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KeystoPalaearcticFauna

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

VolumesinPreparationforFuturePublication

KeystoNeotropicalandAntarcticFauna (2019)

KeystoAustralasianFauna (2020)

KeystoAfrotropicalFauna (2023)

PlannedFutureVolumes

KeystoOrientalandOceanaHexapoda

KeystoOrientalandOceanaFauna

KeystoPalaearcticHexapoda

KeystoNearcticHexapoda

RelatedPublications

FieldGuidetoFreshwaterInvertebratesofNorthAmerica

JamesH.ThorpandD.ChristopherRogers FirstEdition(2011)

KeystoPalaearcticFauna

FourthEdition

KansasBiologicalSurveyandTheBiodiversityInstitute UniversityofKansas,Lawrence,KS,USA JamesH.Thorp

KansasBiologicalSurveyandDepartmentofEcologyandEvolutionaryBiology UniversityofKansas,Lawrence,KS,USA

AcademicPressisanimprintofElsevier 125LondonWall,LondonEC2Y5AS,UnitedKingdom 525BStreet,Suite1650,SanDiego,CA92101,UnitedStates 50HampshireStreet,5thFloor,Cambridge,MA02139,UnitedStates TheBoulevard,LangfordLane,Kidlington,OxfordOX51GB,UnitedKingdom

Copyright 2019ElsevierInc.Allrightsreserved.

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ThisbookandtheindividualcontributionscontainedinitareprotectedundercopyrightbythePublisher(otherthanasmaybenoted herein).

Notices

Knowledgeandbestpracticeinthisfieldareconstantlychanging.Asnewresearchandexperiencebroadenourunderstanding, changesinresearchmethods,professionalpractices,ormedicaltreatmentmaybecomenecessary.

Practitionersandresearchersmustalwaysrelyontheirownexperienceandknowledgeinevaluatingandusinganyinformation, methods,compounds,orexperimentsdescribedherein.Inusingsuchinformationormethodstheyshouldbemindfuloftheirown safetyandthesafetyofothers,includingpartiesforwhomtheyhaveaprofessionalresponsibility.

Tothefullestextentofthelaw,neitherthePublishernortheauthors,contributors,oreditors,assumeanyliabilityforanyinjuryand/or damagetopersonsorpropertyasamatterofproductsliability,negligenceorotherwise,orfromanyuseoroperationofanymethods, products,instructions,orideascontainedinthematerialherein.

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CoverPhotographscourtesyof(fromtop): VadimTakhteev,RafaelAraujo,andGintautasSteiblys

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Toourauthorswhomadethisbookpossible.

D.ChristopherRogers

Tomylovingwifeofnearlyhalfacenturywhohastoleratedmyoccasional frustrationswithmyprofessionwhileshowinggreatinterestinmystudies, andtothestudentsworkinginmylaboverthedecadeswhohavenever failedtoinspireandchallengemewhilereinforcinginmymindthe truevalueofbeingaprofessor.

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ContributorstoVolumeIVxiii

AbouttheEditorsxvii

PrefacetotheFourthEditionxix

PrefacetoVolumeIVxxi

AcknowledgmentsforVolumeIVxxiii

1. Introduction

JamesH.Thorp,D.ChristopherRogers

IntroductiontoThisSeries,Volume, andChapter1

NatureandFaunaofthePalaearctic Bioregion2 ComponentsofTaxonomicChapters2 HowtoUseThisVolume2

KeystoMajorTaxaofInlandWater InvertebratesofthePalaearctic3 References4

PartI

2. Protozoa

AlanWarren,GenovevaF.Esteban

Introduction8 Limitations8

TerminologyandMorphology8

MaterialPreparationandPreservation17 Acknowledgments27

KeystoProtozoa28 References42

3. PhylumPorifera

RenataManconi,RobertoPronzato Introduction43 Limitations43

TerminologyandMorphology44 MaterialPreparationandPreservation45 Acknowledgments45 KeystoSpongillida45 References91

4. PhylumCnidaria

ThomasJankowski,BorisAnokhin

Introduction93 Limitations93 TerminologyandMorphology94 MaterialPreparationandPreservation96 KeystoCnidaria97 References110

5. PhylumPlatyhelminthes

CarolinaNorena,AndreyPorfiriev, OlegTimoshkin

Introduction113 Limitations114 TerminologyandMorphology114 AbbreviationsUsedinFigures116 MaterialPreparationand Preservation117 KeystoPlatyhelminthes118 Acknowledgments144 References144

6. PhylumNemertea

MalinStrand,PerSundberg

Introduction145 Limitations145 TerminologyandMorphology145 MaterialPreparationandPreservation146 KeystoNemertea146 References147

7. PhylumGastrotricha

MariaBalsamo,Jean-Loupd’Hondt,PaoloGrilli

Introduction149 Limitations150

TerminologyandMorphology150

MaterialPreparationandPreservation150

KeystoGastrotricha151 References218

8. PhylumRotifera

RobertL.Wallace,T.W.Snell,E.J.Walsh, S.S.S.Sarma,HendrikSegers

Introduction219

Limitations221

TerminologyandMorphology221

MaterialPreparationandPreservation223

KeytoRotifers224 References264

9. PhylumNematoda

WilfridaDecraemer,UrsulaEisendle-Flockner, EyualemAbebe

Introduction270

Limitations270

TerminologyandMorphology271

MaterialPreparationandPreservation274

KeystoNematoda276 References298

10. PhylumNematomorpha

AndreasSchmidt-Rhaesa

Introduction301

Limitations301

TerminologyandMorphology302

MaterialPreparationandPreservation303

KeystoGordiida303 References307

11. PhylumMollusca

IntroductiontothePhylum310

D.ChristopherRogers

Introduction310

Limitations310

MaterialPreparationandPreservation310

KeystoMollusca310 Reference310

ClassGastropoda310

MaximV.Vinarski

Introduction310 Limitations311 TerminologyandMorphology311

MaterialPreparationandPreservation314 Acknowledgments316 KeystoGastropoda323 References343

ClassBivalvia346

RafaelAraujo

Introduction346 Limitations346 TerminologyandMorphology346

MaterialPreparationandPreservation347 KeystoBivalvia348 References355

12. PhylumAnnelida

IntroductiontothePhylum360 JamesH.Thorp,LawrenceL.Lovell Introduction360 Limitations360 TerminologyandMorphology360 MaterialPreparationandPreservation360 KeystoAnnelida364 References364

ClassClitellata:SubclassOligochaeta364 TarmoTimm,PatrickMartin Introduction365 Limitations365 TerminologyandMorphology365 MaterialPreparationandPreservation367 KeystoOligochaeta369 References482

SubclassBranchiobdellida483

StuartR.Gelder

Introduction483 Limitations483 TerminologyandMorphology483

MaterialPreparationandPreservation484 KeystoBranchiobdellida487 References490

SubclassHirudinida491

FredricR.Govedich,WilliamE.Moser, TakafumiNakano,AleksanderBielecki, BonnieA.Bain,SergeUtevsky

Introduction491

TerminologyandMorphology492

MaterialPreparationandPreservation494

KeystoLeeches495 References507

SubclassAcanthobdella507

ClassAphanoneura508

JoaoGil,ChristopherJ.Glasby,DanielMartin Introduction508 Limitations508

TerminologyandMorphology508

MaterialPreparationandPreservation508

KeystoAphanoneura508 References511

ClassPolychaeta512

JoaoGil,ChristopherJ.Glasby,DanielMartin Introduction512 Limitations513

TerminologyandMorphology513

MaterialPreparationandPreservation514

KeystoPolychaeta515 References517

13. PhylumEctoprocta

TimothyS.Wood

Introduction519 Limitations519

TerminologyandMorphology520

MaterialPreparationandPreservation521

KeystoEctoprocta(Bryozoa)525 References529

14. PhylumEntoprocta

TimothyS.Wood

Introduction531

TerminologyandMorphology531

MaterialPreparationandPreservation532 References532

15. PhylumTardigrada

DianeR.Nelson,RobertoGuidetti, LorenaRebecchi

Introduction533

Limitations535

TerminologyandMorphology535

MaterialPreparationandPreservation540 KeystoTardigrada544 References547

PartII

16. PhylumArthropoda:Introductionand Arachnida

IntroductiontoArthropoda551 D.ChristopherRogers

Introduction551

TerminologyandMorphology551

KeystoArthropoda552 References553

Arachnida:Acari:Trombidiformes: Halacaridae554 IlseBartsch

Introduction554

TerminologyandMorphology554

MaterialPreparationandPreservation554 KeystoHalacaridae555 References562

16.1. Arthropoda:IntroductiontoCrustacea andHexapoda

IntroductiontotheSubphylumCrustacea565 D.ChristopherRogers

Introduction565

TerminologyandMorphology565 KeystoCrustacea566 References566

Crustacea:Hexapoda566

R.EdwardDeWalt,VincentH.Resh

Introduction566

Limitations567

TerminologyandMorphology567

MaterialPreparationandPreservation567

Acknowledgments570

KeystoHexapoda570 References639

16.2. Arthropoda:ClassBranchiopoda

D.ChristopherRogers,AlexeyA.Kotov, ArtemY.Sinev,SergeiM.Glagolev, NikolaiM.Korovchinsky,NikolaiN.Smirnov, EugeniyaI.Bekker

Introduction644

Limitations644

TerminologyandMorphology645

MaterialPreparationandPreservation645

KeystoBranchiopoda646 References724

16.3. Arthropoda:Ostracoda

DavidJ.Horne,ClaudeMeisch, KoenMartens

Introduction725 Limitations726

TerminologyandMorphology726

MaterialPreparationandPreservation726

Acknowledgments727

KeystoOstracoda728 References760

16.4. Arthropoda:Copepoda DongJuLee,WonchoelLee

Introduction761

Limitations761

TerminologyandMorphology761

MaterialPreparationandPreservation762

KeystoCopepoda766 References780

16.5. Arthropoda:Thecostraca

IntroductiontoThecostraca781

D.ChristopherRogers Introduction781

KeystoClassThecostraca781 Reference782

Thecostraca:SubclassSessilia782

RobertJ.VanSyoc

Introduction782

TerminologyandMorphology782

MaterialPreparationandPreservation783 KeytoSubclassSessilia786 References787

16.6. PhylumArthropoda:Malacostraca

IntroductiontoMalacostraca791

D.ChristopherRogers

Introduction791

KeystoMalacostraca791

Malacostraca:Cumacea799

LesWatling

Introduction799 Limitations799 TerminologyandMorphology799 MaterialPreparationandPreservation799 KeystoCumacea800 References801

Malacostraca:Bathynellacea802

D.ChristopherRogers

Introduction802 Limitations802 TerminologyandMorphology802 MaterialPreparationandPreservation802 KeystoBathynellacea803 References808

Malacostraca:Amphipoda808

BorisSket,HiroshiMorino,VadimTakhteev, D.ChristopherRogers

Introduction808 Limitations808

TerminologyandMorphology809

MaterialPreparationandPreservation809 Acknowledgments809 KeystoAmphipoda809 Ponto-CaspianRegionAmphipoda823 LakeBaikalRegionAmphipoda825 References836

Malacostraca:Tanaidacea836

TomHansknecht

Introduction836 Limitations836 TerminologyandMorphology836

MaterialPreparationandPreservation836

KeystoTanaidacea837 References837

Malacostraca:Isopoda837

D.ChristopherRogers,JulianJ.Lewis

Introduction837

Limitations838

TerminologyandMorphology838

MaterialPreparationandPreservation838

Acknowledgments840

KeystoIsopoda840 References844

Malacostraca:Thermosbanacea844

D.ChristopherRogers

Introduction844

Limitations844

TerminologyandMorphology844

MaterialPreparationandPreservation845

KeystoThermosbaenacea845 Reference845

Malacostraca:Decapoda845

SebastianKlaus,NganKeeNg,SameerKumarPati, TohruNaruse,DirkBrandis,D.ChristopherRogers, DarrenC.J.Yeo

Introduction846 Limitations846

TerminologyandMorphology846

MaterialPreparationandPreservation847

KeystoDecapoda847 References865

Malacostraca:MysidaandStygiomysida866

MikhailE.Daneliya,VictorV.Petryashov, RistoVa¨inola

Introduction866 Limitations867

TerminologyandMorphology867

MaterialPreparationandPreservation868

KeystoMysidaandStygiomysida868 References889

TaxonomicIndex891

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EyualemAbebe [Chapter9]DepartmentofNatural Sciences,PharmacyandHealthProfessions,Elizabeth StateUniversity,ElizabethCity,NC,UnitedStates; email:ebabebe@mail.ecsu.edu

BorisAnokhin [Chapter4]ZoologicalInstituteRAS,Universitetskayanab.1,St.Petersburg,Russia;email: cnidaria@nm.ru

RafaelAraujo [Chapter11(Bivalvia)]MuseoNacionalde CienciasNaturales,Madrid,Spain;email: rafael@mncn.csic.es

BonnieA.Bain [Chapter12(Hirudinida)]Departmentof BiologicalSciences,SouthernUtahUniversity,Cedar City,UT,UnitedStates;email:bain@dixie.edu

MariaBalsamo [Chapter7]DepartmentofBiomolecular Sciences(DISB),UniversityofUrbino,“CarloBo”, Urbino,Italy;email:maria.balsamo@uniurb.it

IlseBartsch [Chapter16]ForschungsinstitutSenckenberg, Hamburg,Germany;email:bartsch@meeresforschung.de

EugeniyaI.Bekker [Chapter16.2]A.N.SevertsovInstituteofEcologyandEvolution,Moscow,Russia;email: evbekker@ya.ru

AleksanderBielecki [Chapter12(Hirudinida)]DepartmentofZoology,FacultyofBiologyandBiotechnology,UniversityofWarmiaandMazuryinOlsztyn, Olsztyn,Poland;email:alekb@uwm.edu.pl

DirkBrandis [Chapter16.6(Decapoda)]Zoological Museum,UniversityofKiel,Kiel,Germany;email: brandis@zoolmuseum.uni-kiel.de

MikhailE.Daneliya [Chapter16.6(MysidaandStygomysida)]FinnishMuseumofNaturalHistory,University ofHelsinki,Helsinki,Finland;email:mikhail.daneliya@ helsinki.fi

WilfridaDecraemer [Chapter9]DepartmentofBiology, GhentUniversity,Ghent,Belgium,Departmentof Taxonomy&Phylogeny,RoyalBelgianInstituteof NaturalSciences,Brussels,Belgium;email: wilfrida.decraemer@ugent.be

R.EdwardDeWalt [Chapter16.1]UniversityofIllinois, PrairieResearchInstitute,IllinoisNaturalHistory

Survey,Champaign,IL,UnitedStates;email: edewalt@inhs.illinois.edu

Jean-Loupd’Hondt [Chapter7]De ´ partement“Milieuxet PeuplementsAquatiques”,Muse ´ umNational d’HistoireNaturelle,Paris,France;email: dhondt@mnhn.fr

UrsulaEisendle-Flockner [Chapter9]Departmentof CellBiologyandPhysiology,UniversityofSalzburg, Salzburg,Austria;email:Ursula.Eisendle@sbq.ac.at

GenovevaF.Esteban [Chapter2]BournemouthUniversity,FacultyofScienceandTechnology,Department ofLifeandEnvironmentalSciences,Dorset,United Kingdom;email:gesteban@bournemouth.ac.uk

StuartR.Gelder [Chapter12(Branchiobdellida)]DepartmentofScienceandMath,UniversityofMaineatPresqueIsle,PresqueIsle,ME,UnitedStates;email: stuart.gelder@umpi.edu

Joa ˜ oGil [Chapter12(Aphanoneura;Polychaeta)] CCMAR,UniversidadedoAlgarve,Faro,Portugal; email:joaocfgil@gmail.com

SergeiM.Glagolev [Chapter16.2]MoscowSouth-West HighSchool,Moscow,Russia;email: sglagolev@yandex.ru

ChristopherJ.Glasby [Chapter12(Aphanoneura;Polychaeta)]Museum&ArtGalleryNorthernTerritory, Darwin,NT,Australia;email:chris.glasby@nt.gov.au

FredricR.Govedich [Chapter12(Hirudinida)]DepartmentofBiologicalSciences,SouthernUtahUniversity, CedarCity,UT,UnitedStates;email: govedich@suu.edu

PaoloGrilli [Chapter7]DepartmentofBiomolecular Sciences(DISB),UniversityofUrbino“CarloBo”, Urbino,Italy;email:paolo.grilli2@tin.it

RobertoGuidetti [Chapter15]DepartmentofLife Sciences,UniversityofModenaandReggioEmilia, Modena,Italy;email:roberto.guidetti@unimore.it

TomHansknecht [Chapter16.6(Tanaidacea)]BarryVittor&Associates,Inc.,Mobile,AL,UnitedStates; email:bvataxa@bvaenviro.com

DavidJ.Horne [Chapter16.3]SchoolofGeography, QueenMary,UniversityofLondon,London,United Kingdom;email:d.j.horne@qmul.ac.uk

ThomasJankowski [Chapter4]Teningen,Germany; email:thomas.jankowski@posteo.de

SebastianKlaus [Chapter16.6(Decapoda)]Departmentof Ecology&Evolution,Goethe-University Biologicum, FrankfurtamMain,Germany;email:klaus@bio. uni-frankfurt.de

NikolaiM.Korovchinsky [Chapter16.2]A.N.Severtsov InstituteofEcologyandEvolution,Moscow,Russia; email:nmkor@yandex.ru

AlexeyA.Kotov [Chapter16.2]A.N.SevertsovInstitute ofEcologyandEvolution,Moscow,Russia;email: alexey-a-kotov@vandex.ru

DongJuLee [Chapter16.4]DepartmentofLifeScience, HanyangUniversity,Seoul,RepublicofKorea;email: velocy00@gmail.com

WonchoelLee [Chapter16.4]DepartmentofLifeScience, HanyangUniversity,Seoul,RepublicofKorea;email: wlee@hanyang.ac.kr

JulianJ.Lewis [Chapter16.6(Isopoda)]Lewis&AssociatesLLC,Borden,IN,UnitedStates;email: lewisbioconsult@aol.com

LawrenceL.Lovell [Chapter12(Introductiontothe Phylum)]ResearchandCollections,NaturalHistory MuseumofLosAngelesCounty,LosAngeles,CA, UnitedStates;email:lllpolytax@gmail.com

RenataManconi [Chapter3]DipartimentodiMedicina Veterinaria,Universita ` diSassari,Sassari,Italy;email: r.manconi@uniss.it

KoenMartens [Chapter16.3]RoyalBelgianInstituteof NaturalSciences,FreshwaterBiology,Brussels, Belgium;email:darwinula@gmail.com

DanielMartin [Chapter12(Aphanoneura;Polychaeta)] CEAB-CSIC,Blanes,Catalunya,Spain;email: dani@ceab.csic.es

PatrickMartin [Chapter12(Oligochaeta)]RoyalBelgian InstituteofNaturalSciences,OperationalDirectorate Taxonomy&Phylogeny,Brussels,Belgium;email: patrick.martin@sciencenaturelles.be

ClaudeMeisch [Chapter16.3]Muse ´ enationald’histoire naturelle,Luxembourg;email:claude.meisch@ education.lu

HiroshiMorino [Chapter16.6(Amphipoda)]Department ofZoology,NationalMuseumofNatureandScience, Tsukuba,Ibaraki,Japan;email:hiroshi.morino.talitrids @vc.ibaraki.ac.jp

WilliamE.Moser [Chapter12(Hirudinida)]Department ofInvertebrateZoology,NationalMuseumofNatural History,SmithsonianInstitution,Suitland,MD,United States;email:moserw@si.edu

TakafumiNakano [Chapter12(Hirudinida)]Department ofScienceEducation,GraduateSchoolofEducation,HiroshimaUniversity,Higashihiroshima,Japan;Department ofZoology,GraduateSchoolofScience,KyotoUniversity,Kyoto,Japan;email:tnakano@hiroshima-u.ac.jp

TohruNaruse [Chapter16.6(Decapoda)]Tropical BiosphereResearchCenter,UniversityoftheRyukyus, Taketomi,Japan;email:naruse@lab.u-ryukyu.ac.jp

DianeR.Nelson [Chapter15]DepartmentofBiological Sciences,EastTennesseeStateUniversity,Johnson City,TN,UnitedStates;email:janddnelson@yahoo.com

NganKeeNg [Chapter16.6(Decapoda)]Departmentof BiologicalSciences,NationalUniversityofSingapore, RepublicofSingapore;email:ngankee@nus.edu.sg

CarolinaNorena [Chapter5]DepartamentoBiodiversidadyBiologı´aEvolutiva,MuseoNacionalde CienciasNaturales(CSIC),Madrid,Espana;email: mcnnj92@mncn.csic.es

SameerKumarPati [Chapter16.6(Decapoda)]ZoologicalSurveyofIndia,WesternRegionalCentre,Pune, India;email:sameerkumarpati@gmail.com

VictorV.Petryashov [Chapter16.6.(MysidaandStygiomysida)]ZoologicalInstituteoftheRussianAcademy ofSciences,SaintPetersburg,Russia;email: taxalab@taxonomicum.com

AndreyPorfiriev [Chapter5]KazanStateUniversity, Kazan,Russia;email:andpor@rambler.ru

RobertoPronzato [Chapter3]DipartimentodiScienze dellaTerra,dell’AmbienteedellaVita(DISTAV),Universita ` diGenova,Genova,Italy;email: pronzato@dipteris.unige.it

LorenaRebecchi [Chapter15]DepartmentofLife Sciences,UniversityofModenaandReggioEmilia, Modena,Italy;email:lorena.rebecchi@unimore.it

VincentH.Resh [Chapter16.1(Hexapoda)]Department ofEnvironmentalScience,Policy,andManagement, UniversityofCalifornia,Berkeley,CA,UnitedStates; email:resh@berkeley.edu

D.ChristopherRogers [Chapters1,11,16,16.1,16.2, 16.5,16.6]KansasBiologicalSurveyandTheBiodiversityInstitute,UniversityofKansas,Lawrence,KS, UnitedStates;email:branchiopod@gmail.com

S.S.S.Sarma [Chapter8]UniversidadNacionalAuto ´ nomadeMe ´ xicoCampusIztacala,Me ´ xicoCity, Me ´ xico;email:sssarma@gmail.com

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

HendrikSegers [Chapter8]RoyalBelgianInstituteof NaturalSciences,Brussels,Belgium;email: Hendrik.Segers@naturalsciences.be

ArtemY.Sinev [Chapter16.2]DepartmentofInvertebrate Zoology,BiologicalFaculty,M.V.LomonosovMoscowStateUniversity,LeninskieGory,Moscow,Russia; email:artem.sinev@gmail.com

BorisSket [Chapter16.6(Amphipoda)]Biologydepartment,BiotechnicalFaculty,UniverzavLjubljani, Slovenia;email:sidorov@biosoil.ru

NikolaiN.Smirnov [Chapter16.2]A.N.SevertsovInstituteofEcologyandEvolution,Moscow,Russia;email: smirnov08520@mail.ru

T.W.Snell [Chapter8]SchoolofBiology,GeorgiaInstituteofTechnology,Atlanta,GA,UnitedStates;email: terry.snell@biology.gatech.edu

MalinStrand [Chapter6]TheSwedishSpeciesInformationCentre,SwedishUniversityofAgricultural Sciences,Uppsala,Sweden;email:Malin.Strand@slu.se

PerSundberg [Chapter6]DepartmentofMarine Sciences,UniversityofGothenburg,Gothenburg, Sweden;email:P.Sundberg@zool.gu.se

VadimTakhteev [Chapter16.6(Amphipoda)]Department ofInvertebrateZoology&Hydrobiology,IrkutskState University,Russia;email:amphipoda@yandex.ru

JamesH.Thorp [Chapters1,12]KansasBiological SurveyandDepartmentofEcologyandEvolutionary Biology,UniversityofKansas,Lawrence,KS,United States;email:thorp@ku.edu

TarmoTimm [Chapter12(Oligochaeta)]Instituteof AgriculturalandEnvironmentalSciences,Centrefor Limnology,EstonianUniversityofLifeSciences, Tartu,Estonia;email:tarmo.timm@emu.ee

OlegTimoshkin [Chapter5]LimnologicalInstitute,UlanBatorskaya,Irkutsk,Russia;email:tim@lin.irk.ru

SergeUtevsky [Chapter12(Hirudinida)]Departmentof ZoologyandAnimalEcology,V.N.KarazinKharkiv NationalUniversity,Kharkiv,Ukraine;email: serge.utevsky@gmail.com

RistoVa ¨ ino ¨ la ¨ [Chapter16.6(MysidaandStygiomysida)] FinnishMuseumofNaturalHistory,Universityof Helsinki,Helsinki,Finland;email:risto.vainola@ helsinki.fi

RobertJ.VanSyoc [Chapter16.5(Sessilia)]California AcademyofSciences,SanFrancisco,CA,United States;email:bvansyoc@calacademy.org

MaximV.Vinarski [Chapter11(Gastropoda)]SaintPetersburgStateUniversity,Saint-Petersburg,Russia; email:radix.vinarski@gmail.com

RobertL.Wallace [Chapter8]DepartmentofBiology, RiponCollege,Ripon,WI,UnitedStates;email: wallacer@ripon.edu

E.J.Walsh [Chapter8]DepartmentofBiologicalScience, UniversityofTexasatElPaso,ElPaso,TX,United States;email:ewalsh@utep.edu

AlanWarren [Chapter2]DepartmentofLifeSciences, NaturalHistoryMuseum,London,UnitedKingdom; email:a.warren@nhm.ac.uk

LesWatling [Chapter16.6(Cumacea)]Departmentof Biology,UniversityofHawaiiatManoa,Honolulu, HI,UnitedStates;email:watling@hawaii.edu

TimothyS.Wood [Chapters13-14]DepartmentofBiologicalSciences,WrightStateUniversity,Dayton, OH,UnitedStates;email:tim.wood@wright.edu

DarrenC.J.Yeo [Chapter16.6(Decapoda)]Department ofBiologicalSciences,NationalUniversityof Singapore,RepublicofSingapore;email: darrenyeo@nus.edu.sg

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AbouttheEditors

Dr.D.ChristopherRogers isaresearchzoologistatthe UniversityofKansaswiththeKansasBiologicalSurvey andisaffiliatedwiththeBiodiversityInstitute.Hereceived hisPh.D.degreefromtheUniversityofNewEnglandin Armidale,NSW,Australia.Christopherspecializesin freshwatercrustaceans(particularlyBranchiopodaand Malacostraca)andtheinvertebratefaunaofseasonally astaticwetlandsonaglobalscale.Hehasnumerouspeerreviewedpublicationsincrustaceantaxonomyandinvertebrateecology,aswellaspublishedpopularandscientific fieldguidesandidentificationmanualstofreshwater invertebrates.ChristopherisanAssociateEditorforthe JournalofCrustaceanBiologyandafoundingmemberof theSouthwestAssociationofFreshwaterInvertebrate Taxonomists.Hehasbeeninvolvedinaquaticinvertebrate conservationeffortsallovertheworld.

Dr.JamesH.Thorp hasbeenaProfessorintheDepartmentof EcologyandEvolutionaryBiologyattheUniversityofKansas (Lawrence,KS,USA)andaSeniorScientistintheKansas BiologicalSurveysince2001.Priortoreturningtohisalma mater,Prof.ThorpwasaDistinguishedProfessorandDeanat ClarksonUniversity,DepartmentChairandProfessoratthe UniversityofLouisville,AssociateProfessorandDirectorof theCalderEcologyCenterofFordhamUniversity,Visiting AssociateProfessoratCornell,andResearchEcologistatthe UniversityofGeorgia’sSavannahRiverEcologyLaboratory. HereceivedhisBaccalaureatefromtheUniversityofKansas (KU)andbothMastersandPh.D.degreesfromNorthCarolina State.Thosedegreesfocusedonzoology,ecology,andmarine biology,withanemphasisontheecologyoffreshwaterand marineinvertebrates.Dr.Thorphasbeenontheeditorialboard ofthreefreshwaterjournalsandisaformerPresidentofthe InternationalSocietyforRiverScience.Heteachesfreshwater, marine,andinvertebratecoursesatKU,andhisMastersand Doctoralgraduatestudentsworkonvariousaspectsofthe ecologyofcommunitiesthroughmacrosystemsinrivers, reservoirs,andwetlands.Prof.Thorp’sresearchinterestsand backgroundarehighlydiverseandspanthegamutfrom organismalbiologytocommunity,ecosystem,andmacrosystemecology.Heworksonbothfundamentalandapplied researchtopicsusingdescriptive,experimental,andmodeling approachesinthefieldandlab.Whilehisresearchemphasizes aquaticinvertebrates,healsostudiesfishecology,especiallyas relatedtofoodwebs.Hehaspublishedmorethanonehundred refereedjournalarticles,books,andchapters,includingthree single-volumeeditionsof EcologyandClassificationofNorth AmericanFreshwaterInvertebrates (editedbyJ.H.Thorpand A.P.Covich)andthreevolumesinthecurrentfourtheditionof ThorpandCovich’sFreshwaterInvertebrates

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Thosereadersfamiliarwiththefirstthreeeditionsofour invertebratebook(EcologyandClassificationofNorth AmericanFreshwaterInvertebrates,editedbyJ.H.Thorp andA.P.Covich)willnotethatthefourtheditionhas expandedfromaNorthAmericanfocustoworldwide coverageofinlandwaterinvertebrates.Wegaveourbook seriesoninlandwaterinvertebratesthename Thorpand Covich’sFreshwaterInvertebrates to:(1)associatepresent withpasteditions,unitecurrentvolumes,andlinktofuture editions;(2)establishaconnectionbetweentheecological andgeneralbiologycoverageinVolumeIwiththetaxonomickeysintheremainingvolumes;and(3)givecreditto ProfessorAlanCovichforhisworkonthefirstthree editions.Forthesakeofbrevity,werefertothecurrent editionasT&CIV.WhetherafiftheditionofT&Cwillever appeariscertainlyproblematic,butwhoknows?Atpresent weareconsideringproducingupto12volumesinthe fourthedition.

WhileIamthesoleeditorofthe“series”atthispoint,Dr. D.ChristopherRogershasbeenamajorandhighlyvalued partnerindevelopingideasforthefourtheditionandisthus faracoeditoronthefirstfourvolumesinprint(senioreditor onthefourth)aswellasonthethreevolumesindevelopment.Heisalsolikelytoplayamajorroleinmanyofthe remainingvolumesbecauseofhisdiverseandglobal knowledgeoffreshwaterinvertebrates,especiallyinthearea oftaxonomy.Aswemadesignificantprogressonthefirst sevenvolumes,webegancontactingpotentialcoeditorsto developtwomorevolumesforanotherzoogeographic region,andnegotiationswiththosepotentialeditorsarenow underway.However,wearestillseekingexpertsinfieldsof invertebratetaxonomyforvariouszoogeographicregionsto serveashighlydependablecoeditors,especiallythosewho bothworkandliveinthezoogeographicregionscoveredby thevariousfuturevolumes.

OurconceptforT&CIVincludedproducingonebook (Vol.Iin2015)withsixchaptersongeneralenvironmental issuesapplicabletomanyinvertebrates,followedby35 chaptersdevotedtoindividualtaxaatvariouslevels(order tophylum,orevenmultiplephylainthecaseofthe protozoa).VolumeIwasdesignedbothasanindependent bookonecologyandgeneralbiologyofthevariousfreshwaterinvertebratetaxaandasacompanionvolumefor usersofthekeysintheregionaltaxonomicvolumes, therebyreducingtheamountofinformationduplicatedin thetaxonomicvolumes.Theperhaps11taxonomic volumesweforeseepublishinginthisserieswillcontain bothkeysforidentifyinginvertebratesinspecificzoogeographicregionsanddescriptionsofdetailedanatomical featuresneededtoemploythosekeys.

WhilethevastmajorityofauthorsinT&CI IIIwere fromtheUSAorCanada,weattemptedintheglobal volumesofT&CIVtoattractauthorsfrommanyadditional countriesonsixcontinents.Althoughwelargelysucceeded inthisgoal,weexpectthefiftheditionofT&C ifitisever published tocontinueincreasingtheproportionof authorsfromoutsideNorthAmericaasourbooksbecome betterknowninternationally.

OurgoalsforT&CIVaretoimprovethestateof taxonomicandecologicalknowledgeofinlandwater invertebrates,tohelpprotectouraquaticbiodiversity,and toencouragemorestudentstodevotetheircareersto workingwiththesefascinatingorganisms.Thesegoalsare especiallyimportantbecausetheverifiedandprobable lossesofspeciesinwetlands,ponds,lakes,creeks,and riversaroundtheglobeexceedthoseinmostterrestrial habitats.

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Thisisthefourthvolumeofthefourtheditionof Thorpand Covich’sFreshwaterInvertebrates (T&CIV),andthe secondtofocusonabioregionoutsidetheNearctic. Informationontheecologyandgeneralbiologyofthe groupscanbefoundinVolumeI(EcologyandGeneral Biology,editedbyThorp&Rogers,2015),thecompanion textforthecurrentandallremainingbooksinthisseries. Alltaxonomicvolumes(otherthanthosefocusedexclusivelyonHexapoda)areexpectedtoconsistofanintroductorychapter,achapteronprotozoa(multiple kingdoms),andmultiplechaptersonindividualphylafrom CnidariatovariousgroupswithinArthropoda.Someofthe chaptersareverysmall(e.g.,Chapter14onEntoprocta), whereasothersarequitelarge.AmajorchangeinVolume IVisthedivisionofthesinglechapteronArthropodafound inVolumeII(whichrepresentedmorethanhalfthepagesof thatvolume)intoaseparatesectionencompassingsix chapters.

Atypicalchapterincludesashortintroduction,abrief discussionoflimitstoidentificationoftaxainthatchapter, importantinformationonterminologyandmorphologythat isneededtousethekeys,techniquesforpreparingand preservingmaterialforidentification(alsocoveredin VolumeI),thetaxonomickeys,andafewreferences.Inthe largechaptersonMollusca(11),Annelida(12),and Arthropoda(16 22),differentindividualshavecontributed separatesections,andthustherearemultiplesectionson introductionthroughkeysandreferences.Whilethismay confusesomereaders,ithasallowedustogaincontributionsfromanincreasednumberofexpertsaroundthe world.

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

backgroundexperience.Wealsodesignedmostkeys, wherepossible,toproceedfromageneraltoaspecific characterwithinacouplet.

Wehaveaskedauthorstoincludeonlytaxathatare recognizedinternationallybypublicationinreputable scientificjournalswhichfollowtheInternationalCodeof ZoologicalNomenclature.Thus,notaxathathavemerely beenproposedshouldhavebeenincluded,eveniftheyhave beenidentifiedbytheworld’sexpertonthatgroup. “Common”speciesarenotdesignatedinthekeysthemselvesbecauseacommonspeciesinoneareamaynotbe commoninanother,andthisdesignationcanleadtooverly frequentandfalseidentifications.Authorshavebeen encouragedtoendthekeysatthepointwherefurther identificationwithoutgeneticanalysisisnotpracticalor whenitisclearthattoomanyoftheextantfaunahaveyetto bedescribedinscientificpublications.

Usersofthesekeysneedtorealizethattaxonomyis agrowingandvibrantfieldinwhichnewtaxaarebeing describedandpreviouslyacceptedrelationshipsre-evaluated.Forsomeusers,thisvolumemaybesufficientfortheir needs,butforothersacompaniontextlistingknown speciesinasmallergeographicregionmayalsobehelpful.

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

Itisourhopethatscientistsandstudentsfromaround theworldwillbenefitfromthisvolume,especiallythose studyingorganismsinthePalaearctic.Suggestionsfor improvingfuturevolumesarewelcomed.

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Manypeoplecontributedtothisvolumeinadditiontothe chapterauthorsandthoseacknowledgedinindividual chapters.Wegreatlyappreciateallourcolleagueswhohave contributedinformation,figures,orreviewstoVolumeIII andalsothankthosewhoprovidedsimilarservicesforthe earliereditions,uponwhichthepresentbookpartially relies.Weareagaingratefultothehighlycompetentpeople atAcademicPress/Elsevierwhohelpedinmanyaspectsof

AcknowledgmentsforVolumeIV

thebook’sproductionfromtheoriginalconcepttothefinal marketing.Inparticular,weappreciateourassociationwith ElseviereditorsincludingEmilyThomsonandLouisa MunroandearlierLauraKelleher,aswellastheentire globalproductionteamsforElsevier.

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Introduction1

JamesH.Thorp

KansasBiologicalSurveyandDepartmentofEcologyandEvolutionaryBiology,UniversityofKansas,Lawrence,KS,UnitedStates

D.ChristopherRogers

KansasBiologicalSurveyandTheBiodiversityInstitute,UniversityofKansas,Lawrence,KS,UnitedStates

ChapterOutline

IntroductiontoThisSeries,Volume,andChapter1 NatureandFaunaofthePalaearcticBioregion2 ComponentsofTaxonomicChapters2 HowtoUseThisVolume2

INTRODUCTIONTOTHISSERIES, VOLUME,ANDCHAPTER

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

Inadditiontothefirstfourvolumescurrentlyinprint (Thorp&Rogers,2015,2016; Hamadaetal.,2019;andthe currentvolume),twovolumesintheseriesmayappearin 2019 2020:(1)KeystoNeotropicalandAntarcticFauna (Damboreneaetal.,2019,inprep.);and(2)Keysto AustralasianFauna(Ahyongetal.,2020,inprep).Future

1Thischapterwaswrittentobeausefulstartingpointfortheprevious, current,andfuturetaxonomicvolumesinallzoogeographicregions. Consequently,therewillbeonlyminortomoderatedifferencesinthis chapteramongvariousvolumes.

KeystoMajorTaxaofInlandWaterInvertebratesofthe Palaearctic3 References4

volumeswillcoverAfrotropicalFauna(Dayetal.;perhaps in2022)andOriental/Oceanafauna(eachwithonetotwo volumes)aswellasmoredetailedcoverageofHexapodain theNearcticandPalaearcticBioregions.Weconsiderthis seriesof10 12volumestobeagloballyexpanded,fourth editionoftheearlier EcologyandClassificationofNorth AmericanFreshwaterInvertebrates (editedbyThorp& Covichin1991,2001,and2010).Whilesomelabsmay havemultiplecopiesofthe“KeystoFauna”intheirregion, wealsorecommendthattheyhaveatleastonecopyof VolumeItoobtainusefulbackgroundinformationoneach invertebrategroup.

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

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