Parasitism: The Diversity and Ecology of Animal Parasites

524 Pages·2014·17.79 MB·English

Parasitism
The Diversity and Ecology of Animal Parasites
SECOND EDITION
Reflectingtheenormousadvancesmadeinthefieldoverthepast10years,thistextsynthesizes
thelatestdevelopmentsintheecologyandevolutionofanimalparasitesagainstabackdrop
ofparalleladvancesinparasitesystematics,biodiversity,andlifecycles.Ithasbeenthoroughly
revisedtomeettheneedsofanewgenerationofparasitologystudents,whethertheirinterest
isinecology,conservationbiology,evolution,immunology,orhealthsciences.
Balancingtraditionalapproachesinparasitologywithmodernstudiesinparasiteecology
andevolution,theauthorspresentbasicecologicalprinciplesasaunifyingframeworktohelp
studentsunderstandthecomplexphenomenonofparasitism.Richlyillustratedwithover
300figures,thetextisaccompaniedbycasestudyboxesdesignedtohelpstudentsappreciate
thecomplexityanddiversityofparasitesandthescientistswhostudythem.Thisunique
approach,whichispresentedclearlyandwithaminimumofjargonandmathematicaldetail,
encouragesstudentstothinkgenerallyandconceptuallyaboutparasitesandparasitism.
TimothyM.GoaterisProfessorandformerChairintheBiologyDepartmentatVancouverIsland
University,BritishColumbia,Canada.Duringthepast20yearshehastaughtcoursesin
introductorybiology,parasitology,ecologicalparasitology,invertebratezoology,andento-
mology.Hisresearchinterestsfocusonthepopulationandcommunityecologyofparasites.
CameronP.GoaterisAssociateProfessorandformerChairintheDepartmentofBiological
SciencesattheUniversityofLethbridge,Alberta,Canada.Hisparasitologicalresearchrootsare
inthecommunityecologyofhelminthsofwaterfowlontheCanadianprairies,andoverthe
past15yearshehastaughtcoursesinintroductorybiology,invertebratebiology,fieldbiology,
andsymbioticinteractions.Hiscurrentresearchinterestsareintheexperimentalecologyof
helminth–hostinteractions.
GeraldW.EschisCharlesM.AllenProfessorofBiologyatWakeForestUniversity,North
Carolina,USA,wherehehastaughtfor47years.Heiswidelyregardedtobeoneofthe
world’sleadingecologicalparasitologists,andservedasEditoroftheJournalofParasitologyfor
19years.
AdvancepraiseforParasitism:TheDiversityandEcologyofAnimal
Parasites, Second Edition
“Theirapproachissynthetic,refreshinglyoriginalandeffectivelyblendscoverageoflong-
standingfundamentalsofparasitologywithmodernadvancesinthefield.”
JanineN.Caira,UniversityofConnecticut,USA
“Thisisanextremelywellwrittenbookthatdoesanexcellentjobofintegratingconceptualand
organismalaspectsofparasitology.”
DaleH.Clayton,UniversityofUtah,USA
“Thereisawealthofdetailforwell-selectedexamples,buildingontherichexperienceofthe
authorsastop-notchresearchersandeducators.”
MarkR.Forbes,CarletonUniversity,Canada
“Parasitismgivesthestudentboththesystematicandzoologicalbackgroundtounderstand
parasitologyandtheecologicalandevolutionarycontexttounderstandwhyitisimportant...
Asateam,theirapproachisclearandscholarly,withmanyimportantupdatessincethefirst
edition.”
KevinD.Lafferty,USGeologicalSurvey,UniversityofCalifornia,SantaBarbara,USA
“Thisneweditionwillbeawonderfulresourceforteachersofundergraduateparasitology
courses.Thewell-illustratedandeasy-to-readtextisunrivalledatthemomentandwillbea
greattooltoturnonanewgenerationofyoungmindstothewondersofparasiticorganisms.A
trueparasitologicaltourdeforce!”
RobertPoulin,UniversityofOtago,NewZealand
“Awell-organizedintegrationofthediversityofideasandmethodsthatcharacterizethisnew
fieldofparasiteecology.Thestyleiseasilyreadable,thedetailsextraordinary,andthestoryis
toldfromtheperspectiveofevolutionarythought...eventheproswilllearnfromthisbook.”
MichaelV.KSukhdeo,RutgersUniversity,USA
Parasitism
The Diversity and Ecology
of Animal Parasites
SECOND EDITION
TIMOTHY M. GOATER
VancouverIslandUniversity,BritishColumbia,Canada
CAMERON P. GOATER
UniversityofLethbridge,Alberta,Canada
GERALD W. ESCH
WakeForestUniversity,NorthCarolina,USA
UniversityPrintingHouse,CambridgeCB28BS,UnitedKingdom
PublishedintheUnitedStatesofAmericabyCambridgeUniversityPress,NewYork
CambridgeUniversityPressispartoftheUniversityofCambridge.
ItfurtherstheUniversity’smissionbydisseminatingknowledgeinthepursuitof
education,learning,andresearchatthehighestinternationallevelsofexcellence.
www.cambridge.org
Informationonthistitle:www.cambridge.org/9780521190282
©CambridgeUniversityPress2001,2014
Thispublicationisincopyright.Subjecttostatutoryexception
andtotheprovisionsofrelevantcollectivelicensingagreements,
noreproductionofanypartmaytakeplacewithoutthewritten
permissionofCambridgeUniversityPress.
Firstpublished2001
Secondedition2014
PrintedintheUnitedStatesofAmericabyEdwardsBrothersMalloy
AcatalogrecordforthispublicationisavailablefromtheBritishLibrary
LibraryofCongressCataloginginPublicationdata
Goater,TimothyM.,1959–
Parasitism:thediversityandecologyofanimalparasites/TimothyM.Goater,Cameron
P.Goater,GeraldW.Esch.–Secondedition.
pages cm.
Firstpublishedas:Parasitism/AlbertO.Bush...[andothers],2001.
Includesbibliographicalreferencesandindex.
ISBN978-0-521-19028-2(hardback)–ISBN978-0-521-12205-4(paperback)
1. Parasites–Textbooks. 2. Parasitism–Textbooks. 3. Parasites–Ecology–Textbooks.
4. Biodiversity–Textbooks. 5. Parasitology–Textbooks. I. Goater,CameronP.
II. Esch,GeraldW. III. Parasitism. IV. Title.
QL757.P287 2013
578.6′5–dc23 2013016194
ISBN978-0-521-19028-2Hardback
ISBN978-0-521-12205-4Paperback
Additionalresourcesforthispublicationatwww.cambridge.org/parasitism
CambridgeUniversityPresshasnoresponsibilityforthepersistenceoraccuracyof
URLsforexternalorthird-partyinternetwebsitesreferredtointhispublication,
anddoesnotguaranteethatanycontentonsuchwebsitesis,orwillremain,
accurateorappropriate.
We dedicate this book to our students,
past, present, and future

CONTENTS
Listofboxes pageix 5.5 Phylogeneticrelationships
ForewordbyJohnC.Holmes xi andclassification 106
Prefaceandacknowledgments xiii
6 Platyhelminthes:theflatworms
113
1 Introduction
1 6.1 Generalconsiderations 113
1.1 Encounterswithparasites 1 6.2 Temnocephalidea 114
1.2 Scope 2 6.3 Udonellida 115
1.3 Terminology 5 6.4 Aspidobothrea 115
1.4 Overview 10 6.5 Digenea 116
6.6 Monogenea 145
2 Immunologicalaspectsofparasitism 16 6.7 Gyrocotylidea 152
2.1 Generalconsiderations 16 6.8 Amphilinidea 153
2.2 Vertebrateimmunity 21 6.9 Eucestoda 153
2.3 Invertebrateimmunity 31 6.10 Phylogeneticrelationshipsand
2.4 Ecologicalimmunology 34 classification 169
3 Protista:theunicellulareukaryotes 7 Acanthocephala:thethorny-headed
40
worms
3.1 Generalconsiderations 40 179
3.2 Formandfunction 41 7.1 Generalconsiderations 179
3.3 Biodiversityandlife-cyclevariation 43 7.2 Formandfunction 179
3.4 Phylogeneticrelationshipsand 7.3 Nutrientuptakeandmetabolism 186
classification 80 7.4 Developmentandgenerallifecycle 188
7.5 Biodiversityandlife-cyclevariation 191
4 Microsporida:theintracellular, 7.6 Phylogeneticrelationshipsand
spore-formingfungi classification 192
86
4.1 Generalconsiderations 86
8 Nematoda:theroundworms
4.2 Formandfunction 86 199
4.3 Developmentandgenerallifecycle 88 8.1 Generalconsiderations 199
4.4 Biodiversityandlife-cyclevariation 88 8.2 Formandfunction 199
8.3 Nutrientuptakeandmetabolism 208
4.5 Phylogeneticrelationshipsand
classification 93 8.4 Developmentandgenerallifecycle 209
8.5 Biodiversityandlife-cyclevariation 211
5 Myxozoa:thespore-formingcnidarians 8.6 Phylogeneticrelationshipsand
96
classification 237
5.1 Generalconsiderations 96
5.2 Formandfunction 97
9 Nematomorpha:thehairworms
244
5.3 Developmentandgenerallifecycle 97
5.4 Biodiversityandlife-cycle 9.1 Generalconsiderations 244
variation 100 9.2 Formandfunction 244
viii Contents
9.3 Developmentandgenerallifecycle 246 14 Parasitebiogeographyand
9.4 Biodiversityandecology 247 phylogeography 379
9.5 Phylogeneticrelationshipsand 14.1 Generalconsiderations 379
classification 249 14.2 Historicalbiogeography 380
14.3 Ecologicalbiogeography 386
10 Pentastomida:thetongueworms
252 14.4 Appliedaspectsofparasitebiogeography
10.1 Generalconsiderations 252 andphylogeography 389
10.2 Formandfunction 252
10.3 Nutrientuptakeandmetabolism 256 15 Effectsofparasitesontheirhosts:
10.4 Developmentandgenerallifecycle 256 fromindividualstoecosystems 396
10.5 Biodiversityandlife-cyclevariation 258 15.1 Generalconsiderations 396
10.6 Phylogeneticrelationshipsand 15.2 Effectsofparasitesonhostindividuals 396
classification 260
15.3 Effectsofparasitesonhostpopulations 411
15.4 Effectsofparasitesonhostcommunities
11 Arthropoda:thejoint-leggedanimals
263 andecosystems 422
11.1 Generalconsiderations 263
11.2 Crustacea 264 16 Evolutionofhost–parasiteinteractions 432
11.3 Chelicerata 289 16.1 Generalconsiderations 432
11.4 Hexapoda 308 16.2 Parasite-mediatednaturalselectionand
11.5 Phylogeneticrelationshipsand evolution 432
classification 328 16.3 Geneticstructureofparasite
populations 441
12 Parasitepopulationecology 335 16.4 Introductiontohost–parasite
12.1 Generalconsiderations 335 coevolution 446
12.2 Terminologyandgeneralapproaches 336
12.3 Introductiontoparasitepopulation 17 Environmentalparasitology:parasites
ecology 337 asbioindicatorsofecosystemhealth 459
17.1 Generalconsiderations 459
13 Parasitecommunityecology
356 17.2 Parasitesaseffectindicatorsofpollutant
13.1 Generalconsiderations 356 stress 460
13.2 Introductiontoparasitecommunity 17.3 Parasitesasenvironmentalsentinels 470
ecology 357
13.3 Thestructureofparasiteinfra-
Glossary 477
communities:restrictedniches 361
Index 489
13.4 Thestructureofparasitecommunities:
speciesrichness 368 Thecolorplatesarelocatedbetweenpages248
and249.