS,USA

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ToLaura,Cecilia,andSantiago,mywonderful children,andtoRaulfortheircomprehension. Marı´aCristinaDamborenea

Toourauthorswhomadethisbookpossible.

Toourwonderfulchildren,SaraandZachary,forlovingthe naturalworldandunderstandingtheirfather’sobsessionwithit.

RobertoG.Alves [Chapter12:PhylumAnnelida]DepartamentoZoologia,UniversidadeFederaldeJuizde Fora,JuizdeFora,MG,Brazil;email: gama.alves@ ufjf.edu.br

MonicaAmmonFernandez [Chapter11:PhylumMollusca]Laborato ´ riodeMalacologia/Laborato ´ riode Refere ˆ nciaNacionalparaEsquistossomose-Malacologia,InstitutoOswaldoCruz,Fundac¸a ˜ oOswaldoCruz, RiodeJaneiro,Brazil;email: ammon@ioc.ﬁocruz.br

SoniaBarbosadosSantos [Chapter11:PhylumMollusca]Laborato ´ riodeMalacologiaLı´mnicaeTerrestre, DepartamentodeZoologia,UniversidadedoEstado doRiodeJaneiro(UERJ),Brazil;email: gundlachia@ yahoo.com.br

GaryM.Barker [Chapter11:PhylumMollusca]LandcareResearch,Lincoln,Canterbury,NewZealand; email: barkerg@landcareresearch.co.nz

MariaIsabelBerning [Chapter11:PhylumMollusca] BavarianStateCollectionofZoology(ZSM),Section Mollusca,Munich,Germany;email: gomez.berning@ googlemail.com

FelipeBezerraRibeiro [Chapter23:ClassMalacostraca] DepartamentodeZoologia(Laborato ´ riodeCarcinologia),InstitutodeBiocie ˆ ncias,UniversidadeFederal doRioGrandedoSul,Brazil;email: fbribeiro.bio@ gmail.com

MatthewG.Bolek [Chapter10:PhylumNematomorpha] DepartmentofIntegrativeBiology,OklahomaState University,Stillwater,Oklahoma,USA;email: bolek@ okstate.edu

GeorginaBond-Buckup [Chapter23:ClassMalacostraca]IndependentResearcher,retiredfromtheUniversidadeFederaldoRioGrandedoSul,PortoAlegre,RS, Brazil;email: gbond_buckup@yahoo.com.br

FranciscoBrusa [Chapter5:PhylumPlatyhelminthes] Divisio ´ nZoologı´aInvertebrados,MuseodeLaPlata, FCNyM,UNLP/CONICET,Argentina;email: fbrusa@ fcnym.unlp.edu.ar

AlessandraBueno [Chapter23:ClassMalacostraca] DepartamentodeBiologia,UniversidadeFederalde Lavras,Brazil;email: aapbueno@yahoo.com.br

LucianaBurdman [Chapter2:Protozoa]Instituteof EnvironmentalResearchandEngineering,University ofSanMartin,CONICET,BuenosAires,Argentina; email: lburdman@unsam.edu.ar

LudimilaCalheira [Chapter3:PhylumPorifera]DepartamentodeZoologia,UniversidadeFederaldePernambuco,Recife,Brazil;email: calheiralaurindo@gmail.com

AnaIsabelCamacho [Chapter23:ClassMalacostraca] MuseoNacionaldeCienciasNaturales(CSIC),DepartamentodeBiodiversidadyBiologı´aEvolutiva,Madrid, Spain;email: mcnac22@mncn.csic.es

AlineCarvalhodeMattos [Chapter11:PhylumMollusca]Laborato ´ riodeMalacologia,/Laborato ´ riode Refere ˆ nciaNacionalparaEsquistossomose-Malacologia,InstitutoOswaldoCruz,Fundac¸a ˜ oOswaldoCruz, RiodeJaneiro,Brazil;email: amattos@ioc.ﬁocruz.br

SilvanaCarvalhoThiengo [Chapter11:PhylumMollusca]Laborato ´ riodeMalacologia/Laborato ´ riode Refere ˆ nciaNacionalparaEsquistossomose-Malacologia,InstitutoOswaldoCruz,Fundac¸aoOswaldoCruz, RiodeJaneiro,Brazil;email: sthiengo@ioc.ﬁocruz.br

Marı´aCristinaClaps [Chapter2:Protozoa]Institutode Limnologı´a"Dr.R.A.Ringuelet",LaPlata,Buenos Aires,Argentina;email: claps@ilpla.edu.ar

RosaGracielaCohen [Chapter19:ClassBranchiopoda] DepartamentodeBiodiversidadyBiologı´aExperimental,FacultaddeCienciasExactasyNaturales,UniversidaddeBuenosAires,Argentina;email: cohen@ bg.fcen.uba.ar

GonzaloA.Collado [Chapter11:PhylumMollusca]FacultaddeCiencias,DepartamentodeCienciasBa ´ sicas, UniversidaddelBı´o-Bı´o,Chilla ´ n,Chile;email: collado. gonzalo@gmail.com

YanisCruz-Quintana [Chapter21:ClassCopepoda] EscueladeAcuiculturayPesquerı´a,FacultaddeCienciasVeterinarias,UniversidadTe ´ cnicadeManabı´, Ecuador;email: cqyanis@gmail.com

MariaGabrielaCuezzo [Chapter11:PhylumMollusca] InstitutodeBiodiversidadNeotropical(CONICETUNT),UniversidadNacionaldeTucuma ´ n,Tucuma ´ n, Argentina;email: gcuezzo@webmail.unt.edu.ar

CristinaDamborenea [Chapter1:Introduction,Chapter 2:Protozoa,Chapter5:PhylumPlatyhelminthes, Chapter11:PhylumMollusca,Chapter12:Phylum Annelida]Divisio ´ nZoologı´aInvertebrados,Museode LaPlata,FCNyM,UNLP/CONICET,Argentina; email: cdambor@fcnym.unlp.edu.ar

PaulDeLey [Chapter9:PhylumNematoda]Department ofNematology,UniversityofCaliforniaRiverside, Riverside,CA,USA;email: ude.rcu@yeled.luap

CristinadeVillalobos [Chapter10:PhylumNematomorpha]MuseodeCienciasNaturales,FCNyM,UNLP, Argentina;email: villalo@fcnym.unlp.edu.ar

Marı´aIreneDeserti [Chapter4:PhylumCnidaria]InstitutodeInvestigacionesMarinasyCosteras,Universidad NacionaldeMardelPlata-CONICET,MardelPlata, BuenosAires,Argentina;email: mdeserti@mdp.edu.ar

EdinaldoNelsondosSantos-Silva [Chapter21:Class Copepoda]InstitutodePesquisasdaAmazo ˆ nia,Manaus,AM,Brazil;email: nelson@inpa.gov.br

MarceloE.Doucet [Chapter9:PhylumNematoda]InstitutodeDiversidadyEcologı´aAnimal(CONICETUNC)andCentrodeZoologı´aAplicada,Universidad NacionaldeCo ´ rdoba,Co ´ rdoba,Argentina;email: mdoucet@efn.uncor.edu

MariaCristinaDreherMansur [Chapter11:Phylum Mollusca]GrupodePesquisadoCNPq,Biodiversidade deMoluscosContinentais,MuseudeCie ˆ nciasNaturais,PortoAlegre,RS,Brazil;email: mcrismansur@ gmail.com

ElizangelaFeitosadaSilva [Chapter11:PhylumMollusca]Laborato ´ riodeRefere ˆ nciaNacionalparaEsquistossomose,Malacologia,InstitutoOswaldoCruz, Fundac¸a ˜ oOswaldoCruz,RiodeJaneiro,Brazil;email: efeitosa@ioc.ﬁocruz.br

LeonardoFerna ´ ndez [Chapter2:Protozoa]CIRENYS, UniversidadBernardoO’Higgins,Santiago,Chile; email: limnoleo@gmail.com

NelsonFerretti [Chapter16:PhylumArthropoda:IntroductionandArachnida]InstitutodeCienciasBiolo ´ gicasy Biome ´ dicasdelSur(INBIO-SUR-CONICET-UNS), BahıaBlanca,BuenosAires,Argentina;email: nferretti@conicet.gov.ar

DiegoFontaneto [Chapter8:PhylumRotifera]Water ResearchInstitute(IRSA),NationalResearchCouncil (CNR),VerbaniaPallanza,Italy;email: d.fontaneto@ ise.cnr.it

NataliaA.Fredes [Chapter16:PhylumArthropoda: IntroductionandArachnida]DepartamentodeBiologı´a,UniversidadNacionaldeMardelPlata, Argentina;email: fredesnaty@gmail.com

StuartR.Gelder [Chapter12:PhylumAnnelida]DepartmentofScienceandMath,UniversityofMaineatPresqueIsle,PresqueIsle,Maine04769,USA;email: stuart.gelder@umpi.edu

Joa ˜ oGil [Chapter11:PhylumAnnelida]Centred’Estudis Avanc¸atsdeBlanes,CEAB-CSIC,Blanes,Catalunya, Spain;email: gil@ceab.csic.es

ChristopherJ.Glasby [Chapter12:PhylumAnnelida] Museum&ArtGalleryoftheNorthernTerritory,Darwin,Australia;email: Chris.glasby@nt.gov.au

SamuelGo ´ mez [Chapter21:ClassCopepoda]Institutode CienciasdelMaryLimnologı´a,UNAM.Unidad Acade ´ micaMazatla ´ n,Mazatla ´ n,Mexico;email: samuelgomez67@gmail.com

ExequielR.Gonza ´ lez [Chapter23:ClassMalacostraca] UniversidadSantoToma ´ s,Santiago,Chile;email: egonzalezb@santotomas.cl

PriscilaA.Grohmann [Chapter4:PhylumCnidaria] InstitutodeBiologia,DepartamentodeZoologia,UniversidadeFederaldoRiodeJaneiro,RiodeJaneiro, Brazil;email: grohmann@biologia.ufrj.br

Ju ¨ rgenGuerrero-Kommritz [Chapter23:ClassMalacostraca]DevelopmentofAppliedBiology,Fundabas Foundation,Bogota ´ ,Colombia;email: greledone@ hotmail.com

RobertoGuidetti [Chapter15:PhylumTardigrada] DepartmentofLifeSciences,UniversityofModena andReggioEmilia,Modena,Italy;email: roberto. guidetti@unimore.it

MarthaA.Gutie ´ rrez-Aguirre [Chapter21:ClassCopepoda]UniversidaddeQuintanaRoo,Cozumel, Mexico;email: margutierrez@uqroo.edu.mx

DiegoEduardoGutie ´ rrezGre ´ goric [Chapter11:Phylum Mollusca]Divisio ´ nZoologı´aInvertebrados,Museode LaPlata,FCNyM,UNLP/CONICET,Argentina; email: dieguty@yahoo.com.ar

NeusaHamada [Chapter.17:PhylumArthropoda:IntroductiontoCrustaceaandHexapoda]Coordenac¸aode Biodiversidade,InstitutoNacionaldePesquisasda Amazo ˆ nia-INPA,Manaus,AM,Brazil;email: nhamada@inpa.gov.br

BenHanelt [Chapter10:PhylumNematomorpha]DepartmentofBiology,UniversityofNewMexico,Albuquerque,NM,USA;email: bhanelt@unm.edu

BrendaJ.Hann [Chapter19:Branchiopoda]Department ofBiologicalSciences,UniversityofManitoba, Canada;email: hann@cc.umanitoba.ca

JanetHiguti [Chapter20:Ostracoda]StateUniversityof Maringa ´ ,CentreforResearchonLimnology,IchthyologyandAquaculture,GraduateProgrammein EcologyofInlandWaterEcosystems,Maringa ´ ,Parana ´ , Brazil;email: janethiguti@gmail.com

Cristia ´ nItuarte [Chapter11:PhylumMollusca]Divisio ´ n Zoologı´aInvertebrados,MuseoArgentinodeCiencias Naturales,BuenosAires,Argentina;email: ituarte@ macn.gov.ar

CarlosG.Jara [Chapter23:ClassMalacostraca]Instituto deCienciasMarinasyLimnolo ´ gicas,Universidad AustraldeChile,Valdivia,Chile;email: cjara@ uaChapter.cl

SusanaB.Jose ´ dePaggi [Chapter8:PhylumRotifera] InstitutodeLimnologia,SantaFe,Argentina;email: sjdepaggi@gmail.com

q ukaszKaczmarek [Chapter15:PhylumTardigrada] DepartmentofAnimalTaxonomyandEcology, AdamMickiewiczUniversityinPoznan,Poznan, Poland;email: kaczmar@amu.edu.pl

AnushKosakyan [Chapter2:Protozoa]InstituteofParasitology,BiologyCentreoftheCzechAcademyof Sciences, Ceske ´ Budejovice,CzechRepublic/Institute ofBiosciences,DepartmentofZoology,Universityof SaoPaulo,Brazil;email: anna.kosakyan@gmail.com

ValentynaKrashevska [Chapter2:Protozoa]J.F.BlumenbachInstituteofZoologyandAnthropology, GeorgAugustUniversityGo ¨ ttingen,Go ¨ ttingen, Germany;email: valentyna.krashevska@biologie.unigoettingen.de

GabrielaC.Ku ¨ ppers [Chapter2:Protozoa]Divisio ´ n Invertebrados,MuseoArgentinodeCienciasNaturales "BernardinoRivadavia",BuenosAires,Argentina; email: gabrielakuppers@gmail.com

EnriqueM.Lara [Chapter2:Protozoa]Laboratoryof SoilBiodiversity,UniversityofNeucha ˆ tel,Neucha ˆ tel, Switzerland/RealJardı´nBota ´ nicodeMadrid,CSIC, Spain;email: enrique.lara@rjb.csic.es

PaolaLax [Chapter9:PhylumNematoda]Institutode DiversidadyEcologı´aAnimal(CONICET-UNC)and CentrodeZoologı´aAplicada,UniversidadNacionalde Co ´ rdoba,5000Co ´ rdoba,Argentina;email: laxpaola@ gmail.com

AnaMariaLeal-Zanchet [Chapter5:PhylumPlatyhelminthes]InstitutodePesquisasdePlana ´ riasandProgramadePo ´ s-Graduac¸a ˜ oemBiologia,Universidade doValedoRiodosSinos,UNISINOS,SaoLeopoldo, RioGrandedoSul,Brazil;email: zanchet@unisinos.br

LuizEduardoMacedodeLacerda [Chapter11:Phylum Mollusca]Laborato ´ riodeMalacologiaLı´mnicaeTerrestre,DepartamentodeZoologia,Universidadedo EstadodoRiodeJaneiro(UERJ),Brazil;email: lacerdauerjbio@yahoo.com.br

Ce ´ lioMagalha ˜ es [Chapter23:ClassMalacostraca]InstitutoNacionaldePesquisasdaAmazo ˆ nia,Manaus, AM,Brazil;email: celiomagalhaes.inpa@gmail.com

DanielMansurPimpa ˜ o [Chapter11:PhylumMollusca] InstitutoBrasileirodoMeioAmbienteedosRecursos NaturaisRenova ´ veis(IBAMA),GO,Brazil;email: danielpimpao@yahoo.com.br

FernandoL.Mantelatto [Chapter23:ClassMalacostraca]Laborato ´ riodeBioecologiaeSistema ´ ticade Crusta ´ ceos(LBSC),DepartamentoofBiologia,FaculdadedeFilosoﬁa,Cie ˆ nciaseLetrasdeRibeiraoPreto (FFCLRP),UniversidadedeSaoPaulo(USP),Ribeirao Preto,SP,Brazil;email: ﬂmantel@usp.br

MercedesR.Marchese [Chapter12:PhylumAnnelida]InstitutoNacionaldeLimnologı´a(CONICET-UNL),Santa Fe,Argentina;email: mercedes.marchese@gmail.com

Marı´aCristinaMarinone [Chapter8:PhylumRotifera] DepartamentodeBiodiversidadyBiologı´aExperimental,FacultaddeCienciasExactasyNaturales,UniversidaddeBuenosAires,Argentina;email: marinone@ bg.fcen.uba.ar

TaisaM.Marques [Chapter21:ClassCopepoda]Laborato ´ riodeEcologiaMoleculareParasitologiaEvolutiva, UniversidadeFederaldo,Parana ´ ,Curitiba,Parana ´ , Brazil;email: taisaorsi@gmail.com

KoenMartens [Chapter20:ClassOstracoda]Royal BelgianInstituteofNaturalSciences(RBINS),Brussels,Belgium;DepartmentofBiology,Universityof Ghent,Gent,Belgium;email: darwinula@gmail.com

DanielMartin [Chapter11:PhylumAnnelida]Centre d’EstudisAvanc¸atsdeBlanes,CEAB-CSIC,Catalunya,Spain;email: dani@ceab.csic.es

PabloA.Martı ´ nez [Chapter16:PhylumArthropoda: IntroductionandArachnida]DepartamentodeBiologı ´ a, UniversidadNacionaldeMardelPlata,Argentina; email: pamartin@mdp.edu.ar

SandraMcInnes [Chapter15:PhylumTardigrada]British AntarcticSurvey,NaturalEnvironmentResearch Council,Cambridge,UK;email: s.mcinnes@bas.ac.uk

NancyF.Mercado-Salas [Chapter21:ClassCopepoda] CentrumfurNaturkunde,CeNak/CenterofNatural HistoryHamburg,Universita ¨ tHamburg,Zoological Museum,Hamburg,Germany;Martin-Luther-KingPlatz3,20146Hamburg,Germany;email: nancy. mercado.salas@uni-hamburg.de

IgorChristoMiyahira [Chapter11:PhylumMollusca] UniversidadeEstadualdoRiodeJaneiro,Riode Janeiro,Brazil;email: icmiyahira@yahoo.com.bre

DianeR.Nelson [Chapter15:PhylumTardigrada]DepartmentofBiologicalSciences,EastTennesseeState University,USA;email: nelsond@etsu.edu

CarolinaNoren ˜ a [Chapter5:PhylumPlatyhelminthes] DepartamentoBiodiversidadyBiologı´aEvolutiva, MuseoNacionaldeCienciasNaturales(CSIC), Madrid,Spain;email: mcnnj92@mncn.csic.es

ThiagoNunesAntoniazzi [Chapter11:PhylumMollusca] UniversidadeFederaldoRioGrandedoSul,Porto Alegre,Brazil;email: thiago.antoniazzi@hotmail.com

AlejandroOceguera-Figueroa [Chapter12:Phylum Annelida]LaboratoriodeHelmintologı´a,Institutode Biologı´a,UniversidadNacionalAuto ´ nomadeMe ´ xico; email: aoceguera@ib.unam.mx

JanineOliveiraArruda [Chapter11:PhylumMollusca] GrupodePesquisadoCNPq,BiodiversidadedeMoluscosContinentais,MuseudeCie ˆ nciasNaturais,Porto Alegre,RioGrandedoSul,Brazil;email: arrudajo@ gmail.com

MariaCristinaOrellanaLiebbe [Chapter13:Phylum Ectoprocta(Bryozoa)]FacultaddeCiencias,UniversidadCato ´ licadelaSantı´simaConcepcio ´ n,Concepcio ´ n, Chile;email: mcorella@ucsc.cl

XimenaMariaConstanzaOvando [Chapter11:Phylum Mollusca]Laborato ´ riodeMalacologiaLı´mnicaeTerrestre,DepartamentodeZoologia,Universidadedo EstadodoRiodeJaneiro(UERJ),Brazil;email: velliger08@gmail.com

ManuelPedraza [Chapter23:ClassMalacostraca]Programa dePo ´ s-Graduac¸aoemSistema ´ tica,TaxonomiaAnimale Biodiversidade,MuseudeZoologiadaUniversidadede Sa ˜ oPaulo,Brazil;email: manupedrazam@gmail.com

CarlosPedraza-Lara [Chapter23ClassMalacostraca] LicenciaturaenCienciaForense,FacultaddeMedicina,UniversidadNacionalAuto ´ nomadeMe ´ xico,CiudadUniversitaria,DistritoFederal,Mexico;email: pedrazal@gmail.com

AlmirRoge ´ rioPepato [Chapter16:PhylumArthropoda: IntroductionandArachnida]DepartamentodeZoologia,UniversidadeFederaldeMinasGerais,Belo Horizonte,Brazil;email: apepato@gmail.com

MarcelaPeralta [Chapter23:ClassMalacostraca]InstitutodeInvertebrados,Fundacio ´ nMiguelLillo,San MigueldeTucuma ´ n,Argentina;email: maperalta@ csnat.unt.edu.ar

GilmarPerbiche-Neves [Chapter21:ClassCopepoda] DepartamentodeHidrobiologia,UniversidadeFederal deSaoCarlos,SP,Brazil;email: gilmarperbi Chaptere83@gmail.com

UlissesPinheiro [Chapter3:PhylumPorifera]DepartmentodeZoologiaUniversidadeFederaldePernambuco,Recife,Brazil;email: uspinheiro@hotmail.com

FabioBettiniPitombo [Chapter18:ClassCirripedia] DepartamentodeBiologiaMarinha,Universidade FederalFluminense,Nitero ´ i,RJ,Brazil;email: fpitombo@gmail.com

Jean-PierrePointier [Chapter11:PhylumMollusca] LaboratoireEcosyste ` mesAquatiquesTropicauxet Me ´ diterrane ´ ens,Universite ´ dePerpignan,France; email: pointier@univ-perp.fr

GabrielPompozzi [Chapter16:PhylumArthropoda: IntroductionandArachnida]Seccio ´ nEntomologı´a, FacultaddeCiencias,UniversidaddelaRepu ´ blica, Montevideo,Uruguay;email: gabrielpompozzi@ gmail.com

DanielPreviattelli [Chapter21:ClassCopepoda]DepartamentodeZoologia,InstitutodeBiocie ˆ ncias,UniversidadedeSa ˜ oPaulo,Sa ˜ oPaulo,SP,Brazil;email: dpreviattelli@gmail.com

W.WaynePrice [Chapter23:ClassMalacostraca] DepartmentofBiology,UniversityofTampa,Tampa, Florida,USA;email: wprice@ut.edu

ThiagoQuinta ˜ oArau ´ jo [Chapter7:PhylumGastrotricha] Laborato ´ riodeEvoluc¸a ˜ odeOrganismosMeiofaunais, DepartamentodeBiologiaAnimal,InstitutodeBiologia,UniversidadeEstadualdeCampinas,Campinas, SaoPaulo,Brazil;email: araujotq@gmail.com

LorenaRebecchi [Chapter15:PhylumTardigrada]DepartmentofLifeSciences,UniversityofModenaand ReggioEmilia,Italy;email: lorena.rebecchi@unimore.it

CarlosE.F.daRocha [Chapter21:ClassCopepoda] DepartamentodeZoologia,InstitutodeBiocie ˆ ncias, UniversidadedeSaoPaulo,SaoPaulo,SP,Brazil; email: cefrocha@usp.br

D.ChristopherRogers [Chapter1:Introduction,Chapter 16:PhylumArthropoda:IntroductionandArachnida, Chapter17:PhylumArthropoda:IntroductiontoCrustaceaandHexapoda,Chapter19:ClassBranchiopoda, Chapter23:ClassMalacostraca]KansasBiological Surveyand,TheBiodiversityInstitute,UniversityofKansas,Lawrence,KS,USA;email: branchiopod@gmail.com

ErichHarryRudolphLatorre [Chapter23:ClassMalacostraca]DepartamentodeCienciasBiolo ´ gicasyBiodiversidad,UniversidaddeLosLagos,Osorno,Chile; email: erudolph@ulagos.cl

AnaMarı´aSantana-Pineros [Chapter21ClassCopepoda] FacultaddeCienciasVeterinarias,UniversidadTe ´ cnica deManabı´,Ecuador;email: anasantana4@gmail.com

SandroSantos [Chapter23:ClassMalacostraca]DepartamentodeEcologiaeEvoluc¸a ˜ o,UniversidadeFederal deSantaMaria,SantaMaria,RS,Brazil;email: sandro.santos30@gmail.com

AndreasSchmidt-Rhaesa [Chapter10:PhylumNematomorpha]ZoologicalMuseum,UniversityHamburg, Hamburg,Germany;email: andreas.schmidt-rhaesa@ uni-hamburg.de

Andre ´ RinaldoSennaGarraffoni [Chapter7:Phylum Gastrotricha]Laborato ´ riodeEvoluc¸a ˜ odeOrganismos Meiofaunais,DepartamentodeBiologiaAnimal,InstitutodeBiologia,UniversidadeEstadualdeCampinas, Sa ˜ oPaulo,Brazil;email: arsg@unicamp.br

FerrySiemensma [Chapter2:Protozoa]Julianaweg10, 1241VWKortenhoef,TheNetherlands;email: ferry@ arcella.nl

PedroHenriquedaSilvaConceic¸a ˜ o [Chapter16: PhylumArthropoda:IntroductionandArachnida] DepartamentodeZoologia,UniversidadeFederalde MinasGerais,BeloHorizonte,Brazil;email: pedro. katekyo@gmail.com

ThiagodaSilvaPaiva [Chapter2:Protozoa]Laborato ´ rio deProtistologia,DepartamentodeZoologia,Instituto deBiologia,CCS,UniversidadeFederaldoRiode Janeiro,RJ,Brazil;email: tpaiva@biologia.ufrj.br

Se ´ rgioN.Stampar [Chapter4:PhylumCnidaria]DepartamentodeCie ˆ nciasBiolo ´ gicas,UniversidadeEstadual Paulista(UNESP),FCL/CampusdeAssis,SaoPaulo, Brazil;email: stampar@assis.unesp.br

MalinStrand [Chapter6:PhylumNemertea]Swedish SpeciesInformationCentre,SwedishUniversityof AgriculturalSciences,Uppsala,Sweden;email: malin.strand@slu.se

EduardoSua ´ rez-Morales [Chapter21:ClassCopepoda, Chapter22:ClassBranchiura]ElColegiodelaFronteraSur,Chetumal,Mexico;email: esuarez@ecosur.mx

PerSundberg [Chapter6:PhylumNemertea]Department ofMarineSciences,UniversityofGothenburg,Gothenburg,Sweden;email: per.sundberg@marine.gu.se

JamesH.Thorp [Chapter1:Introduction;Chapter17: PhylumArthropoda:IntroductiontoCrustaceaand Hexapoda]KansasBiologicalSurveyandDepartment ofEcologyandEvolutionaryBiology,Universityof Kansas,USA;email: thorp@ku.edu

TarmoTimm [Chapter12:PhylumAnnelida]Estonian UniversityofLifeSciences,CentreforLimnology,Tartumaa,Estonia;email: Tarmo.Timm@emu.ee

AntonioAlejandroVa ´ zquez [Chapter11:PhylumMollusca]LaboratoryofMalacology,InstituteofTropical MedicinePedroKourı´,Havana,Cuba;email: antonivp@ipk.sld.cu

RobertWallace [Chapter8:PhylumRotifera]Department ofBiology,RiponCollege,Ripon,WI,USA;email: WallaceR@ripon.edu

TimothyS.Wood [Chapter13:PhylumBryozoa,Chapter 14:PhylumEntoprocta]DepartmentofBiological Sciences,WrightStateUniversity,Dayton,Ohio, USA;email: tim.wood@wright.edu

FernandaZanca [Chapter10:PhylumNematomorpha] CentrodeEstudiosParasitolo ´ gicosydeVectores (CEPAVE,CONICET-UNLP),LaPlata,Argentina; email: fmzanca@fcnym.unlp.edu.ar

AbouttheEditors

Dr.M.CristinaDamborenea isaresearchzoologistatthe ConsejoNacionaldeInvestigacionesCientificas&Te ´ cnicas(Argentina),CuratorofInvertebrateCollectionsof MuseodeLaPlata(Argentina),andProfessorofInvertebrateZoologyatFacultaddeCienciasNaturalesyMuseo, LaPlataUniversity(Argentina).Shenotonlyspecializesin freelivingPlatyhelminthesbutalsostudiesothernoninsect groupsofaquaticinvertebratesaswellasaquaticinvasive speciesinSouthAmerica.CristinaDamboreneacreated andleadsaresearchgroupinfreelivingPlatyhelminthesof theNeotropicalregion.Shehasnumerouspeer-reviewed publicationsfocusedonthetaxonomyandecologyof invertebrates,invasivemollusks,scientificfieldguides,and studiesofthedisseminationofscientificknowledge.

Dr.D.ChristopherRogers isaresearchzoologistatthe UniversityofKansaswiththeKansasBiologicalSurvey andisaffiliatedwiththeBiodiversityInstitute,with numerousresearchprojectsallovertheworld.Hereceived hisPhDdegreefromtheUniversityofNewEnglandin Armidale,NSW,Australia.Christopherspecializesin freshwaterandterrestrialcrustaceans(particularlyBranchiopodaandMalacostraca)andtheinvertebratefaunaof seasonallyastaticwetlandsonaglobalscale.Hehas morethan150peer-reviewedpublicationsincrustacean taxonomyandinvertebrateecology,aswellaspublished popularandscientificfieldguidesandidentification manualstofreshwaterinvertebrates.Christopherisan AssociateEditorfortheJournalofCrustaceanBiology andafoundingmemberoftheSouthwestAssociationof FreshwaterInvertebrateTaxonomists.Hehasbeen involvedinaquaticinvertebrateconservationeffortsall overtheworld.

Dr.JamesH.Thorp hasbeenaProfessorintheDepartment ofEcologyandEvolutionaryBiologyattheUniversityof Kansas(Lawrence,KS,USA)andaSeniorScientistinthe KansasBiologicalSurveysince2001.Priortoreturningto hisalmamater,ProfessorThorpwasaDistinguished ProfessorandDeanatClarksonUniversity,Department ChairandProfessorattheUniversityofLouisville,AssociateProfessorandDirectoroftheCalderEcologyCenterof FordhamUniversity,VisitingAssociateProfessoratCornell, andResearchEcologistattheUniversityofGeorgia’s

SavannahRiverEcologyLaboratory.Hereceivedhis BaccalaureatefromtheUniversityofKansas(KU)andboth MastersandPhDdegreesfromNorthCarolinaState.Those degreesfocusedonzoology,ecology,andmarinebiology, withanemphasisontheecologyoffreshwaterandmarine invertebrates.Dr.Thorphasbeenontheeditorialboardof threefreshwaterjournalsandisaformerPresidentofthe InternationalSocietyforRiverScience.Heteachesfreshwater,marine,andinvertebratecoursesatKU,andhis MasterandDoctoralgraduatestudentsworkonvarious aspectsoftheecologyofcommunitiesthroughmacrosystemsinrivers,reservoirs,andwetlands.Professor Thorp’sresearchinterestsandbackgroundarehighlydiverse andspanthegamutfromorganismalbiologytocommunity, ecosystem,andmacrosystemecology.Heworksonboth fundamentalandappliedresearchtopicsusingdescriptive, experimental,andmodelingapproachesinthefieldandlab. Whilehisresearchemphasizesaquaticinvertebrates,healso studiesfishecology,especiallyasrelatedtofoodwebs.He haspublishedmorethan130refereedjournalarticles,books, andchapters,includingthreesingle-volumeeditionsof EcologyandClassificationofNorthAmericanFreshwater Invertebrates (editedbyJ.H.ThorpandA.P.Covich)and fivevolumesinthecurrentfourtheditionof Thorpand Covich’sFreshwaterInvertebrates

Thosereadersfamiliarwiththefirstthreeeditionsofour invertebratebook(EcologyandClassificationofNorth AmericanFreshwaterInvertebrates,editedbyJ.H.Thorp andA.P.Covich)willnotethatthefourtheditionhas expandedfromaNorthAmericanfocustoworldwide coverageofinlandwaterinvertebrates.Wegaveourbook seriesoninlandwaterinvertebratesthename Thorpand Covich’sFreshwaterInvertebrates to:(a)associatepresent withpasteditions,unitecurrentvolumes,andlinktofuture editions;(b)establishaconnectionbetweentheecological andgeneralbiologycoverageinVolumeIwiththetaxonomickeysintheremainingvolumes;and(c)givecreditto ProfessorAlanCovichforhisworkonthefirstthree editions.Forthesakeofbrevity,werefertothecurrent editionasT&CV.WhetherthefiftheditionofT&Cwill everappeariscertainlyproblematic,butwhoknows?At present,weareconsideringproducingupto11volumesin thefourthedition.

WhileIamthesoleserieseditoratthispoint,Christopherhasbeenamajorandhighlyvaluedpartnerindevelopingideasforthefourtheditionandhasbeenacoeditoron allﬁvevolumesnowinprint(senioreditoronVolumeIV). Hewillalsolikelyplayamajorroleinmanyofthe remainingvolumesbecauseofhisdiverseandglobal knowledgeoffreshwaterinvertebrates,especiallyinthe areaoftaxonomy.Inthenextdecade,wehopetopublish 4 6morevolumes,withanAustralasianvolumelikelyto appearin2021.Wearestillseekingexpertsinﬁeldsof invertebratetaxonomyforvariouszoogeographicregions toserveashighlydependablecoeditors,especiallythose whobothworkandliveinthezoogeographicregions coveredbythevariousfuturevolumes.

PrefacetotheFourthEdition

OurconceptforT&CIVincludedproducingonebook (VolumeI,publishedin2015)withsixchaptersongeneral environmentalissuesapplicabletomanyinvertebrates, followedby35chaptersdevotedtoindividualtaxaat variouslevels(ordertophylum,orevenmultiplephylain thecaseoftheprotozoa).VolumeIwasdesignedbothasan independentbookonecologyandgeneralbiologyofthe variousfreshwaterinvertebratetaxaandasacompanion volumeforusersofthekeysintheregionaltaxonomic volumes,therebyreducingtheamountofinformation duplicatedinthetaxonomicvolumes.Alltaxonomic volumestobepublishedoverthenextdecadewillcontain bothkeysforidentifyinginvertebratesinspeciﬁczoogeographicregionsanddescriptionsofdetailedanatomical featuresneededtoemploythosekeys.

WhilethevastmajorityofauthorsinvolumesofT&C editionsI IIIwerefromtheUnitedStatesorCanada,we attemptedinT&CIVtoattractauthorsfrommanyadditionalcountriesinothercontinents.Althoughwelargely succeededinthisgoal,weexpecttheﬁftheditionof T&C ifitiseverpublished tocontinueincreasingthe proportionofauthorsfromoutsideNorthAmericaasour booksbecomebetterknowninternationally.

OurgoalsforT&CIVaretoimprovethestateoftaxonomicandecologicalknowledgeofinlandwaterinvertebrates,tohelpprotectouraquaticbiodiversity,andto encouragemorestudentstodevotetheircareerstoworking withthesefascinatingorganisms.Thesegoalsareespecially importantbecausetheveriﬁedandprobablefuturelossesof speciesinwetlands,ponds,lakes,creeks,andriversaround theglobeexceedthoseinmostterrestrialhabitats.

JamesH.Thorp

Thisistheﬁfthvolumeofthefourtheditionof Thorpand Covich’sFreshwaterInvertebrates (T&CIV).Information ontheecologyandgeneralbiologyofthegroupscanbe foundinVolumeI(EcologyandGeneralBiology,editedby ThorpandRogers,2015),thecompaniontextforthe currentandallremainingbooksinthisseries.Alltaxonomicvolumes(otherthanthosefocusedexclusivelyon Hexapoda)areexpectedtoconsistofanintroductory chapter,achapteronprotozoa(multiplekingdoms),and multiplechaptersonindividualphylafromCnidariato Arthropoda.Someofthechaptersareverysmall (e.g.,Chapter14onEntoprocta),whereasothersarevery large(especiallyChapter23onMalacostraca).

Atypicalchapterincludesashortintroduction,abrief discussionoflimitstoidentiﬁcationoftaxainthatchapter, importantinformationonterminologyandmorphologythat isneededtousethekeys,techniquesforpreparingand preservingmaterialforidentiﬁcation(alsocoveredin VolumeI),thetaxonomickeys,andafewreferences.In largechapters,suchasthoseonMollusca(11),Annelida (12),andArthropoda(16 23),differentindividualshave contributedseparatesections,andthustherearemultiple sectionsonintroductionthroughkeysandreferences. Whilethismayconfusesomereaders,ithasallowedusto gaincontributionsfromanincreasednumberofexperts aroundtheworld.

Themultilevelkeysareformattedtoenableusersto workeasilyattheleveloftheirtaxonomicexpertiseandthe needsoftheirproject.Forthatreason,weseparatedkeysby majortaxonomicdivisions.Forexample,astudentin acollegecoursemightworkthroughoneormoreofthe initialcrustaceankeystodeterminethefamilyinwhich afreshwatershrimpbelongs.Incontrast,someoneworking onanenvironmentalmonitoringprojectmightneedto identifyasnailorcrabtogenusorevenspecies,andthus wouldusetherelevant,detailedkeysthatrequiremore backgroundexperience.

Wehaveaskedauthorstoincludeonlytaxathatare recognizedinternationallybypublicationinreputable

scientificjournalsthatfollowtheInternationalCodeof ZoologicalNomenclature.Thus,notaxathathavemerely beenproposedshouldhavebeenincludedeveniftheyhave beenidentifiedbytheworld’sexpertonthatgroup. “Common”speciesarenotdesignatedinthekeysthemselvesbecauseacommonspeciesinoneareamaynotbe commoninanother,andthisdesignationcanleadtooverly frequentandfalseidentifications.Authorshavebeen encouragedtoendthekeysatthepointwherefurther identificationwithoutgeneticanalysisisnotpracticalor whenitisclearthattoomanyoftheextantfaunahaveyetto bedescribedinscientificpublications.

Usersofthesekeysneedtorealizethattaxonomyis agrowingandvibrantﬁeldinwhichnewtaxaarebeing describedandpreviouslyacceptedrelationshipsreevaluated.Forsomeusers,thisvolumemaybesufﬁcientfor theirneeds,butforothers,acompaniontextlistingknown speciesinasmallergeographicregionmayalsobehelpful.

Thiseditionisstronglyfocusedonspeciesfoundin freshthroughsalineinlandwaters,withanonexclusive emphasisonsurfacewaters,therebyreﬂectingthebiasof existingscientiﬁcliterature.Again,mostestuarineand parasiticspeciesarenotcoveredinthisbook,butwedo discussspecieswhoselifecycleincludesafree-livingstage (e.g.,Nematomorpha)andspeciesthatliveinhardfreshwatersthroughtobrackishwaterseventhoughtheymaybe normallyassociatedwithestuarineormarinehabitatsin somepartsoftheirlifecycles(e.g.,someshrimpandcrabs).

Itisourhopethatscientistsandstudentsfromaround theworldwillbeneﬁtfromthisvolume,especiallythose studyingorganismsintheNeotropics.Suggestionsfor improvingfuturevolumesarewelcome.

Editors

Manypeoplecontributedtothisvolumeinadditiontothe chapterauthorsandthoseacknowledgedinindividual chapters.Wegreatlyappreciateallourcolleagueswhohave contributedinformation,ﬁgures,orreviewstoVolumeV andalsothankthosewhoprovidedsimilarservicesforthe earliereditions,uponwhichthepresentbookpartially relies.Weareagaingratefultothehighlycompetentpeople atAcademicPress/Elsevierwhohelpedinmanyaspectsof thebook’sproductionfromtheoriginalconcepttotheﬁnal

AcknowledgmentsforVolumeV

marketing.Inparticular,weappreciateourassociationwith ElseviereditorsincludingEmilyThomsonandLouisa HutchinsandearlierLauraKelleher,aswellastheentire globalproductionteamsforElsevier,especiallythoseled byPaulPrasadChandramohan.

Introduction1

JamesH.Thorp

KansasBiologicalSurvey,Lawrence,KS,UnitedStates,DepartmentofEcologyandEvolutionaryBiology,UniversityofKansas,Lawrence,KS,United States

D.ChristopherRogers

KansasBiologicalSurvey,Lawrence,KS,UnitedStates,TheBiodiversityInstitute,UniversityofKansas,Lawrence,KS,UnitedStates

CristinaDamborenea

Divisio´nZoologı´aInvertebrados,MuseodeLaPlata,FCNyM-UNLP,BuenosAires,Argentina.CONICET

ChapterOutline

IntroductiontothisVolumeandChapter11 NatureandFaunaoftheNeotropicsandAntarctica2 FaunaofAntarcticaandNearbyIslands 2 ComponentsofTaxonomicChapters2

INTRODUCTIONTOTHISVOLUMEAND CHAPTER1

Thefirstvolumeof ThorpandCovich’sFreshwaterInvertebrates (Thorp&Rogers,2015)providedglobal coverageoftheecology,generalbiology,phylogeny,and collectiontechniquesforinlandwaterinvertebrates.The focusofallvolumesinourseriesafterVolumeIhasbeenor willbeontaxonomickeysforindividualbioregionsofthe world.Thetaxonomickeysprovideuserswithmeansto workattheirlevelofneedandexpertise.Ourauthorshave triedtolimitthespecificityofthekeystothosetaxawhere comprehensiveknowledgeisavailableformosttaxaina particulargrouporwhereidentificationisonlypossiblefor certainlifestages.Itisourhopethatthesebooksdemonstratethelimitsofourknowledgeonthebiodiversityfor differentregionsandtaxa,andthuswillinspirefuture studiesinthoseareas.

Inadditiontotheﬁrstﬁvevolumescurrentlyinprint (Thorp&Rogers,2015,2016;Hamadaetal.,2018;

1Thischapterwaswrittentobeausefulstartingpointfortheprevious, current,andfuturetaxonomicvolumesinallzoogeographicregions. Consequently,therewillbeonlymoderatedifferencesinthischapter amongvariousvolumes.

HowtoUseThisVolume2 KeytoKingdomsandPhylainThisVolume5 References7

Rogers&Thorp,2019,andthecurrentvolume),asixth volumeshouldappearin20 21(KeystoAustralasian Fauna; Ahyongetal.,2021)andworkhasrecently commencedonaseventhvolume(KeystoAfrotropical Fauna;Dayetal.,2024?).Thegeneralinvertebratesand theHexapodaoftheOriental/Oceanabioregionsareexpectedtobecoveredintwofuturevolumes,alongwith moredetailedcoverageofHexapodaintheNearcticand Palaearcticbioregions.

TheNeotropicalregionasdeﬁnedinthisvolumemay havefaunaloverlapwiththeNearcticfaunaintheBaja PeninsulaattheTropicofCancer,intheMexicanLowlands,andthroughthesouthernFloridaPeninsula.For specimenscollectedadjacenttotheseregionsthatdonot closelycorrespondwiththekeyspresentedherein,please consultVolumeIIofthisbookseries.Notealsothatthe currentvolumetreatsinsectsonlytoorderbecausegreater detailappearedinVol.III:KeystoNeotropicalHexapoda (Hamadaetal.,2018).

Weconsiderthisseriesof10 12volumestobea globallyexpandedfourtheditionoftheearlier Ecologyand ClassiﬁcationofNorthAmericanFreshwaterInvertebrates (editedbyThorp&Covichin1991,2001,and2010). Whilesomelabsmayhavemultiplecopiesofthe“Keysto Fauna”intheirregion,wealsorecommendthattheyhave

atleastonecopyofVolumeItoobtainusefulbackground informationoneachinvertebrategroup,includingglobal coverageoftheecology,generalbiology,phylogeny,and collectiontechniquesforinlandwaterinvertebrates.

Thecurrentchapterisorganizedintoanintroduction,a descriptionofthetaxonomicfocusofthisparticularvolume,asectionexplainingtheorganizationofmosttaxonomicchapters,andakeytolargertaxonomicgroups.This chapter’skeyisdesignedtohelpthereaderlocatethemost pertinentchapter(ataskprobablyusefulmostlyforstudentsandrelativelynovicetaxonomists)andbeginidentifyingorganismsintheirsamples.Readerswillnotethat chapterswithinandamongvolumesvaryinthespecificity oftheirtaxonomickeys.Thisreflectsboththelikely percentofthefaunathathasbeennamedandhoweasily taxacanbeseparatedbyalphataxonomicmethodsand associatedkeys.Whilethereareoccasionalreferencesto geneticseparationoftaxa,thekeysareprimarilybasedon microscopeidentifications.

NATUREANDFAUNAOFTHE NEOTROPICSANDANTARCTICA

Asisclearfromthetitle,thisvolumecoversinlandwater invertebratesofboththeNeotropicsandAntarctica,even thoughtheformerbioregionwilllikelyrepresentover 99.9%ofthetaxaidentiﬁedinthisvolume.Antarctica neededtobecoveredinoneofourvolumes,however,and wechosetoputitherebecausemanyinlandwaterspecies onAntarcticaandsurroundingislandsoccurinthesouthern portionoftheNeotropicalregion,withafewalsopresentin thesoutherntipofAfricaandAustralia.Wehaveasked authorstoincludeanyspeciesintheirkeysthathavebeen identiﬁedfromAntarcticaandtoindicatetheirgeneral location.AmapofthegeneralregioncoveredbyAntarctica andnearbypolar-typeenvironmentsappearsas Fig.1.1

Forsomereaders,especiallythosefromtemperate zones,“neotropical”maysuggestawarmclimate.Itis importanttorecognize,however,thattheNeotropical bioregionstretchesfromthemiddleofMexico(warm) throughtheequatorialcountries(oftenhot)tothefrigid areasofTierradelFuego(Argentina)andtheislandsof CapeHorn(Chile).Italsoincludesbothlowandhigh altitudesiteswhichvarygreatlyintheirtemperaturerange. Moreover,someregionsarehotorcolddesertsandothers rangeuptoveryhumidenvironments.Thefreshwater invertebratefaunaofthesediverseregionswillvary accordingly.Manyofthesediversehabitatsaredescribedin Chapter2ofVolumeI.

FaunaofAntarcticaandNearbyIslands

WithinthegeneralAntarcticregioncoveredinthischapter areavarietyofmostlysmallislands(Fig.1.1),someof

whichhavemorereportedspeciesthanonthefrozen continent.Forexample,continentalAntarcticalackspermanentcolonizationofinsects,whereastheirdiversityon islandsrangefrom2to200species,mostofwhichare Diptera(flies)andColeoptera(beetles)(Chown&Convey, 2016).Withinaquaticsystems,zooplanktonspecies (e.g.,rotifersandcopepods)areabundantrelativetomost otheraquaticinvertebrates(otherthanprotozoa).While additionaltaxacontinuetobefound,scientistsfacethe problemofdeterminingwhethertheyarenativespeciesor newlyarrivedimmigrants(e.g., Hughes&Convey,2012). Table1.1 liststheknowncompositionofhighertaxaof inlandwaterinvertebratesreportedfromthemainlandand/or islandsnearAntarcticaandcoldSouthernOcean.Thislistis undoubtedlyincomplete,butitwillgivethereaderanideaof whichchaptersandkeystoexamineinthisvolumeinsearch ofspecifictaxainthesesouthernpolarhabitatsandsomeof thescientificliteratureonthesecoldwaterinvertebrates.

COMPONENTSOFTAXONOMIC CHAPTERS

Thisvolumeisanidentificationmanualtotheinlandwater invertebratesoftheNeotropicalandAntarcticregionswhere wepresentinformationneededtodiagnoseanddetermine theseorganismstovarioustaxonomiclevels.Otherinformationconcerningecology,morphology,physiology,phylogeny,andbothcollectingandculturingtechniquescanbe foundinVolumeIofthisseries.Allremainingchaptersin thecurrentvolumearelimitedtoasinglephylumorclass, exceptforChapter2’scoverageofmultiplephylaofunicellularprotists.Chapter2isdesignedforreaderswhoonly needgeneralinformationaboutprotists.Wehaveattempted toincludethefollowingfivesectionsinthosechapters:(1)a briefoverviewofthehigherleveltaxon;(2)adescriptionof identificationlimitationsforeachtaxon;(3)detailsof pertinentterminologyandmorphology;(4)informationon preparingandpreservingspecimensforidentification;and (5)taxonomickeys(separatedbylevelofidentification).A restrictednumberofespeciallypertinentreferencesare givenineachchapterfollowingappropriatetaxonomic sections.Readerscanfindamuchmoreextensivelistof referencestotheirgroupinVolumeI(Chapters3and7 41) alongwithmoredetailsoncollecting,preparation,and preservingmajortaxa.Figuresineachchapterarelimitedto thoseneededforeffectiveuseofthekeys.Foradditional anatomicalinformation,includingfigures,seetherelevant chapterinVolumeI.

HOWTOUSETHISVOLUME

Thereisanoldmaximthatsays“keysarewrittenby peoplewhodonotneedthemforpeoplewhocannotuse

FIGURE1.1 MapofAntarcticaandsurroundingislandsandcontinents.Themapshowstheapproximatelocationsofanumberofislandswhere collectionofinlandwaterinvertebrateshavetakenplace. MapbasedonavarietyofsourcesandproducedbyNicholasKotlinksi.

them.”Wehavemadeeveryefforttomakethesekeysas user-friendlyaspublicationlimitationswouldpermit.

Eachchapterincludesabasicintroductiontothe morphologyandterminologyusedindiagnosingthetaxaof thatsection.Limitationstothecurrentstateoftaxonomic knowledgearealsopresentedsothatthereadermaygauge thereliabilityoftheinformationpresented.Onlythe established,peerreviewedscientificliteratureisusedto definethetaxonomiccategoriesandepithetsincluded.All names,asfarasweareaware,conformtotheInternational CodeofZoologicalNomenclature.Allnominaandtaxonomicarrangementsused,aswellastherejectionofold names,wasbasedonpeerreviewedscientificliterature. Namesfromunpublishedmanuscripts,dissertations,“in

house”designations,orrecordsthathavenotbeenvalidated arenotacceptable.Provisionalnamesandspeciesdesignated“taxon1”or“species1”werenotusedunlessthey werepreviouslyrecognizedandacceptedinthepeer reviewedscientiﬁcliterature(Richards&Rogers,2011). Nonewspeciesdescriptionsorpreviouslyunpublished taxonomicarrangementsarepresented.

Thekeysaredichotomous(notripletsorquadrupletsare used)andarehierarchical.Thus,foragivengroup,theﬁrst keysaretothehighesttaxonomiccategory.Thesecondset ofkeysistothenextlevel,thethirdsettothelevelbelow thatone,andsoon,downtothelowestjustiﬁabletaxonomiclevelbasedoncurrentknowledgeofthatgroup.This levelisdifferentfordifferentgroupsdependinguponthe

TABLE1.1 CurrentlyReportedInvertebrateTaxaFoundonAntarcticaandSurroundingIslands

TaxonReportedLocationCommentsandRecentReferences

GeneralinvertebratesISL

Downieetal.(2000);Dartnall(2003,2005);Barber-James(2007);Coulson (2007);Dartnall&Smith(2012);Velasco-Castrillonetal.(2014)

KingdomProtistaANT,ISLLabculturesfromwatercollectedatfreshwatersites. Tongetal.(1997);Xuetal.(2016)

PhylumPlatyhelminthesISL

PhylumGastrotrichaISL

Ball&Hay(1977);Dartnall(1995,2005);Dartnalletal.(2005);Dartnall& Smith(2012)

Dartnall(1995,2005);Downieetal.(2000); Dartnall&Smith(2012)

PhylumRotiferaANT,ISLLowdiversityinAntarctica,butgroupisrelativelyabundantcomparedto othertaxaonsouthernoceanislands. Dartnall(1995,2005);Janiec(1996);Downieetal.(2000);Izaguirreetal. (2003);Dartnalletal.(2005);Camacho(2006);Segers(2008);Dartnall& Smith(2012);Fontanetoetal.(2015)

PhylumNematodaANT,ISL

PhylumAnnelidaANT,ISL

Dartnall(1995,2005);Janiec(1996);Downieetal.(2000);Dartnalletal. (2005);Andra ´ ssy&Gibson(2007);Dartnall&Smith(2012);Raymondetal. (2013)

Downieetal.(2000);Dartnall(2005);Dartnalletal.(2005);Rodriguez&Rico (2008);Dartnall&Smith(2012)

PhylumTardigradaANT,ISLIncludeslimnoterrestrialspecies.

Janiec(1996);Dartnall(1995,2005);McInnes&Pugh(1998);McInnesetal. (2001); Downieetal.(2000);Dartnall&Smith(2012)

PhylumArthropoda

SubphylumChelicerataISL

SubphylumCrustacea

ClassHexapoda

Pugh&Dartnall(1994);Dartnall(1995,2005);Dartnalletal.(2005);Dartnall &Smith(2012)

SubclassEntognathaQuestionableIce-freeareasonANT,butmayreallybeaccidentaloccurrencesofterrestrial species.

Dartnall&Smith(2012)

SubclassInsectaISLMostlymidgesanddivingbeetles;nonereportedfromAntarctica. Arnold&Convey(1998);Jonesetal.(2003);Dartnalletal.(2005);Chown& Marshall(2008);Chown&Convey(2016); ANT Allegruccietal.(2012)

ClassBranchiopodaANT,ISLMostlyCladocerabutoneanostracan. Janiec(1996);Dartnall(1995);Pughetal.(2002);Dartnalletal.(2005);Camacho(2006);Dartnall&Hollwedel(2007);Hawes(2009);Dartnall&Smith (2012)

ClassMaxillopodaANT Bayly(1994),Baylyetal.(2003) ISL Dartnall(1995,2005);Janiec(1996);Downieetal.(2000);Pughetal.(2002); Izaguirreetal.(2003);Camacho(2006);Dartnall&Smith(2012)

ClassOstracodaISL Pughetal.(2002);Dartnall(2005);Dartnall&Smith(2012)

ClassMalacostraca Pughetal.(2010)

OrderIsopodaISL Pughetal.(2002);Dartnalletal.(2005)

OrderAmphipodaISL Pughetal.(2002)

Note: Thislistisalmostcertainlyincompleteandwillneedtobeupdatedasmoreinlandwaterhabitatsareexploredinthisarea.Climatechangemayeventually leadtoadditionalcolonizationopportunitiesbyfreshwaterinvertebrates.Thisactionandongoingpossibilitiesforaerialcolonizationfromothercontinentsof sometaxamayeventuallyincreasethefreshwateranimaldiversityonthecontinentandsurroundingislands.Seefullcitationinthechapterreferences. ANT,presenceoncontinentalAntarcticaandadjacentislands; ISL,islandsdistantfromtheAntarcticcontinentbutwithintheSouthernOcean.

stateofresolutioninthescientiﬁcliterature.Organismsnot identiﬁablebeyondaparticulartaxonomiclevelareleftat thatlevel.

Properlypreparedkeystypicallyemployspeciﬁc, primary,diagnosticcharacters.Olderkeysoftenuse differentcharactersthanthemorerecentkeys.Thisshift inprimarycharactersresultsfromsystematistsand taxonomiststestingtheimportanceofcharacters.Theultimategoalofthesystematististohavethetaxonomic charactersusedinourkeysconvergeascloselyaspossible withbiologicalreality.Toanontaxonomist,theresulting processmayerroneouslyseemmerelytobe“lumping andsplitting”taxonomicgroupsratherthantheresult ofemployingthescientiﬁcmethodtorevealnatural relationships.

Surprisingly,manyusersdonotknowhowtointerpreta dichotomouskey,makingthefundamentalassumptionthata correctidentiﬁcationanswerisalwayspresentinthekey.This assumptiongenerallytakesoneofthefollowingthreeforms:

1. Allspeciesareidentiﬁableusingagivenkey. Manynew specieshaveyettobedescribed,letalonediscovered. Generalizedgeographicrangesareprovidedformost taxapresentedherein,yetspeciesrangesshrink, swell,andchangeelevationconstantly,particularlyas weatherandclimatepatternsshift.Speciesdisperse, colonize,andsufferstochasticlocalextinctions.In additiontothesenaturalprocesses,somespeciesare introducedintentionallyoraccidentallybyhumans,and sometimestheirestablishmentallowsotherspeciesto invadeaswell.Furthermore,someregionsarebetter studiedthanothers:manypartsoftheworldhavenot beenexplored,northefaunadescribed.Therearestill plentyofnewtaxatoencounter.

2. Allvariationisaccountedforinthekey. Asstated above,identificationkeysusespecific,primary, diagnosticcharacters.Problemsinidentificationare compoundedbytaxathat:(a)havedifferentcharacter statesatdifferenttimes;(b)onlyhavediagnosticcharactersatcertainlifestagesorincertaingenders;and/or (c)haveseverelytruncatedmorphology(oftendueto lackofsexualselection)andlackmorphologicalcharacterstoseparatethespecies.Furthermore,newvaria-

tionwithintaxaiscontinuallydeveloping,andthusone cannotassumethatspeciesareimmutableordevelop toolspredictingthosechanges.

3. Thekeyisasufficientidentificationtoolinandofitself. A keyisjustatool.Thefactthatonehasaboltthatneeds removingandawrenchofthecorrectsizedoesnot meanthattheboltcanbeloosened.Similarly,identificationkeysaretoolstoaidintaxonidentification.They areprimarilytoolstoeliminateincorrecttaxafromthe rangeofpossiblechoices,narrowingthefieldtothe namesthatmaybeapplicable.Keysaretheprocessof elimination.Thepossibilitythatthespecimentobe identifiedisnew,ahybrid,anomalous,orarecent invasivecolonistisalwaysapossibleanswer.Thisis fundamentaltousinganyidentificationkey.

Onceonearrivesatanameorgroupofpossiblenamesfor aspecimeninhand,thespecimenshouldthenbe comparedagainstdescriptions,distributionmaps,and figuresofthatandothertaxainthatgroup.The descriptions,figures,andmapsareothertoolstobeusedin identification.Directcomparisonofthespecimenathand withidentifiedmuseummaterialorusingmolecular comparisonsisalsosometimesnecessaryforacorrect identification.

Inmanychaptersyouwilldiscoverthatmanyor sometimesmostofthespeciesinagiventaxonhavenot beenidentifiedorthatmostoftheidentificationsarefrom onecountryorregion.Inthosecasesitisextremely importantthatyoudonotselectaspeciesnamefromthe keyunlessallthedescriptionsfitthatspecimen.Instead, youshouldonlyselectthenexthighertaxon(e.g.,genus insteadofspeciesorsubfamilyinsteadofgenus).Todo otherwisewillintroduceavoidableerrorsintoyourproject andpossiblyintothescientificliterature.

Speciesarenotimmutable,fixedinlocationandform. Theychangeconstantlyandwillcontinuetodoso,confoundingkeysandanyotheridentificationmethod,suchas traittables,charactermatrices,orevengeneticanalyses. Thisiswhybiologyisfarbehindphysicsinthedevelopmentofunifiedtheories:biologyisfarmorecomplexthan physics,asitinvolvesmoremutableinteractingpartsand processes.

KEYTOKINGDOMSANDPHYLAINTHISVOLUME

Amajorchangeintheidentificationkeysforourfourtheditionhasbeentoincludemultiplekeysperchapterthatgenerallystartwithaclasslevelkey and proceedtofinerandfinerdivisions.Theseallowuserstoworkattheirlevelsofinterest,need,andskillwithouthavingtowadethroughextraneoustaxanot inthedirectlinetothetaxonofinterest.

ThefollowingkeywasderivedinpartfromChapter1inVolumeIofthefourthedition.Itismeanttoallowyoutomovetothenextlevelofkeys, whichwillbeinindividualchapters.Theterm“kingdom”andthespeciﬁcnamesassociatedwithit(i.e.,AnimaliaandProtista)areusedherelooselyto designatecommongroupingsofphyla,butthesehavehighlyquestionabletaxonomicsigniﬁcance.

1Multicellular,heterotrophicorganismsasindividualsorcolonies(sometimeswithsymbioticautotrophs).......kingdomAnimalia.......2

10 Unicellular(oracellular)organismspresentasindividualsorcolonieswithnucleiirregularlyarranged;heterotrophicand/orautotrophic; multiplephylawithintheautotrophicprotozoaphyla...................................................................................kingdomProtista[Chapter2]

2(1)Radiallysymmetricorasymmetricorganismslivingindividuallyorincolonies.......................................................................................3

20 Individualsbilaterallysymmetric..................................................................................................................................................................4

3(2)Surfacenotporous;oraltentaclesalwayspresentaroundacloseablemouth;colonialorsingle,mostlysinglepolypforms(primarilyhydra) orrarelymedusoidform(freshwaterjellyﬁsh;adultswithasinglecentralbodycavityopeningtotheexteriorandsurroundedbycellular endoderm,acellularmesoglea,andcellularectoderm....................................................................................phylumCnidaria[Chapter4]

30 Surfaceporous;colonial;tentaclesabsent;noclosableoriﬁces;withoutdiscreteorgans;cellular-level(orincipienttissue-level)construction;variable,nondistinctcolonyshapes,includingencrusting,rounded,ordigitiformgrowthforms;skeletonofindividualsiliceous spiculesandacollagenmatrix;internalwatercanalsystem;maycontainsymbioticalgae;thesponges.....phylumPorifera[Chapter3]

4(2)Oralregionwithnumeroustentaclesorciliadistributedaroundthemouth;organismneverwitheversiblejawsandneververmiformas adult...............................................................................................................................................................................................................5

4Oralregionwithtwoornotentacles,ortentaclesbehindthemouth..........................................................................................................7

5(4)Oralregionwithtentacles,organismsingelatinoidsorbranchingcolonies................................................................................................6

50 Oralregionringedwithcilia,muscularpharynx(mastax)withcomplexsetofjaws;singlefreeswimming,orsemi-sessilelivingsinglyorin smallcolonies;wheelanimals,orrotifers........................................................................................................phylumRotifera[Chapter8]

6(5)Oraltentacles(thelophophore)ina“U”or“horseshoe”shapearoundmouth;anusopensoutsideoflophophore;colonialanimals,oftenin massivecoloniesattachedtohardsurfaces;truebryozoans.....................................................phylumEctoprocta(Bryozoa)[Chapter13]

60 Bothmouthandanusopenwithintheoraltentacles;individual(noncolonial)animalswithacalyxcontainingasinglewhorlof8 16 ciliatedtentacles.........................................................................................................................................phylumEntoprocta[Chapter14]

7(4)Notwiththecombinationofcharacteristicsdescribedbelow.....................................................................................................................8

70 Small(50 800 mm),spindle-ortenpin-shaped,ventrallyﬂattenedwithamoreorlessdistinctheadbearingsensorycilia;cuticle usuallyornamentedwithspinesorscalesofvariousshapes;posteriorofbodyoftenformedintoafurcawithdistaladhesivetubes; gastrotrichs..................................................................................................................................................phylumGastrotricha[Chapter7] 8(7)Anteriormouthandposterioranuspresent..................................................................................................................................................9

80 Flattenedorcylindrical,acoelomatewormswithonlyone,ventraldigestivetractopening;sometimeswithevidenthead;turbellarian ﬂatworms(commonlycalledplanaria,anonspeciﬁcandusuallyincorrectname)............................phylumPlatyhelminthes[Chapter5] 9(8)Vermiformornot,eversibleoralproboscisnotpresent,althougheversiblejawsorothermouthpartsmayoccur..................................10

90 Long,ﬂattened,unsegmentedwormswithaneversibleproboscis;ribbonworms......................................phylumNemertea[Chapter6] 10(9)Bodynotenclosedinasingle,spiraledshellorinahinged,bivalvedshell;orifabivalvedshellispresent,thenanimalhasjointedlegs.....11

100 Soft-bodiedcoelomateswhosevisceraiscovered(infreshwaterspecies)byasingleordual(hinged),hardcalcareousshell;withaventral muscularfoot;ﬂeshymantlecoversinternalorgans;snails,clams,andmussels.......................................phylumMollusca[Chapter11] 11(10)Segmentedlegsabsentinalllifestages;ifjawsarepresent,thenbodywithatleast15segments........................................................12

110 Adultsandmostlarvalstageswithlegs;iflarvaewithoutlegsorprolegs(someinsects),thencephalicregionwithpairedmandibles,or eversiblehead,alwayswith14orfewerbodysegments...........................................................................................................................14

12(11)Organismvermiform,notsegmented..........................................................................................................................................................13

120 Organismvermiformornot,bodysegmented..............................................................................................phylumAnnelida[Chapter12]

13(12)Bodycylindrical,usuallytaperingatbothends;cuticlewithoutcilia,oftenwithstriations,punctuations,minutebristles,etc.;1cmlong (exceptfamilyMermithidae, < 6cm);nematodes,roundworms.................................................................phylumNematoda[Chapter9]

130 Bodywithanteriortipnormallyobtuselyroundedorblunt,posteriortipmaybebi-ortrilobed;cuticleopaquetodarkbrownorblack,and epicuticleusuallycrisscrossedbyminutegrooves;lengthseveralcmto1m,width0.25 3mm;onlyadultswithfree-livingstage; hairwormsorhorsehairworms............................................................................................................phylumNematomorpha[Chapter10] 14(11)Fourpairsofclawed,nonjointedlegs;waterbears...................................................................................phylumTardigrada[Chapter15]

140 Adultsandmostlarvaewithjointedlegs,orlegslacking,ormoreorlessthanfourpairs....................phylumArthropoda[Chapter16]2

2Chapter16includesakeythatwillleadtosubsequentchapterscovering thisphylum.

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Protozoa

GabrielaC.Ku¨ppers,AnushKosakyan,FerrySiemensma,Marı´aCristinaClaps,ThiagodaSilvaPaiva,Leonardo D.Ferna´ndez,LucianaBurdman,ValentynaKrashevska,EnriqueLaraandCristinaDamborenea

ChapterOutline

IntroductiontoFree-LivingProtozoa10

Amoeba:Arcellinida:Hyalospheniidae:Genera 33

Amoeba:Rhizaria:FamiliesandEuglyphida IncertaeSedis 34 Rhizaria:Euglyphidae:Genera 34

Amoebae:Phyla 20

Amoeba:Amoebozoa:Classes 20

Amoebae:Amoebozoa:Archamoebae:Families 21

Amoebae:Amoebozoa:Tubulinea(¼ Lobosea):Orders 21

Amoebae:Amoebozoa:Tubulinea(¼ Lobosea): Euamoebida:Families 21

Amoebae:Amoebozoa:Tubulinea(¼ Lobosea): Euamoebida:Amoebidae:Genera 21

Amoebae:Amoebozoa:Tubulinea(¼ Lobosea): Euamoebida:Hartmannellidae:Genera 24

Amoebae:Amoebozoa:Tubulinea(¼ Lobosea): Leptomyxida:Families 24

Amoebae:Amoebozoa:Tubulinea(¼ Lobosea): Leptomyxida:Flabellulidae:Genera 24

Amoebae:Amoebozoa:Tubulinea(¼ Lobosea): Echinamoebida:Families 24

Amoebae:Amoebozoa:Discosea:Orders 24

Amoebae:Amoebozoa:Discosea:Thecamoebida:Families 25

Amoebae:Amoebozoa:Discosea:Himatismenida:Families 25

Amoebae:Amoebozoa:Discosea:Centramoebida:Families 25

Amoebae:Amoebozoa:Discosea:Glycostylida:Families 25

Amoebae:Amoebozoa:Discosea:Dermamoebida:Families 25

Amoebae:Amoebozoa:Variosea:Families 25

Amoeba:Percolozoa:Heterolobosea:Families 26

Amoeba:Arcellinida:Familiesand Incertaesedis 26

Amoeba:Arcellinida:Plagiopyxiidae:Genera 27

Amoeba:Arcellinida:Trigonopixidae:Genera 27

Ciliophora:Phyllopharyngea:Suctoria:Families 44

Ciliophora:Phyllopharyngea:Suctoria:Podophryidae: Genera 45

Ciliophora:Phyllopharyngea:Suctoria:Acinetidae:Genera 45

Ciliophora:Phyllopharyngea:Cyrtophoria:Families

Ciliophora:Phyllopharyngea:Cyrtophoria:Chilodonellidae: Genera 45

Ciliophora:Phyllopharyngea:Dysteriidae:Genera 45

Ciliophora:Spirotrichea:MajorGroups

Ciliophora:Spirotrichea:OligotrichidaandHalteriida: Families

Ciliophora:Spirotrichea:Halteriidae:Genera 47

Ciliophora:Spirotrichea:Strombidiidae:Genera 47

Ciliophora:Spirotrichea:Choreotrichida:Families 47

Ciliophora:Spirotrichea:Choreotrichida:Codonellidae: Genera 49

Ciliophora:Spirotrichea:Choreotrichida:Strobilidiidae: Genera 49

Ciliophora:Spirotrichea:Hypotricha:Families 49

Ciliophora:Spirotrichea:Hypotricha:Spiroﬁlidae:Genera 50

Ciliophora:Spirotrichea:Hypotricha:Holostichidae:Genera 50

Ciliophora:Spirotrichea:Hypotricha:Pseudourostylidae: Genera 50

Ciliophora:Spirotrichea:Hypotricha:Kahliellidae:Genera 50

Ciliophora:Spirotrichea:Hypotricha:Amphisiellidae: Genera 50

Ciliophora:Spirotrichea:Hypotricha:Oxytrichidae:Genera 50

Ciliophora:Spirotrichea:Euplota:Families 52

Ciliophora:Spirotrichea:Euplota:Uronychiidae:Genera 54

Ciliophora:Spirotrichea:Euplota:Euplotidae:Genera 54

Ciliophora:Oligohymenophorea:MajorGroups 54

Ciliophora:Oligohymenophorea:Peritrichia:Families 55

Ciliophora:Oligohymenophorea:Peritrichia:Trichodinidae: Genera 55

Ciliophora:Oligohymenophorea:Peritrichia:Astylozoidae: Genera 55

Ciliophora:Oligohymenophorea:Peritrichia:Vorticellidae: Genera 57

Ciliophora:Oligohymenophorea:Peritrichia:Epistylididae: Genera 57

Ciliophora:Oligohymenophorea:Peritrichia:Vaginicolidae: Genera 57

Ciliophora:Oligohymenophorea:Peniculia:Families 57

Ciliophora:Oligohymenophorea:Peniculia:Frontoniidae: Genera 59

Ciliophora:Oligohymenophorea:Scuticociliatia:Families 59

Ciliophora:Oligohymenophorea:Scuticociliatia: Cyclidiidae:Genera 59

Ciliophora:Oligohymenophorea:Scuticociliatia: Philasteridae:Genera 60

Ciliophora:Oligohymenophorea:Scuticociliatia: Cinetochilidae:Genera 60

Ciliophora:Oligohymenophorea:Scuticociliatia: Uronematidae:Genera 60

Ciliophora:Oligohymenophorea:Scuticociliatia: Loxocephalidae:Genera 60

Ciliophora:Oligohymenophorea:Hymenostomatia: Families 60

Ciliophora:Oligohymenophorea:Hymenostomatia: Turaniellidae:Genera 60

Ciliophora:Oligohymenophorea:Hymenostomatia: Glaucomidae:Genera 60

Ciliophora:Armophorea:Families 61

Ciliophora:Armophorea:Metopidae:Genera 61

Ciliophora:Heterotrichea:Families 61

IntroductiontoFree-Living Protozoa

LeonardoD.Ferna´ndez

CentrodeInvestigacio´nenRecursosNaturalesySustentabilidad (CIRENYS),UniversidadBernardoO’Higgins,Santiago,Chile

CristinaDamborenea

CONICET,Divisio´nZoologı´aInvertebrados,MuseodeLaPlata,La Plata,Argentina

Ciliophora:Heterotrichea:Folliculinidae:Genera 61

Ciliophora:Heterotrichea:Blepharismidae:Genera 61

Ciliophora:Heterotrichea:Climacostomidae:Genera 61

Ciliophora:Heterotrichea:Condylostomatidae:Genera 62

Ciliophora:Prostomatea:Families 62

Ciliophora:Prostomatea:Holophryidae:Genera 64

Ciliophora:Nassophorea:MajorGroups 64

Ciliophora:Nassophorea:Microthoracida:Families 64

Ciliophora:Nassophorea:Microthoracida:Microthoracidae: Genera 64

Ciliophora:Nassophorea:Microthoracida: Leptopharyngidae:Genera 64

Ciliophora:Nassophorea:Synhymeniida:Familiesand Genera 66

Ciliophora:Nassophorea:Nassulida:Families 66

Ciliophora:Karyorelictea:FamiliesandGenera 66

Ciliophora:Litostomatea:Families 66

Ciliophora:Litostomatea:Tracheliidae:Genera 67

Ciliophora:Litostomatea:Enchelyidae:Genera 68

Ciliophora:Litostomatea:Lacrymariidae:Genera 68

Ciliophora:Litostomatea:Trachelophyllidae:Genera 68

Ciliophora:Litostomatea:Litonotidae:Genera 68

Ciliophora:Litostomatea:Spathidiidae:Genera 68

Ciliophora:Litostomatea:Actinobolinidae:Genera 68

Ciliophora:Litostomatea:Mesodiniidae:Genera 70

Ciliophora:Litostomatea:Didiniidae:Genera 70

Ciliophora:Litostomatea:Acropisthiidae:Genera 70

Ciliophora:Plagiopylea:MajorGroups 70

Ciliophora:Plagiopylea:Plagiopylida:FamiliesandGenera 70

Ciliophora:Plagiopylea:Odontostomatida:Families 70

Ciliophora:Plagiopylea:Epalxellidae:Genera 70

Ciliophora:Plagiopylea:Mylestomatidae:Genera 71

Ciliophora:Colpodea:Families 71

Ciliophora:Colpodea:Bryometopidae:Genera 71 Acknowledgments71 References71 FlagellatedProtozoa74 Introduction74 KeytoFlagellatedProtozoa75 KeytoMajorGroups 75 References77

INTRODUCTION

Classificationofunicellulareukaryoteshasbeenhistorically underreview.In1677,AntonyvanLeeuwenhoekfirst describedtheseorganismsunderthenameof“animalcules.” Almostacenturylater,unicellulareukaryotesweregrouped togetherunderthenameof“Infusoria,”atermpossibly coinedbyMartinF.Ledermullerin1763(Kudo,1931). TheywerelaterincludedwithinProtozoa(i.e.,“earlyanimals”),ataxonoriginallycreatedasaclass(Goldfuss,1820) andthenelevatedtophylum(vonSiebold,1854)andfinally

tokingdom(Owen,1860).Aroundthesametimeofthe creationofthekingdomProtozoa,unicellulareukaryotes werealternatelyincludedwithinthekingdomProtoctista (i.e.,firstcreatedbeings)toformallyincludemixotrophic representativesandmulticellularorganismsthatdidnot behavelike“trueanimals”or“zoa”(Hogg,1860).Sixyears later,thekingdomProtista(“thefirstofall,primordial”)was createdtoincludeallthoseorganismsthatwereneitheranimalsnorplants,suchasbacteriaandunicellulareukaryotes (Haeckel,1866).Fromthen,untiltheendofthe20thcentury,kingdomsProtozoa,Protoctista,andProtistawere redefined,revalidated,and/orinvalidatedbyseveralresearchers.However,inthelightofcurrentmolecular phylogeneticevidence,thesekingdomsarenolonger acceptedbymostmodernprotistologists(Hausmannetal., 2003; Adletal.,2005,2012; O’Malleyetal.,2012;Geisen etal.,2017).Onlysomeclassicterms,suchas“protist,”are stillretainedtoinformallydescribeeukaryoteswithaunicellularleveloforganization,withoutcelldifferentiation intotissues(Adletal.,2005).Theterm“protist”isalso increasinglybeingusedbyprotistologistsworkingin freshwaterunicellulareukaryotesfromtheNeotropicalregion(e.g., Ferna ´ ndezetal.,2015; Schiaffinoetal.,2016).

Unicellulareukaryoteshavereceivedspecialattention inthepastdecades.Recentadvanceswroughtbymolecular phylogeneticstudiesintroducednewperspectivesinthe knowledgeoftheseorganisms.Termssuchasalgae(phototrophicprotists)andprotozoa(mostheterotrophicprotists,otherthanthosethatcloselyresemblefungi, predominantlynonﬁlamentousheterotrophicspecies)lost theirboundariesandhavenotaxonomicvalue.Theterm protistdescribeseukaryoteswithaunicellularleveloforganization,withoutcelldifferentiationintotissues.Alternatevegetativemorphologyrestrictedtosexual reproduction,quiescent,orresistantstages,suchascysts, mightoccur(Adletal.,2005).Free-livingprotozoaare unicellularandheterotrophicandmayalsobemixotrophic (occurrenceofphagotrophyandphototrophyinthesame organism).Theyusuallyfeedonbacteria,algae,andeven zooplankton,dependingontheirsize.

Free-livingprotozoaplayavitalecologicalrolein aquaticenvironment.Becauseoftheirhighsensitivityto environmentalchanges,theyhavebeenusedasbioindicatorsandbiomonitorsofpollution.Hydrologicalcycles,increasednutrients,physicalandchemicalchanges, andphytoplanktonbloomsinwaterbodiesinterfereinthe dynamicsofthiscommunityaffectingthetrophicnetwork.

Thischapter,asintheothervolumesoftheseries, providesasynthesisonthediversity,ecology,anddistributionoffree-livingprotozoafoundinfreshwater Neotropicalecosystems,suchasciliates,ﬂagellates,and sarcodines.Thetwolattergroupsconsistofanumberof largelyunrelatedevolutionarylineages.

LIMITATIONS

Free-livingprotozoaarediverseandabundantinfreshwater ecosystems.Thestudyoftheirdiversity,ecology,and geographicaldistributionisincipientcomparedwiththatof multicellularorganisms,particularlyintheNeotropical region(Ferna ´ ndezetal.,2016; Schiaffinoetal.,2016). Eventhoughtheyplaypivotalrolesinnaturalecosystems (Geisenetal.,2018),theirstudyhascontributedlittletothe knowledgeofecologyandevolutionarybiology,mainly becauseresearchonprotozoaexhibitssomelimitations (Fenchel,1987).Forinstance,mostfree-livingprotozoa exhibitmicroscopicsizes,makingtheirmorphological identificationandcountingdifficult.Often,speciesidentificationrequirestraininginspecificmicroscopyand stainingtechniques,skillsthatnotmanyresearchers managetodate.TheyalsosufferfromasignificantLinnean shortfall,duetotheglobaldeficitoftaxonomistsdedicated tothestudyofthedifferentgroupsthatmakeupProtozoa. Inaddition,thereisacomparativelypoorknowledgeon theirsystematics,mainlydueto:(1)lackofadequate methodsforthefixationandlong-termpreservation;(2) absenceoftypespecimensformostspecies;(3)lackof morphologicalfeaturesforspeciesdelimitation;(4)inadequatespeciesdescriptions;and(5)greatareasundersampled(e.g.,theNeotropics)(Estebanetal.,2015).

Accordingtotheaforementionedbackground,thekeys presentedhereinreachdifferentlevelsofidentiﬁcation, accordingtothegroups.

TERMINOLOGYANDMORPHOLOGY

Mostprotozoansaremicroscopic,usually5 250 mmlong. Somespecieshavethesizerangeofbacteriaandrarely exceeding6 7mm.Themorphologyisvaried,e.g.,single independentcellsthatmayormaynotbemotile,ﬁlamentousspecies,andcolonial(linkedbyacommonstalk,or sheath,orcytoplasmicextensions)andmultinucleateforms.

Theypossessdifferentlocomotorystructures:cilia, ﬂagella,orpseudopodia(ﬁngerlikecytoplasmicextensions).Anotherrelevantcharacteristicisthemembranethat allowstheexchangewiththeenvironment.Thismembrane maybepresentalone(nakedforms),butmanyprotozoa haveathickenedmembraneandformasemirigidorrigid pellicle.Theyalsocansecretescalesorashell-liketestor canaccumulateexternalparticles.

Thetypesofnutritionarevaried.Theboundariesbetweenautotrophsandheterotrophsarenotstrict:some heterotrophsretaintheirprey’splastids;otherheterotrophs formsymbioseswithphotosyntheticspecies;andmany photosyntheticspeciesarealsophagotrophicandosmotrophic.