Ecological parasitology reflections on 50 years of research in aquatic ecosytems

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TableofContents
Cover
TitlePage
Preface
Acknowledgments
ThePlayers
Microbiographies
Closingcomments
References
1TheBeginning
References
2TheEndoftheBeginning
References
3GullLakeandtheW.K.KelloggBiologicalStation
References
4GullLakeandtheConnectionwiththeSavannahRiverEcologyLaboratory
References
5DevelopmentofSomeConceptualNotions
References
6ThePond:PartI
References
7ThePond:PartII
References
8TheBigLake

References
9TheStrigeids
References
10SomeSmallStreamsandSmallPonds
References
11RedSoreDisease
References
12TheEnd,Almost

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References
13TheCatastrophicCollapseoftheLarvalTrematodeComponentCommunityinCharlie’s
Pond(NorthCarolina)
References
14AnEpilogue:What’sInvolvedwithGraduateSchool?
Whattolookforifyouarebeingrecruited
Planningyourprogram
Thestudent’sadvisorycommittees
Specialskillrequirements
Handlingthespecialexams
Writinganddefendingthethesis/dissertation
Publishingpapersandselectionofthejournal
Collaboration
Graduation(andsometimesapostdocexperience)
Wouldyoudoitagain?
EndUserLicenseAgreement

ListofIllustrations

Preface
Figure0.1AphototakenattheAmericanaHotel,SanJuan,PuertoRico,November
1966,41stannualmeeting,theAmericanSocietyofParasitologists.Fromlefttoright:
me,AnnEsch,HoraceBailey,JayneBailey,SueMcDaniel,JimMcDaniel,Mrs.Ida
Self,Dr.J.TeagueSelf,andKarenJanovy.
Chapter01
Figure1.1AcoenurusofTaeniamulticeps.Thebladderwasabouttwicethesizeofan
ordinarygolfball.Notethatthescolicesareinseveralrows(hencetheoldname
Multiceps[manyheads]serialis[inaseriesofrows]).Thecoenuruswasremoved
fromajackrabbitshotontheshortgrassprairieofeasternColoradoduringmysenior
yearatColoradoCollege.
Figure1.2AcysticercusofTaeniacrassiceps,about5 mminsize,fromtoptobottom.
ThiscysticercusbelongstotheanomalousOntarioResearchFoundation(ORF)strain
isolatedbyReino(Ray)FreemanattheUniversityofToronto.Thescolexdevelopment
hasbegunatthetop,butitwouldnothaveproceededanyfurtherthanwhatcanbeseen
here.Exogenousbuddingisoccurringattheotherend.
Chapter03


Figure3.1GulllakeadjacenttotheW.K.KelloggBiologicalStationinMichigan.
Researchersatthelabhadprimaryaccesstotheboatpresentintheforeground.
Chapter04
Figure4.1ParPondontheSavannahRiverPlantsiteinSouthCarolina.Inthe
foregroundisa“shockingboat.”Thecraftwasequippedwithageneratorfor
producingelectricityusedforstunningfishinwateraheadoftheboat.The
electroshockedfisharestunnedandrisetothesurfacewheretheyareremovedfromthe
waterwithadipnetandthendepositedinalargetubcarriedintheboat.Inanhourof
shocking,50largemouthbasscouldbeeasilycaptured,withoutkillingthem.
Chapter06
Figure6.1Charlie’sPondinNorthCarolina.Thepondwasprobablycreatedduring

constructionofthecoolingreservoir(BelewsLake)in1970.Ourresearchfirstbegan
inthepondduring1983whenAmyCrewsandIbegancollectingsnails.
Figure6.2TwocercariaeofHalipegusoccidualis.Thedeliverytubeandbodyofthe
trematodearestillinsidethetail.
Figure6.3Inanexplosiveepisode,adeliverytubeemerges,andsimultaneously,the
bodyofthecercariaisshotthroughthetubeandintothehemocoelofanostracod.
Figure6.4Usingitsventralsucker,theadultHal.occidualisattachestotissueatthe
bottomofthemouthandunderthetongueofgreenfogs,Ranaclamitans.
Figure6.5Agreenfrog,R.clamitans.
Figure6.6ThelifecycleofHal.occidualis.
Chapter08
Figure8.1AgroupofstudentstakealookatBelewsLake.Charlie’sPondislocated
behindthem,about20 maway.
Chapter09
Figure9.1Lepomismacrochiruswithblackspot,eachofwhichisametacercariaof
Uvuliferambloplitis.
Chapter11
Figure11.1LesionsonthesurfaceofMicropterussalmoides,causedbyAeromonas
hydrophila.
Chapter12
Figure12.1MaleandfemaleDaubayliapotomaca,rhabditidnematodesofHelisoma
ancepsinMallardLake,NorthCarolina.(a)Thefemalesareslightlylargerthanmales
andareeasilydistinguishedbythehooklikeposteriorendofthebody.(b)Males
possessagubernaculumandspicule.


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Chapter13
Figure13.1Rarefiedspeciesrichness.Meanspeciesrichnessand95%confidence
intervalswerecalculatedusing2000replicates(randomsampleswithoutreplacement)

forasamplesizeof275(thesamplesizefor2013).Thedashedlinemarksthespecies
richnessforthe2013sample.Rawdatawereusedfor1984,2002,2006,2012,and
2014.
Figure13.2(a)EastbankofCharlie’sPondshowingemergentvegetationin1988.
(Thislocationmatchessite24inZelmeretal.(1999).)(b)WestcoveofCharlie’s
Pondshowingvegetationin1988.(Thislocationmatchessite15inZelmeretal.
(1999).)(c)EastbankofCharlie’sPondwithoutvegetationin2014.(d)Northwest
coveofCharlie’sPondwithoutvegetationin2014.

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EcologicalParasitology:Reflectionson50Yearsof
ResearchinAquaticEcosystems
GeraldW.Esch
CharlesM.Allen,ProfessorofBiology
DepartmentofBiology
WakeForestUniversity
Winston-Salem,NorthCarolina,USA






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Dedicatedtothememoryofmyfriend,ProfessorRalphD.Amen,whodiedjustafewmonths
agofollowingafriendshipofsome50years.Unfortunately,my“personaleditor”wasnot
abletohelpmethistime.


Preface

Fromthebeginningofmyacademiccareernearly50yearsago,Ihavebeenblessedbystrong
associationswithqualitygraduateandundergraduatestudents.Throughoutthisperiod,ithas
beenmypleasuretowatchthemsuccessfullyapplytheireffortsinboththefieldandthe
laboratory.
Overthepastseveralyears,Ihavegivenseriousthoughttowritingabouttheirresearch
experiencesandmyconnectiontothem.WardCooperwasaSeniorCommissioningEditorat
Wiley-Blackwell.Ihaveknownhimformanyyears,andhehasenthusiasticallyhelpedme
withotherprojectsalongtheway.Acoupleofyearsago,IpersuadedWardthatstoriesI
wantedtotellaboutmystudentsandtheirresearchwereestimableandworthyofincorporating
intoabook.Heagreedandwasabletoconvincethe“editorialgroup”atWiley-Blackwellto
supportitspublication.
Necessarily,someofwhatIhavewrittenforthepresentbookalsoinvolvesmyprofessional
beginning,makingthefirstcoupleofchaptersseemlikethebookwillbeautobiographical,but
itisn’t.AsIwillemphasizelater,thereisnowayofseparatingmycareerfromthebeginning
oftheirs.Theyarenaturallyintertwined.Ihavealsoincludeddescriptionsofcontributions
madetomycareerandtothoseofmystudentsbysomeofmynonstudentcollaborators,
mentors,andcolleagues.Thegroupincludessomereallygreatpeople,forexample,Robert
(“Doc”)Stabler,MaryAliceHamilton,J.TeagueSelf,JimMcDaniel,MacWilsonWarren,Jim
Hendricks,GeorgeLauff,DesmondSmyth,CliveKennedy,WhitGibbons,DarwinMurrell,Al
Shostak,RayKuhn,RonDimockJr.,DickSeed,andAlBush.Allofthesepeople,plussome
otherswhoIwillidentifyinthetext,havetouchedmypersonalandprofessionallifeinone
wayoranother,andIfeelthatIamindebttothemforcontributingtowhateversuccessesmy
studentsandIhavehadalongtheway.
Foremost,however,thoughtsregardingmycareeralwaysreturntomystudentsandwhatthey
didinthewayofresearchwhentheyweregraduatestudentswithmehereatWakeForest.
Withoutexception,theywerebright,self-starters,hardworking,innovative,andloyal.During
mycareer,ItakeprideinknowingthatIhavehadjusttwostudentsleavemylabwithout
achievingthesuccessofsecuringtheiradvanceddegree.
In1966,theAmericanSocietyofParasitologistsheldourannualmeetinginSanJuan,Puerto
Rico.Irecallitquitewell.Foronething,itwasthefirsttimeIhadeverflown.Mywife,Ann,

andIdrovetoBaltimoreandstayedwithourfriends,BillandSallyCline,thenightbeforewe
weretoheadforSanJuanaboardanEasternAirlinesElectraturbopropaircraft.Mywifewas
aveteranflyerandfeltitwouldbebesttodrivetoBaltimoreandthenflyfromthererather
thantakingmyfirstplanetripinasmalleraircraft.Moreover,Billwasaphysicianand
promisedtoprovidemewithsomesortofremedythatwouldcalmmynervesonflightday.
Unfortunately,onarrivalattheairport,welearnedthatawindshieldontheplanehadcracked
onlandinginNewarkonitswaytoBaltimoreandwasaboutfourhourslate—bythattime,the


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effectsofthedrughadwornoff.Imustadmitthatitwasstillagreatfirstflight!
ThebestpartoftheSanJuanmeetingwasadinnerontheverandaoftheAmericanaHotelin
whichwewerestaying.Dr.andMrs.Self,JohnandKarenJanovy,JimandSueMcDaniel,
HoraceandJayneBailey,andmywife,Ann,andwerethere(Figure0.1)(John,Jim,Horace,
andIwereallDr.Self’sgraduatestudents).Theeveningwaswarm,butnothot.Therewasa
gentlebreezeblowinginofftheocean,andthesunwassettinginthewest.Inotherwords,it
wasbeautiful!Dr.Selfwasfeelingprettygood(hewasascotchdrinker)butwasinverygood
control.Wetalkedabouttheimportanceofprofessionalmeetingsandwhywestudentsshould
attendregularly.IrememberDr.Selfnostalgicallylookingaroundthetablebeforesaying,“I
reallyenjoycomingtothesemeetings.Itgivesmetheopportunitytobaskinthereflectedglory
ofmystudents.”Thisstatementistheperfectdescriptionofwell-deservedprideinthesuccess
astudentbringstotheirmentor,whetherataprofessionalmeetingorintheresearchthey
accomplish.Ihaveneverforgottenthisfeeling.ItisanotherreasonIwantedtowritethisbook.

Figure0.1AphototakenattheAmericanaHotel,SanJuan,PuertoRico,November1966,41st
annualmeeting,theAmericanSocietyofParasitologists.Fromlefttoright:me,AnnEsch,
HoraceBailey,JayneBailey,SueMcDaniel,JimMcDaniel,Mrs.IdaSelf,Dr.J.TeagueSelf,
andKarenJanovy.
PhotocourtesyofJohnJanovy,Jr.

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Acknowledgments
Iwanttoespeciallythankmymanycolleagueswho,overtheyears,haveencouraged,guided,
andadvisedmeineverywaypossible.Theirgifttometaughtmetogiveasmuchtomy
students.Asmentoringmodels,Icouldnothavehadabetterbunch.
Ialsowanttothankmystudentsfortheircooperationinhelpingmetowritethistome,in
correctinganumberoferrors,andinremindingmeofthingsthatIhadomitted.Theircompany
andfriendshiphavebeenoutstandingeversincetheywalkedintomylabandparticipatedwith
mewhetheritwasatColoradoCollege,theUniversityofOklahoma,WakeForestUniversity,
KelloggBiologicalStation,ImperialCollegeinLondon,ortheSavannahRiverEcology
Laboratory.Myfinalgraduatestudent,KyleLuth,wasparticularlyhelpful.Hereadtheentire
bookandofferedabushelfullofsuggestions,mostofwhichIgratefullyaccepted.Justbefore
submittingthefinalversiontoWiley-Blackwell,Iaskedalong-timefriend,Dr.RichardSeed,
toreadthebookandofferanysuggestionshemighthaveforimprovingthecontent.Hewas
veryhelpful.
ThepeopleatWiley-Blackwell,especiallyWardCooperandKelvinMathews,helpedputit
together.Myprojectmanager,JayavelRadhakrishnan,providedverygoodtechnical
assistance.
Finally,Ithankmychildren,Craig,Lisa,andCharlie,whowillinglyaccompaniedmealmost
allthewayonthislongjourney.Mostespecially,Iamgratefultomywonderfulwife,Ann,
whohasbeensofantasticthroughoutourlives.Sheisoneofmybest,andtoughest,critics.
Thoughnotascientist,shereadtheentirefinaldraftandofferedasignificantnumberofideas
forimprovingthecontentandthewayIpresentedit.


ThePlayers
Thereneverwasintheworldtwoopinionsalike,nomorethantwohairs,ortwograins;

themostuniversalqualityisdiversity.
OftheResemblanceofChildrentoTheirFathers,MicheldeMontaigne(1533–1592)

AsIapproachedtheendofthepresentmanuscript,Irealizedthatthebookwasnotjusta
successionofindividualstudentstories;inmanycases,theywereclearlylinkedtooneanother.
ItwaslikemystudentsandIhadtraveledtogetheronajourney.Aseachofthemcameonthe
scene,theystayedwithmeforawhileandthendepartedtobegintheirowntravels.Afew
haveretired,butmostarestillmakingtheirway.Manyofthemhavebecomeconnectedtotheir
ownstudentsforpartoftheirtripsinthesamewaytheyjoinedwithmeandIwithDr.Self,my
graduateschoolmentor.IhopeIhavebeenassuccessfulasarolemodelasheandDr.Stabler,
myundergraduateguide,wereforme.Imustemphasizeagainthatthebookisnotan
autobiography—itisacollectionofintertwinedbiographies,withmineincluded.
Early,whilewritingtheirstories,Ibeganpreparingmicrobiographiesforeachofstudentwho
hadtakenanecologicalroute—therewereseveralwhodidnotfollowinthisresearch
directionandarenotincluded.Theoriginalideawastoincludetheirnarrativesattheendof
thebook,butIdecidedtomovethisinformationtothebeginning.Thisway,youthereader
wouldhaveatleastanideaabouteachofthemasIbegintotelltheirstories.
Duringmy50yearsasafacultymemberhereatWakeForestUniversity,Ihavehadthe
privilegeofmentoring24masterofscience(MS)and19PhDstudents.OftheMSstudents,
fivehavecontinuedonfortheirPhDdegreeswithme.Unfortunately,Ilosttrackofseven
studentsaftertheirgraduation.However,Iknowthatatleastoneinthisgroupbecamea
dentist,anotherbecameaphysician,twoarehousewives,oneearnedaPhDelsewhere,and
twoothersworkinthecorporateworld.
Iconsiderallofmystudentsasclosefriends,andIamtrulypleasedtoseeagreatmanyof
themeveryyearattheannualmeetingoftheAmericanSocietyofParasitologists.RayKuhn,
mylongtimefriendandcolleague,hasgraduatedaboutthesamenumberofstudentsasI,anda
greatmanyofhisformerstudentsattendtheannualmeetingaswell.Ithinkthelargest
combinednumberwas17,attheASPmeetinginColoradoSpringsin2010.Oneadvantageof
havingsuchalargegroupistheextentof“networking”thatisaccomplishedbythestudents
aftertheyleavehere—itreallymeansalot!

Microbiographies
JohnTrainer
Johnwasmyfirstgraduatestudent—hewasinfactwaitingformewhenIarrivedatWake
Forestinthefallof1965.So,webeganourjourneytogether.Althoughhisresearchdidnot


involveparasitesinaquaticecosystems,collectingforhisthesiswasaccomplishedinthe
field.AfteracquiringhisMSdegree,hewentontotheUniversityofOklahomawherehe
earnedhisPhDwithDr.J.TeagueSelf(mymentoringraduateschool).John’sfirstacademic
positionwasatJacksonvilleUniversity(Jacksonville,Florida)wherehemovedrapidly
throughtheacademicranks,endingupasvicepresidentforacademicaffairsanddeanof
facultyafterjust9yearsinresidence.HiscareerasanadministratorthentookhimtoLenoir–
RhyneCollege(Hickory,NorthCarolina)aspresident.Eventually,hewastobecomepresident
andheadofTheBollesSchoolinJacksonville,oneofthefinestK–12privateschoolsonthe
eastcoastoftheUnitedStates.AfterhisretirementatTheBollesSchool,hereturnedto
JacksonvilleUniversitywherehepresentlyservesastheseniorcampaignofficer.Herecently
sentmeayearbookthatincludedphotographs,warmreminiscesandrecollections,etc.,dealing
withhisretirementgalaatBolles—whatanexcitingexperiencethatmusthavebeenforhim
andhiswonderfulwife,Alice!

Robert(Bob)Morcock
BobwasoneofmyfirstPhDstudents.Hebeganhisdoctoralworkin1969rightafterthePhD
programwasestablishedinourDepartmentofBiologyatWakeForest.Hisdissertation
researchwasfocusedontheintermediarycarbohydratemetabolismoflarvalTaenia
crassiceps(aboutas“nonecological”asonecouldget).AlthoughIhadnotyetmadetheswitch
toecologicalparasitology,Iwasabletoprovidehimsomehelpsincemyowndissertation
workwasaimedatcomparativeintermediarycarbohydratemetabolisminlarvaeandadultsof
Taeniamulticeps.HeisnonethelessincludedinmyecologystudentgroupbecauseitwasBob
whoinadvertentlystartedourworkonthepopulationbiologyofCrepidostomumcooperiin

GullLakeduringitseutrophication/recoveryprocess.Aftercompletinghisdegree,heobtained
ateachingjobatHoodCollege(Frederick,Maryland).Hestayedthereforarelativelyshort
timebeforegoingtoworkattheEnvironmentalProtectionAgency(EPA)inWashington,DC,
wherehespenttherestofhiscareer,retiringjustrecently.

JohnAho
JohnobtainedhisBSandMSdegreeshereatWakeForest.IrememberrecruitingJohnintomy
labwhenhewasasophomore.Ineededsomeone(preferablyastudent—becausetheycame
relativelycheap)tocareformymousecolony.Whileteachingacourseincellphysiology,I
walkedintomyteachinglabonedayandaskediftherewassomeonewhowouldliketomake
afewextradollarsbycaringformyanimals.John’shandshotupimmediately.Hestayedwith
meforseveralyearswhilecompletinghisundergraduateworkandhisfirstfewyearsasa
graduatestudent.OncompletionofhisMSdegree,hesecuredaFulbrightscholarshipandused
ittoobtainhisPhDdegreeattheUniversityofExeter(England)underthetutelageofDr.Clive
Kennedy.Iwaseveninvitedtobehisexternalexaminerforhisdissertation.Aftercompleting
hisPhD,JohnthenspenttimewithDr.JohnHolmesattheUniversityofAlberta(Edmonton,
Alberta,Canada)asapostdocandbackhereatWakeForestasavisitingprofessorbefore
headingforAuburnUniversity(Montgomery,Alabama).


Joseph(Joe)Bourque
IfirstmetJoewhenIwasteachingfieldparasitologyattheW.K.KelloggBiologicalStation
(HickoryCorners,Michigan)ofMichiganStateUniversity.JoecamefromtheUniversityof
Illinois(Champaign,Illinois)toWakeForesttofirstdoanMSdegreeandthenhisPhD.He
wasoneofmyfirsttwo“genuine”parasiteecologystudents.Hisresearchwasconductedat
theSavannahRiverEcologyLaboratory(Aiken,SouthCarolina),wherehestudiedtheeffects
ofthermaleffluentstressonthepopulationandcommunitybiologyofhelminthparasitesin
yellow-belliedturtles.AftersuccessfullycompletinghisdissertationresearchandhisPhD
degree,hejoinedthePeaceCorpsandservedinsoutheasternAsia.Whenhereturnedhome
fromthePeaceCorps(withawife),hefirstworkedasamedicaltechnicianatNorthwestern

University(Chicago,Illinois)andthenwasfinallywasadmittedintotheUniversityofIllinois
MedicalSchool(Urbana,Champaign,Illinois)wherehepursuedanMDdegree.Hespenthis
careerasathoracicsurgeoninCalifornia.

James(Jim)Coggins
JimCogginsobtainedhisBAandMAdegreesatEastCarolinaUniversity(Greenville,North
Carolina),whereheworkedwithDr.JimMcDaniel.Thelatter,Jim,andIhadbeengraduate
studentstogetherattheUniversityofOklahoma(OU)(Norman,Oklahoma).Hewasthefirst
graduatestudentImetwhenIarrivedonthecampusattheOUin1958,andwebecamelifelong
friends.Veryregrettably,hediedwaytooyoung,andIstillmisshiminsomanyways.Jim
CogginscametoWakeForesttoworkonhisPhD.Hisresearchwasfocusedonthe
plerocercoidstageofProteocephalusambloplitis,theso-calledbasstapeworm,withprimary
interestonthefinestructureandhistochemistryoftheapicalendorganofthisunusual
tapeworm.DuringonesummerofhisstayatWakeForest,JimaccompaniedmetoKBSwhere
heservedasmyteachingassistant.HeandIdidsomecollaborativeworkusingwhatwe
called“tethers,”whichinrealitywerewide-meshbasketsmadeofheavytwineinwhichwe
couldplaceuninfectedbluegillscollectedfromanearbypond.Weplantedthe“tethers”in
selectedlocationsoftheGullLakelittoralzonesothatwecouldlearnsomethingaboutthe
ratesofhelminthrecruitmentandturnoverundercontrolledconditions.Itwasaninteresting
undertaking.WhenJimfinishedhisdissertationwork,heheadedofftotheUniversityofNotre
Dame(NotreDame,Indiana)andapostdocwithDr.PaulWeinstein.Hecontinuedasuccessful
careerattheUniversityofWisconsin–Milwaukee(Milwaukee,Wisconsin),fromwhichheis
nowretired.

HermanEure
IwriteatlengthaboutHermaninthetextofChapter7,soIwillnotrepeateverythinghere.
HermancametoWakeForestwithhisundergraduatedegreefromtheUniversityofMaryland–
EasternShore(PrincessAnne,Maryland)ononehandandaprestigiousFordFoundation
Fellowshipontheother.Hedecidedtoskipthemaster’sdegreeandheadstraightforhisPhD.
HisresearchwasconductedonthepopulationbiologyofthebasstapewormP.ambloplitisand

theacanthocephalanNeoechinorhynchuscylindratusinParPondontheSavannahRiverPlant


(Aiken,SouthCarolina).TheyearhefinishedhisPhD,IleftWakeForesttotakealeaveof
absenceattheSavannahRiverEcologyLaboratory(SREL),andHermanwasappointedasmy
“temporary”replacement.Onmyreturnthenextyear,Hermanwashiredasatenure-track
facultymemberinourDepartmentofBiology.Asourgraduateschool’sfirstAfricanAmerican,inadditiontohisfacultyresponsibilities,hewasemployedinavarietyof
administrativepositionsbeforebecomingourDepartmentofBiology’schairman,wherehedid
asuperbjob.HesubsequentlymarriedKelliSappandrecentlyretiredasafacultymemberin
thecollege.Heis,however,stillactiveintheAmericanSocietyofParasitologists,forwhich
heandKelliserveasco-programofficers.

TerryHazen
TerryobtainedhisBSandMSdegreesininterdepartmentalbiologyfromMichiganState
University(EastLansing,Michigan).IfirstmetTerrywhenhecametotheKelloggBiological
Station(KBS)totakemyfieldparasitologycourseinthesummerof1973.Heaccompaniedme
totheSRELasmytechnicianwhileIwasonleavefromWakeForestduringthe1974–1975
academicyear.WhenIreturnedtoWakeForestinthefallof1975,hecamewithmetopursue
hisPhD.HisdissertationresearchfocusedontheecologyofAeromonashydrophila,the
causativeagentforredsorediseaseinfish.Whilehedidsomeparasitologicalresearchearly
on,Terrybecameaninternationallyrenownedmicrobialecologist,withstronginterestsin
bioremediationandbioenergytechnologies.Throughouthiscareer,hehasheldseveral
academic/administrativepositionsofresponsibilityattheUniversityofPuertoRico,the
SavannahRiverPlant,andtheE.O.LawrenceBerkeleyNationalLaboratoryinCalifornia
(Berkeley,California).HeispresentlytheUniversityofTennessee/OakRidgeNational
LaboratoryGovernor’schair(OakRidge,Tennessee)andprofessorintheirDepartmentof
MicrobiologyandtheDepartmentofEarthandPlanetarySciences(Knoxville,Tennessee).

Joseph(Joe)Camp
JoereceivedBSandMSdegreesatIllinoisStateUniversity(Normal,Illinois),wherehe

workedundertheguidanceofalongtimefriend,Dr.HarryHuizinga.HecametoWakeForest
in1977topursueaPhDdegreeinmylab.Hisresearchinvolvedastudyonthepopulation
biologyofTylodelphys(Diplostomulum)scheuringiinmosquitofish(Gambusiaaffinis)in
ParPond,ontheSavannahRiverPlantsite.Onreceiptofhisdegree,hetookapositionat
PurdueUniversityNorthCentral(Westville,Indiana).HesubsequentlytransitionedtoPurdue’s
maincampus(WestLafayette,Indiana),whereservesasaprofessorofveterinarymedicine
andsecretaryoffaculties.
AverycoincidentaloccurrencetookplacewhenItraveledtoLawrence,Kansas,tovisitwith
severalpeoplewithwhomIwouldworkatAllenPress(AP)afterbeingappointedsuccessor
toBrentNickolaseditoroftheJournalofParasitologyin1993.Wewereatlunchandavery
niceladywassittingacrossthetable.IaskedherwhatshedidatAP,andsherepliedthatshe
wasacopyeditorforourjournal.Iinquiredifthiswasheronlyjobandshereplied,“Oh,no.I
amalsotheDeanoftheCollegeofLiberalArtsandSciencesattheUniversityofKansas”in
Lawrence;IalmostchokedonthebiteofsaladIwastryingtoswallow.Iaskedwhyinthe


worldshewouldholdthesetwosortsofjobs.Sheexplainedthatsheenjoyedworkinginthe
eveningwhilesittinginacomfortableeasychairusingawritingboardtoeditwhilelistening
tobeautifulmusic—it“wasrelaxing,”sheremarked.Ilaterlearnedthatshe(SallyMason)had
becomeprovostatPurdueUniversity.ItwasshewhoinvitedJoeCamptocomedowntothe
maincampustobecomesecretaryoftheFacultySenate,aswellasprofessorintheirSchoolof
VeterinaryMedicinewherehealsoteachesparasitology.SallyMasonispresentlythe
presidentattheUniversityofIowa!

Willard(Bill)GranathJr.
BillcompletedhisundergraduatedegreeatDelawareValleyCollegeofScienceand
Agriculture(Doylestown,Pennsylvania)andthenhisMSdegreeatIllinoisStateUniversity
(Normal,Illinois)whereheworkedwithHarryHuizinga.JoeCampandBillhadbecome
friendsatIllinoisStateandJoestronglyrecommendedthatBillcometoWakeForestforhis
PhDdegree.Heappliedandwasaccepted—agreatmoveonourpart!Hisdoctoralresearch

wastofocusonthepopulationecologyoftheAsiantapewormBothriocephalusacheilognathi
inmosquitofishinanearbycoolingreservoiroperatedbyDukeEnergy.Iamespeciallyglad
thatBillcamehere,becauseIseriouslydoubtthatanyresearchonB.acheilognathiwould
haveeverbeenundertaken(infact,Idonotbelievethatwewouldhaveknownaboutthe
presenceofB.acheilognathiinthereservoirbecauseIstillcannotseehowthistapewormcan
fitintothegutofsuchsmallfish!).Moreover,thefishfaunahadbeensignificantlyreducedvia
seleniumpollution,andtheonlypiscinespeciesleftinthemainbodyofthelakewereafew
carp,channelcatfish,andmosquitofish;whilestilllargelydevoidoffish,fatheadminnowsand
redshinerswereintroducedintothelake.ThoseofuswhoworkedinBelewsfigurethata
frustratedbassfishermanwhocouldnotcatchanythinginthemainbodyofthelake,letalone
getabite,“donated”theirbaitfishtothelakeafterafrustrating“no-catch”day.Oncompletion
ofhisPhDdegree,BilldidpostdoctoralworkwithDr.TimYoshinoattheUniversityof
Oklahoma(Norman,Oklahoma)beforeheobtainedatenure-trackpositionattheUniversityof
Montana(Missoula,Montana).Healsoswitchedhisresearchfocustothecnidarian
Myxoboluscerebralisandhasbecomealeadingworldexpertonwhirlingdiseaseinsalmonid
fish,definitive(final)hostsfortheparasite.

AmyCrews
AmystartedasanundergraduateatEmoryUniversity(Atlanta,Georgia)beforetransferringto
WakeForestwhereshecompletedherBSandMSdegrees.Shewasthefirsttoworkin
Charlie’sPondandsetaveryhighstandardforthestudentswhofollowedherintooursmall
impoundment.ShebeganherPhDstudieswithDr.TimYoshinoattheUniversityofOklahoma
andfinishedattheUniversityofWisconsinSchoolofVeterinaryMedicine(Madison,
Wisconsin)afterTimmovedfromOklahoma.ShethendidpostdoctoralworkwithBillCollins
attheCentersforDiseaseControl(CDC)inAtlanta.Amyacceptedateachingpositionatthe
UniversityofNorthernAlabama(Florence,Alabama)whereshepresentlyworksasdirector
oftheirHealthProfessionsCommittee.


Timothy(Tim)Goater

TimwasthefirstofthreeCanadiangraduatestudentstocomedowntotherelativelywarm
southofNorthAmerica.HisundergraduatementorwasDr.AlBushatBrandonUniversity
(Brandon,Manitoba,Canada).Withoutquestion,Timcouldnothavehadamoreinterestingand
knowledgeablepersontointroducehimtotheecologicalaspectsofparasitologythanAlBush,
who,regrettably,diednotverylongagoataveryyoungage.Forhismaster’sresearchatWake
Forest,TimchosetoexaminethecommunityecologyofsalamandersinstreamsoftheSmoky
MountainsofsouthwesternNorthCarolinaandTennessee.HestayedontodohisPhDatWake
ForestandproducedawonderfuldissertationdealingwiththetrematodeHalipegus(=Hal.)
occidualisinCharlie’sPond.ItwasTimwhoalsodiscoveredthecolonizationofPhysa
gyrinaandHalipeguseccentricusintothepond.Oncompletionofhisdissertation,hebecame
afacultymemberofwhatisnowknownasVancouverIslandUniversity(Nanaimo,British
Columbia,Canada).
In2001,AlBush,JackieFernandez,DickSeed,andIcoauthoredabooktitledParasitism:
TheDiversityandEcologyofAnimalParasites.Afewyearsago,CambridgeUniversity
Press,whichpublishedthefirsteditionofthebook,approachedmewithaninvitationtowrite
asecondedition.ItoldthemthatIwould,butIneededtoselectacoupleofnewcoauthorsto
assistsinceAlhaddiedandDickandJackiehadretired.Cambridgegavemethe“go-ahead,”
andIaskedTimandCamGoater(brothers)iftheywouldbeinterested(Irecalledanother
brotherduo,ElmerandGlennNoble,whohadwrittenanotherparasitologytextbookseveral
yearsagoandhowsuccessfulitwas).TimandCamacceptedtheinvitation.Theyassumedthe
leadinrewritingthebookand,byfar,didmostoftheworkingettingtherevisioncompleted.
WhileIwaswritingthissectionofthepresentbook,acopyofthe2ndeditionarrived(Goater
etal.,2014).Itisanexceptionallyfinepieceofwork(ifIdosaysomyself!).

David(Dave)Marcogliese
Davewasthesecond“Canuck”tocomesouth.HeoverlappedwithTim,andtogether,they
wereexcitingguystohavearound—lotsandlotsofenthusiasmandenergyfromthepair.Dave
didhisundergraduateworkatConcordiaUniversity(Montreal,Quebec,Canada)andthen
movedtoDalhousieUniversity(Halifax,NovaScotia,Canada),wherehecompletedhisMS
HeworkedfortheDepartmentofFisheriesandOceansforthreemoreyearsbeforecomingto

WakeForesttobeginhisPhDstudies.WhilehedidsomeworkonB.acheilognathi,mostof
hisdissertationresearchdealtwiththebiologyofcopepods,whichserveasfirstintermediate
hostsfortheparasite.DavehassincegoneonwithafantasticcareerinEnvironmentCanada
(somethingliketheDepartmentofInteriorintheUnitedStates).Hisresearchovertheyears
broughttherecognitionhedeservedfromtheCanadianSocietyofZoologistswithhisreceipt
oftheirRobertArnoldWardleAwardandtheHenryBaldwinWardMedalfromtheAmerican
SocietyofParasitologists.

Jacqueline(Jackie)Fernandez
JackiecametoWakeForestfromChile.Shehadappliedtoworkwithmygoodfriend,thelate


JohnCrites.However,Johnwasintheprocessofretiringandrecommendedthatshecometo
WakeForestanddoherdissertationresearchwithme,whichshedid.Whenshearrived,she
hadmorethan20publicationsandbothundergraduateandMSdegreesfromtheUniversityof
Concepcion(Concepcion,Chile).Shefitintomylaboratorybeautifully.Jackieisnotonlya
charmingandintelligentyoungwoman,butalsosheishighlyenergeticandaveryhardworker.
Herresearchmostlyinvolvedtheinfracommunityandcomponentcommunitydynamicsof
trematodesinHelisoma(Hel.)ancepsinCharlie’sPond.WhilehereatWakeForest,shemeta
young,formermarinewhowasworkingtoward(successfully)hisPhDdegreeinthe
DepartmentofChemistry.Theyweremarriedandhadtwoboyswhoarenowintheprocessof
completingtheircollegeeducations.Jackieispresentlyteachinghighschoolbiologyin
VirginiaandwasrecentlyrecognizedasteacheroftheyearbytheNationalAssociationof
BiologyTeachers,areallygreathonor!

A.DennisLemly
DennisstartedoutwithastronginterestinwildlifebiologyandaBSdegreefromWestern
CarolinaUniversity(Cullowhee,NorthCarolina).Hisinitialcareergoalwastobecomea
wildlifeandfisheriesofficer.Towardthatend,hegraduatedwithanappliedsciencedegree
fromtheHaywoodCommunityCollege(Clyde,NorthCarolina)in1975.Then,heconsidered

teachingasaprofessionandgraduatedfromWakeForestUniversitywithanMAEd.degree.
Denniswasasuperiorstudentandwasluckythathealreadyhadhisdissertationresearchset
upwhenhebeganhisPhDprograminmylab.Inthesecondyearofhismaster’sdegreework,
hetookmyparasitologycourse,andhisfocuschangedagain,thistimetoparasitologyand
ecology.ForhisPhDresearch,hecenteredonthepopulationbiologyofUvuliferambloplitis,
thecausativeagentof“blackspot”infish.Tomywayofthinking,hisdoctoralresearch
representsaclassiccontributiontounderstandingtheconceptofoverdispersionbyparasite
infrapopulations.Healsobeganalifelongcareerconcentratingontheeffectofselenium
pollutioninaquaticecosystemsandhasbecomea“world-class”expertinthisareaof
research.HeispresentlyemployedbytheUSFishandWildlifeServiceandhasan
appointmentasresearchassociateprofessoratWakeForest.

Michael(Mike)Riggs
MikeRiggsobtainedhisundergraduatetrainingattheUniversityofNewHampshire(Durham,
NewHampshire)andthenhismaster’sdegreeatIowaStateUniversity(Ames,Iowa)wherehe
workedwithDr.MartinUlmer.HecametoworktowardhisPhDinmylabatWakeForestin
1979.HisverystrongresearchwasconductedontheAsiantapewormB.acheilognathiin
severalminnowspeciesatBelewsLake.AboutmidwaythroughhisPhDdegreeprogram,
MikedecidedtoalsopursueamasterofpublichealthdegreewithsupportfromaUSPublic
HealthServicetraininggrantinbiostatisticsattheSchoolofPublicHealth,theUniversityof
NorthCarolina–ChapelHill(ChapelHill,NorthCarolina).HereturnedtoWakeForestwhere
hecompletedhisdissertationin1986.HeworkedasastatisticianwiththeDivisionofFish
andWildlife,MinnesotaDepartmentofNaturalResources(Minneapolis,Minnesota)and
returnedasaseniorenvironmentalstatisticianforResearchTriangleInstitute(RTI)inthe


Free ebooks ==> www.Ebook777.com
ResearchTrianglePark,NorthCarolina.MikeisnowlivingbackinNewEngland.

EricWetzel

EricisaPennsylvanianativewhodidhisundergraduatedegreeatMillersvilleUniversity
(Millersville,Pennsylvania)beforecomingtoWakeForest.Hebeganasamaster’sstudentof
Dr.PeterWeiglinourDepartmentofBiology.Petewasalongtimeworkeronflyingsquirrels
inthenearbyAppalachianMountains,andthesewerethe“critters”towhichEricfirstbecame
attached.Eric’sfocuswasonaspeciesofStrongyloidesthatinfectsthesquirrels,andthisis
whatledhimintoparasitologyandmylabaftercompletinghisMSdegree.HisPhDresearch
tookhimintoCharlie’sPondandanemphasisonHal.eccentricus.Aftercompletinghis
degree,hestayedatWakeForestforanotheryearandahalfasavisitinginstructorbefore
securingateachingpositionatWabashCollege(Crawfordsville,Indiana).Itwasaperfectfit
forEricsinceWabashisoneofthefinestliberalartscollegeseastoftheMississippiRiver,
andthisisexactlywhathewantedintermsofateachingcareer.Recently,Erichasenteredinto
anotherareaofparasitology,onethathasledhimintoPeruinSouthAmericawhereheis
focusingoninfectiousparasiticdiseaseresearchamongthepoorestofpeopleinthatpartofthe
WesternHemisphere.HeispresentlytheNormanE.TrevesprofessorofBiologyatWabash.

DerekZelmer
DerekisthethirdCanadiantocomesouth.HisBSandMSdegreeswerebothobtainedatthe
UniversityofCalgary(Calgary,Alberta,Canada),wherehestudiedwiththelateDr.H.P.Arai.
HisPhDdegreewasbroadlyfocusedonseveralaspectsofthelifecycleandpopulation
biologyofHal.occidualis.IncollaborationwithEricWetzel(mentionedearlier),Derekwas
abletoestablishthatthelifecyclesofHal.occidualiscouldbecenteredatfourdifferent
‘hotspots’inthepond,allofwhichhadsimilarmicrohabitat,orlandscape,characteristics.He
alsoclearlydeterminedthatodonatenaiadsarenecessaryparatenichostsfortheparasiteand,
finally,thattransmissionofHal.occidualisinCharlie’sPondisgovernedbyprevalenceand
notintensityofadultparasites.WhenDerekfinishedhisPhDresearch,hewasabletosecurea
teachingpositionatEmporiaStateUniversity(Emporia,Kansas),anopportunitythatbothhe
andIthoughtwasrightforhim.Unfortunately,wewerebothwrongandhislifeinEmporia
wasmiserable.After6years,however,hefoundamuch,muchbetterpositionattheUniversity
ofSouthCarolina–Aiken,andheisquiteathomeinhisrecentlyfoundsurroundings.

KymJacobson
KymcametoWakeForestwithaBSdegreefromtheUniversityofNevada–Reno.She
completedhermaster’sdegreewithmeandthenswitchedintoimmunologyandobtainedher
PhDwithRayKuhnhereatWakeForest.Hermaster’sdegreeinvolvedthecommunitybiology
ofenterichelminthsintheyellow-belliedsliderturtlesdownattheSREL.Shedidnot,
however,enjoyperformingturtlenecropsies,whichledherintoRay’slab.AfterherPhD,the
nextstopwasattheMayoClinic(Rochester,Minnesota)andapostdocinneuroimmunology.
KymthenspenttimeasapostdocattheSeattleBiomedicalResearchInstituteoftheUniversity
ofWashington(Seattle,Washington).Despitehersuccessinimmunology,Ialwayshada“gut

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feeling”thattheecologicalaspectsofparasitologywereherfirstlove,andIwascorrect.
Accordingly,sheacceptedapostdoc,followedbyafull-timestaffposition,asafish
ecologist/parasitologistattheNationalOceanicandAtmosphericAdministration(NOAA)
LaboratoryinNewport,Oregon.Iamsoproudthatshehascreatedasolidreputationfor
herselfinmarineecologyandparasitology.

JulieWilliams
JulieWilliamscompletedherundergraduatedegreewithDr.BrentNickolattheUniversityof
Nebraska–Lincoln(Lincoln,Nebraska)whereshedevelopedaninterestinparasitology.She
cametoWakeForestandsuccessfullypursuedherMSdegreeinmylab.Duringherlastyear
here,sheexplainedthatshewantedtobecomeaphysician,andthatiswhatshedid!She
attendedtheBowmanGraySchoolofMedicine(nowtheWakeForestUniversitySchoolof
Medicine)andisnowsuccessfullyemployedthereintheirDepartmentof
Geriatrics/Gerontology.

ScottSnyder
ScottwasanotherstudentfromtheUniversityofNebraska–Lincoln,wherehewas

coaxed/persuadedintoparasitologybymyoldfriend,Dr.JohnJanovyJr.Scottwassuccessful
inhisMSdegreeresearchhereatWakeForest.HisresearchwasfocusedonHal.eccentricus
inP.gyrinainCharlie’sPond,workthatprovidedareallyexcellentcomparisonwithits
sympatriccongenerinourpond.HereturnedtoNebraskaandJohnwhereheperformedsome
reallyexcellentresearchonfrogtrematodebiologywhileworkingtowardaPhDdegree.This
wasfollowedbystintsattheUniversityofWisconsin–Oshkosh(Oshkosh,Wisconsin)andasa
programdirectorinsystematicsattheNationalScienceFoundation.Heispresentlyafaculty
memberandadministratorattheUniversityofNebraska–Omaha(Omaha,Nebraska).

AnnaSchotthoefer
AnnareceivedherBSdegreeinbiologyatCornellUniversity(Ithaca,NewYork)in1993and
thenspentacoupleofyearsattheVirginiaInstituteofMarineScience(Gloucester,Virginia)
withDr.EugeneBurreson.ShecametoWakeForestin1996andcompletedhermaster’s
researchin1998.Herworkwasfocusedonthecommunitystructureofdigenetictrematodesin
thepulmonatesnailsHel.ancepsandP.gyrina.Shewasoneofthekeyplayersindeveloping
ideasregardingdifferentialtransmissionfociforparasitespeciesinCharlie’sPond.She
completedherPhDwithDr.RebeccaColeandispresentlyahousewifeinAurora,Colorado.

KelliSapp
KellispentasummeratWakeForestasanNSFREUfellowbeforeacquiringher
undergraduatedegreeatMethodistCollege(Fayetteville,NorthCarolina).Shesuccessfully
completedherMSdegreeinmylabbeforeheadingofftotheUniversityofNewMexico
(Albuquerque,NewMexico)andpursuingherPhDwithDr.SamLoker.AsisnotedinChapter
7,sheismarriedtoanotherofmyformerstudents,Dr.HermanEure.Presently,Kelliisavery


successfulfacultymemberatHighPointUniversity(HighPoint,NorthCarolina).Sheand
HermanarecurrentlycoprogramofficersfortheAmericanSocietyofParasitologists.

BrianKeas

BriancompletedhisundergraduatedegreeatHopeCollege(Holland,Michigan)wherehe
cameundertheinfluenceofDr.HarveyBlankespoor,anotheroldfriendofmine.Hecompleted
hisMSdegreehereatWakeForestwithaverycomplicatedefforttounderstandthegrowthand
reproductivecharacteristicsofHel.ancepswithandwithoutinfectionbyHal.occidualis.He
thenmovedontoEastLansing,Michigan,wherehecompletedhisPhDatMichiganState
University.HeispresentlyafacultymemberatOhioNorthernUniversity(Ada,Ohio).

GiselleBroacha-Bruzinski
GiselleobtainedherundergraduatedegreeatBelmontAbbeyCollege(Belmont,North
Carolina).ShesuccessfullyworkedforherMSdegreebyexaminingtrematodepopulation
biologyinmosquitofishinCharlie’sPond.Shewasmarriedjustaftersecuringhermaster’s
degreeandisnowahousewifeinPennsylvania.

JoelFellis
JoelbeganhisundergraduateworkatLewisandClarkCollege(Portland,Oregon)andthen
completeditattheUniversityofNewMexicoandcameimmediatelytoWakeForest.His
researchforhisMSdegreewasfocusedonthecommunitystructureandseasonaldynamicsof
helminthparasitesingreenandbluegillsunfishinCharlie’sPond.Interestingly,thesuitesof
parasitesinthetwofishhostswerequitesimilar,butwhentheirabundancepatternswere
examined,hefoundthemtobedistinctlydifferent.Iwasquitepleasedthathestayedinmylab
todohisdissertationresearch,whichinvolvedtheeffectsofthelifecyclevariationon
communityassemblyandpopulationgeneticstructureofmacroparasitesinbluegillsunfish
fromaseriesofpondssituatedatvaryingdistancesfromeachotherinForsythCounty,North
Carolina.JoeldecidedtopursueanonacademiccareerafterfinishinghisPhDdegreeandhas
becomeaverysuccessfulbusinessmaninCalifornia.

Michael(Mike)Barger
Mikeisa“flatland”guywhoeventuallyendedupintheAppalachianMountainsworkingon
hisPhDHisundergraduateworkandMSdegreeswereobtainedattheUniversityof
Nebraska–Lincoln,whereDrs.JohnJanovyJr.andBrentNickolfirstmentoredhim.Lotic

habitatsincluderiversandstream,whilelentichabitatsarecomprisedofstandingwatersuch
aspondsandlakes.Mostoftheaquaticworkinvolvingparasiteshasbeenaccomplishedin
lenticsettings,soMikedecidedthathewouldbecomeinvolvedwithstreamsystemsupinthe
AppalachianMountainsintheheadwatersoftheYadkinRiver.Acoupleofunexpected
opportunitiesprecededhisstreamecologyresearch,however,whenhefirstencountered
Allopodocotylechiliticorumn.sp.,whichhedescribed.Tocompoundhisearlyventureinto
thestreams,hethenfoundPlagioporussinitsini,averyunusualtrematodeparasitewitha1-,


2-,or3-hostlifecycle,allatonce!Heeventuallymovedintolandscapecommunityecologyin
fishfromtheheadwaterstreamsoftheYadkinRiver,amostproductiveexperience.On
completionofhisdissertationresearch,hefounda“perfect”academicpositionatPeruState
College(Peru,Nebraska)backhomeinhisflatlandsurroundingsofsoutheasternNebraska.

LaurenCamp
LaurenisthedaughterofJoeCamp,aformerPhDstudentofmine.Sheobtainedher
undergraduatedegreeattheUniversityofChicago(Chicago,Illinois)andcametoWakeForest
toacquireherMSdegreeinmylab.Laurenwasthefirstofmygraduatestudentstodoany
researchwiththerhabditidnematodeDaubayliapotomacainMallardLake,asmallpond
locatedonagolfcoursenotfarfromWinston-Salem.Herworkwasverythoroughandpaved
thewayforMikeZimmermannandKyleLuthtosubsequentlyfocusontheparasite.On
completionofhermaster’sdegree,shemovedtotheUniversityofCalifornia–Davis(Davis,
California)whereshehasworkedtowardherPhDunderthedirectionofDr.SteveNadler.

Nicholas(Nick)Negovetich
NickcametoWakeForestfromWabashCollege(Crawfordsville,Indiana)andEricWetzel
(mentionedearlier)in2001.HisMSandPhDdegreesbothinvolvedpopulationand
communityecologyofdigenetictrematodesinCharlie’sPond.Usingmark–release–recapture
methodsforHel.anceps,hewassuccessfulinmathematicallymodelingthesnailpopulation
biologyusingmatrixmodels.AfterreceivinghisPhDin2007,hedidapostdoctoralstintover

a4-yearperiodatSt.JudeChildren’sResearchHospitalinMemphis,Tennessee,wherehe
workedwithDr.RobertWebster,thenotedvirologist/epidemiologist.Nickrecentlyaccepted
anacademicpositionatAngeloStateUniversity(SanAngelo,Texas).

KyleLuthandMichael(Mike)Zimmermann
WhenIbeganwritingthemicrobiographiesofmygraduatestudents,IdecidedtoputKyleand
Miketogether.MikeisfinishednowandpresentlyteachesatShenandoahUniversity
(Winchester,Virginia).Kylewillbemylastgraduatestudent;heisanalyzinghisdataandwill
bedoneinanotheryearorso.Together,theyarrivedhereinthefallof2008andbegantheir
master’sworkonarhabditidnematode,D.potomaca,althoughKylehadtopulloutofhis
initialfocusbecausehisresearchwasnotproducingwhatweexpected.Theybothfinished
theirMSdegreesinthespringof2010.
KyleLuth’sfirstmentorwasnoneotherthanEricWetzelwhodidhisPhDdegreewithmehere
atWakeForest(seeChapter7)andthensecuredapositionatWabashCollege.Bothofthese
guysandIweresimilartoalotofyoungsterswhobeginourundergraduateworkthinkingabout
apremedprogramandmedicalschool.Atsomepointduringtheirundergraduatecareer,
however,theycameundertheinfluenceofaprofessor,orafascinatingaspectofbiology,
whichthenstimulatedateachingandresearchcareer.
MikegraduatedfromMuskingumCollege(NewConcord,Ohio)in2008.Inadditiontoplaying
baseball(hewasapitcher)andfootball(Americanstyle—hewasapunter),hebecame


interestedinparasitologyasanundergraduatestudentwhiledoinganhonorsprojectonfish
parasitesinseveralabandoned“sandpits”eastofNewConcord.Hehadreadsomefish
parasitologypaperscoauthoredbymygraduatestudentsandmewhileworkingonhis
undergraduatedegreeatMuskingum,andhisinterestintheseparasiteswasstimulated.
Iwantedtoendthebiographicsketchesonapositivenote,andIthinkthereisnothingbetter
thantobrieflydescribethecontentoftheirPhDdissertations.Whentheywerecompletingtheir
master’sdegrees,Iapproachedeachofthemwithadifferentresearchproposal,which
capturedtheirattentionandbothdecidedtostay,representingthelasttwoofmygraduate

students.Sincetheyrepresenttheendofmycareer,Ithoughtitwouldbeofinteresttodescribe
theirworktogetherbecauseIthinkitisamongthebestmystudentswouldhaveaccomplished.
Kyle’sfocuswastobeonthegeographicdifferencesinbiologyofautogenicP.ambloplitis,
theso-calledbasstapeworm.Thebasisforhisstudyisrelatedtothefactthatadultsofthe
cestodeoccurinthesummerinthenorth(Eschetal.,1975)andinthewinterinthesouth
(Eure,1974).TheoriginalobjectwastocollectP.ambloplitisinthenorthandsouthandthen
performPCRandsequencingtoseeifpopulationsaregeneticallydifferentgeographically.In
otherwords,arewedealingwithcrypticspecies,orisitsomesortofbehavioral,
physiological,orecologicalphenomenonthatcanaccountforthelifecyclepatterninthenorth
andsouth?
Mike’sstudywastoconcentrateonthegeneticdifferencesoftheallogenictrematode
EchinostomarevolutuminthethreeflywaysusedbyCanadageeseeastoftheRocky
Mountains.Howmuchgeneticmixingandvariabilityoccurintheseparasites?InMike’scase,
thiswouldnotrequireshootingofthegeeseinthethreeflywaystocollectadultsofE.
revolutum.Itwassufficienttocollectsnailsanduseechinostomemetacercariaeforthe
conductofPCR,sequencing,andmolecularcomparisons.Inbothcases,thisisthewaytheir
workstarted.
However,asthingsprogressedearlyduringtheirresearch,Kyledevelopedtheideathatit
mightbeagoodthingifhecouldworkfromeasttowestaswellasnorthandsouth.So,he
approachedmewiththeproposalandIagreed.Mikequicklyjoinedinbecausehehadalready
plannedonworkingeasttowestifhewasgoingtosampleinthethreeflyways.So,whydidn’t
hejoinwiththefieldeffortsproposedbyKyle?Afterall,fourhandsareconsiderablymore
effectivethantwo!Afterahugecollectingeffortinthesummerof2012,theydecidedtoexpand
theircollectingevenfurtherin2013.
Altogether,theywereinthefieldfor70daysovertwosummers.Theycoveredgreaterthan
16,000milesandsampled235sitesin231pondsandlakesin29stateseastoftheRocky
MountainsallthewaytotheAtlanticCoastandsouthoftheCanadianborderdowntotheGulf
Coast.Indoingso,Kylecollected2361centrarchidfish,andMikeexaminedgreaterthan9000
snails,representingfivespecies.IshouldemphasizethatKylehasalsoperformedfull
necropsiesonallofthesefishsothathenotonlyacquiredplerocercoidsofP.ambloplitisbut

entericadultsandparentericlarvalhelminthsaswell.Mikealsocollecteddataforcercariae
shedding.Inotherwords,thetwoofthemassembledtwoofthelargest,ifnotthelargest,sets
offielddataofwhichIamaware.Mikesuccessfullydefendedhisdissertationresearchin


Aprilof2014.Kyledecidedthathewantedtostayonanextrayearsothathecouldcomplete
hisgeneticsworkandthoroughlyconductthedataanalysisforhisdissertation.

Closingcomments
IwassittingwithMikeZimmermannoneafternooninthespringof2014talkingabouthis
research—IhadjustspokenwithKyle30minearlieraboutthesamething.Mikeremindedme
thatinordertosuccessfullyaccomplishsomethinglikethetwoguyshad,theymusthaveagood
workethic,andtheybothdid.Ibeganthinkingaboutthisalittlemoreastheafternoonpassed.
Iagreethatastrongworkethicisessential,butthereismore.Therewasalsoaneedfor
financialbacking,andIhadnograntmoneytosupporttheirwork.Instead,theyusedmoney
fromwhatiscalledtheGradyBrittFund,actuallynamedforthemanIreplacedwhenIcameto
WakeForest.Idonotknowthecircumstancesofhisdepartureonlythathewentontoasmall
collegeintheDeepSouthwherehefinishedhiscareer.Hehadnofamily,andwhenhedied,he
lefthisestatetotheDepartmentofBiologywithinstructionsthatincomegeneratedwastobe
usedbygraduatestudentsinparasitology.WithsupportfromtheGradyBrittFund,Kyleand
Mikehaddollarstosubsidizetheirfieldtripsintothe“boonies”andthentohandletheirPCR
andsequencingwork.
Ialsobelievethesetwostudents,plusalltheothersIhavedirected,hadtohavean“intangible
asset”thatpeoplemustpossesswhopursueeitherorbothofthetwohighestgraduatedegrees.
Theintangibleissomethingdeepinsideaperson.Itisdifficulttodefine,butIknowitisthere.
Personally,IcansenseiteverytimeIseeoneofmystudentsachievesuccess.Iknowitisthere
becauseitisafeelingIhavewhenoneofmystudentssecuresajob,orpublishesapaper,oris
admittedintoanothergraduateprogram,orentersmedicalschool.ThereisaquotationIused
oncebeforeduringmycareer(Esch,1999),andIbelievethatitisareflectionofmypersonal
intangible.ItcomesfromCarolusLinnaeus(1707–1778).Heisquoted,“Aprofessorcannever

betterdistinguishhimselfinhisworkthanbyencouragingacleverpupil,forthetrue
discoverersareamongthem,ascometsareamongthestars.”Whilemypersonalintangibleis
difficulttodefine,IamcertainthatLinnaeus’declarationadequatelyexpressesit.

References
Esch,G.W.1999.Musingsofamentor:AcceptanceoftheClarkP.ReadAward.Journalof
Parasitology85:1008–1010.
Esch,G.W.,W.E.Johnson,andJ.R.Coggins.1975.PopulationbiologyofProteocephalus
ambloplitisinthesmallmouthbass.AcommemorativevolumeoftheOklahomaAcademyof
SciencepublishedinhonorofProfessorJ.TeagueSelf.ProceedingsoftheOklahoma
AcademyofScience55:122–127.
Eure,H.1974.Studiesontheeffectsofthermaleffluentonthepopulationdynamicsofhelminth
parasitesofthelargemouthbass,Micropterussalmoides.Ph.D.dissertation,WakeForest