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Oikos 1997

Oikos 1997

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Published by Dr. Debal Deb
Gut content analysis (GCA) enough to ascertain dietary linkages of species? Unlikely. GCA is not even desirable in the case of rare and endangered species. Instead, direct observations of predation, published literature, and indigenous people's knowledge can be reliable sources of information about food habits of species.
Gut content analysis (GCA) enough to ascertain dietary linkages of species? Unlikely. GCA is not even desirable in the case of rare and endangered species. Instead, direct observations of predation, published literature, and indigenous people's knowledge can be reliable sources of information about food habits of species.

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Published by: Dr. Debal Deb on Sep 28, 2008
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FORUMFORUMFORUM
FORUMisintendedfornewideasornewwaysofinterpretingexistinginformation.Itprovidesachanceforsuggestinghypothesesandforchallengingcurrentthinkingonecologicalissues.Alighterprose,designedtoattractreaders,willbepermitted.Formalresearchreports,albeitshort,willnotbeaccepted,andallcontributionsshouldbeconcisewitharelativelyshortlistofreferences.Asummaryisnotrequired.
Trophicuncertaintyvsparsimonyinfoodwebresearch
DebalDeb,WWF-India,EasternRegion,TataCentre5thfloor,
43
ChowringheeRd,Calcutta700071,India.
Gutcontentanalysis(GCA)isthemostwidelyacceptedmethodforgeneralisingaboutaspecies'foodhabits.GCAisvaluableifthepurposeofthestudyistodeterminethefrequencyorstrengthofinteractionsbetweenspecies,ortoestablishnewfoodlinks.However,determiningallfoodlinksthroughGCAisimpossibleforlargespeciosewebs.Furthermore,GCAmaynotrevealthetruenatureoflinkagedynamicsduetoenvironmentalandphysiologicalstochasticity.Itisthereforeparsimonioustoas-sumethatlinkagesbetweenspeciesrecordedintheliteraturewillbefoundinallfoodwebs,ifthesamepreyandpredatorspeciesoccurinthosesystems.Torevealnewlinkages,freshGCAisdesirable,butimpracticableforlargespeciosewebscontainingmanyrareandendangeredspecies,inwhichcaseitmaybereplacedbyseveralnon-dissectivemethods.High-resolutiondatafortropicalwebscouldbegener-atedthroughobservationsmadebytrainedindigenouspeoples.
Gutcontentanalysis(GCA)isperformedtofindoutwhatananimalhaseaten,andthefindingissubse-quentlygeneralized:whathasbeenfoundinaspeci-men'sstomachwouldrepresentthedietitemsofthespecies.Thiskindofinductivegeneralizationisapow-erfultoolinbiology.However,inferringfromGCAmaynotalwaysrevealthetruenatureofinteractiondynamics(StonerandZimmerman1988).Forexample,thevarietyanddensityofprey,accesstothefooditems,predatorhungerandgustatorypreferencesmayaffecttheinferencesdrawnfromthetypologicalGCA.Notonlyarethetrophiclinksalwaysinastateofflux(Lane1985),butthedirectionalityofthelinksalsovariesaccordingtodevelopmentalstagesoforganisms(Warren1989,Deb1995).Thus,thegutcontentsoftoday'ssamplesarelikelytodifferfromthoseofan-othertime;similarsamplesfromdifferentcommunitiesmayalsoyielddifferentGCAresults,duetotheirdifferentspeciescompositionsandabundances,andalsoperhapsduetodifferentenvironmentalinfluences.ConsideringtheseandotherlimitationsofGCA,manyresearchershaveadoptedseveralalternativemethods,buttheirexplanationsastowhyGCAwasnotconducted(StonerandZimmerman1988,Havens1991,Polis1991,Deb1995)oftenappeartorevealthatpeerreviewofsuchworksdemandedthoseexplana-
OIKOS78:1(1997)
tions.This,combinedwiththefactthatdifferentnon-dissectivemethodshavebeenemployedbyfoodwebresearchersonlyinrecentyears(e.g.Havensetal.1996),seemstoindicatethatGCAhasbeenheldbymanyauthoritiestobethemostreliablemethod.SchoenlyandCohen(1991),forexample,insistthatGCAisindispensableforestablishingfoodlinkages,andoughttobeconductedafreshforeverynewde-scriptionofafoodweb,inordertoascertainthetrophiclinksactuallyexistinginthesystemduringeachperiodofobservation.TrophicindeterminatenessPrecisionofinferencesregardingthestructureofafoodwebcruciallydependsontheresolutionofdata.Todescribeanentirewebrequiresidentifyingalltaxainthesystem,whichappearstooambitioustoaccomplish.Thechecklistofspeciesinamoderatelysizedcommu-nitytendstolengthenwithtimeandeffortspentonidentifyingthem(Cohenetal.1993,Havensetal.1996).Asacorollary,thedietspectrumofanyonespeciesislikelytoincreaseindefinitelywitheffortstodiscoverthem(Polis1991).Onemightcallthis"trophicindeterminateness",whichseemstobecorroboratedbyagrowingbodyofevidence.Polis(1991)derivedhis"species-effortcurve"fromhisstudyofdesertarthropods,andaquaticsystemshavealsoyieldedsimilarresults.Forexample,thewaterflea
Daphnia,
knowntobepurelyherbivorous,werereportedbyGilbertandcoworkers(BurnsandGilbert1986,GilbertandMacIsaac1989)tokillandconsumesmallrotiferssuchas
Keratella
intheprocessoffilteringalgae.Calanoidcopepods,knownaspelagicherbivores,arenowreportedalsotoeatsmallbra-chionidrotifers(WarrenandLawton1987).Untilre-cently,phagotrophicuptakeofbacteriabyphyto-flagellates(Tranviketal.1989)wasalsounknown.Aquaticmicro-organismsawaitintensivestudiestore-vealfurtherdet.ailsoffeedingbehaviour.Terrestrialexamplesincludesuchcommonmammalianherbivores191
 
assheepandreddeerwhooccasionallyconsume
livebirds
-
tomeetmineraldeficiencyinfood(Bazely
1989),andtheArabianwolf
(Canislupusarabs)
intheSarawatmountainrangewhothrivemainlyonfishandbirds,asreportedintheMarch1995issueof
BBCWildlife
reports(p.11).Thus,thenumberofpotentialfooditemsofananimalmaybelargerthanwhatisbelievedtobeitsactualdietspectrum.Bycontrast,apotentialfoodorganismmaynotbeeatenbythepredator,ifthepreyhasanti-predatoradaptations:forexample,
Keratellaslacki,Brachionuscalyciflorus,
and
Polyarthra
spp.mosteffectivelyevadepredationfromthepredatoryrotifer
Asplanchna
(KerfootandSih1987,GilbertandKirk1988).Inspiteofthisexpandedknowledgeoffeedingbiolo-gies,thepresenceorabsenceofthetrophiclinksmaynotalwaysbedetectablebyrandomgutexaminationsduetostochasticenvironmentalandphysiologicalrea-sons.Predatorhunger,forinstance,regulatestheplanktonicclearanceratesofbothusually-resistantandsusceptibleprey(Stemberger1985).Furthermore,afreshGCAmayrevealthataparticularresourcetypeisabsentfromthegutofananimal,butthatmaynotindicatethe"absenceofalink.Rather,itsimplymeansthatthepredatordidnoteatthepreytypeduringtheperiodequaltothegutpassagetime",whenitwasexamined(Havens1991).Suchspatio-tempora1uncer-taintyabouttheconstancyofdietarylinksmaydebili-tatetheveryobjectiveofGCA.Schoener(1989)opinedthatthedecisiontodrawpotentiallinksmustbeeither"vettedbyexperiments...orbyextensivecomparativeobservations,bothhardtocomebyandunlikelytobeobtainedformostfoodwebsineventhefarfuture,howeverdesirable"(p.1586).SchoenlyandCohen(1991),however,notonlydemandsuchexperimentsandrepeatedobservationsbeperformed,butwouldalsoliketoseetheidealstudenttoreportthefluctuationsofpopulationdensities,aswellasseasonalchangesofselectedabioticenvironmen-talparameters.thatmightgivecluestothefluxofdietarylinks.Thus,theysuggestthattrophiclinkagesbe
4educed
fromallempiricaldataineveryparticularcase,insteadof
inducing
themfromafewcases.Thisstandischaracteristicofextremeempiricism,andleavesnoscopeforgeneralization.OnemayalsoarguethatSchoenlyandCohen'srecommendationimpliesthatfirstwedescribeaphenomenon(here,presence/absenceoftrophiclink),andthenseekasuitablecause(e.g._environmentalparameters)byreferencetowhichwe
canexplainthephenomenonunderthatdescription
-
a
procedurethatcorruptsthescientificmethod.
ApplicationofOccam'srazor
Incontrastwiththeirownrecommendation,SchoenlyandCohen(1991)themselvesreliedonthepublished
192dataoffoodhabitsofthespeciescomprisingtheirtime-specific~ebs,constructed"byassumingthatafeedinglinkfrompreyAtopredatorBwaspresentifandonlyifsuchalinkwaspresentinthecumulativewebandspeciesAandBoccurredtogetheratthetimeofobservation".Acumulativewebisawebwhichincorporatesallknowntrophiclinksbetweenpairsofspeciesacrosstime(linksduringsummer,winter,dayandnight,forexample).ThusAisassumedtobelinkedalwaystoB,regardlessofhowoftenAeatsB,orwhetherthislinkisstrongerorweakerthanothersuchlinksinthecommunity.Thisparsimonyofassumptionthatthelinksbetweenpairsof(onto)speciesobservedinthepastwillbeobservedinthefutureandinsimilarsystemsinvolvesapplicationofOccam'srazor,whichisessentialforallrationalenquiry(Lindh1993).Based.essentiallyonthisargument,mostrecentfoodwebstudiesareliterature-dependent,andyetshowimprovedgeneralisationsaboutwebstatistics(e.g.Martinez1991,1992,Havens1992,1993,Deb1995).Theessentialimprovementinthesestudieshasbeenduetofinerresolutionofknowndataratherthanusingnewdietaryinformation.Oneproblemwithcumulativewebsisthatifontoge-neticdietshiftsoccurred,thesimultaneouspresenceofAandBintheirwebswouldnotnecessarilysignifyafeedingrelationbetweenthem(SchoenlyandCohen1991).Thus,ifBiseatenbythejuvenile,butnottheadultofspeciesA,thecalculationoffoodlinksbetweenthemwheneverAandBco-occurwouldsimplyoverestimatethenumberofactuallinksincumulativewebs.Thisoverestimationcouldbeavoidediftherelevantlife-historystagesoforganismsarede-scribedseparatelyasdistinctwebelementsinthecumu-lativewebs(Havens1992,Deb1995).Iwouldliketocallsuchelements
ontospecies
(Deb1995).Thus,speciesAmayberesolvedintoontospeciesX(theyoungofspeciesA)andY(adultA),suchthatB(intheaboveexample)islinkedwithX,butnotwithy.
Estimatingoverestimationlimits
Constructingcumulativewebsimpliesthattheorgan-isms"inthewebunderstudyuse(or,willuse)alltrophicbiologiestheyareknowntouse.Asaresult,thenumberoffoodlinks
(L)
tendstobeoverestimated(Pimmetat.1991).Herethemethodologicalproblemiswhethertheoverestimationof
L
approximatestheto-pologicallymaximumnumberoffeasiblelinks
(Lmax)
amongstallontospecies,inwhichcasethewholeexer-ciseofcumulativewebanalysisisboundtoyieldspuri-ousresults.Lmaxisdefinedas
Lmax
=[S(S
-If-
I
Si(Si
-I)J/2=
I
SiCS
-sJ/2,
OIKOS78:1(1997)
 
wheres;isthenumberofspeciesontheithtrophiclevel,andS=
L
s;.Totestif
L
calculatedfromcumula-tivewebsmightapproximate
LmaX'
IanalysedmyowndataontwofreshwaterpondsfromsouthernBengal,andsimulated11500randomizedwebsoncomputer,usingthesamepoolofontospecies(Deb1995).Alltrophiclinksbetweentheontospecieswereinferredfromtheliterature.Theresult(Fig.1)showsthatthetwoestimatesmoveincreasinglyapartfromeachotherasSincreases,andthelowerlimitofthe
Lmax
rangeisabovetheupperlimitoftherangeof
L.
Thisindicatesthatthemethodofliterature-dependencedoesnotover-estimatelinkagestosuchextentastoexhaustthepossibilityofcountingallpotentiallinks.
Supplementing
GCA
Informationaboutthepresenceorabsenceofpotentiallinksbetweenspeciesmayserveasfundamentalinfor-mationforconstructingfoodwebs,providedthat(a)thestudyconcernsitselfonlywithwebstructure,notwiththefrequencyorstrengthofinteractions(Polis1991),and(b)asimilarassemblageofthesamepreda-torandpreyspecieshavebeenstudiedpreviously.Whenanewspeciesisidentifiedinanecosystem,orwhenthefoodhabitsofaknownspeciesareunknown,GCAseemstobenecessaryforestablishinglinkageswiththeother(onto)speciesinthesystem.However,1000
L5001020405005
Fig.1.Therelationshipoflinkageswithsystemsize
(S)
forII500randomizedanalogsofpondwebs,createdonrulesofnon-randomlinkagesinferredfrompublisheddata.Eachdotisco-incidentforatleast20datapoints.Thecurvesdepictpowerfunctionrelationships:(a)
L=0.12S21O,
and(b)
Lmax
=0.17S2.19
OIKOS78:1(1997)
estimationoflinksthroughGCAwouldbetooarduousandlengthyforlargefoodwebs.GCAfortherarerspecieswould'beespeciallytimeconsuming.Literaturedependenceisperhapsthebestoptionforthemajorityoftherarertaxawhosefoodlinksareknown(Carney1995).Fornewlinkagestudies,Paine(1988)hasac-knowledgedthevariousmeansofestimatingfoodlinks,otherthanGCA:directobservationofthepredationact,scatology,indirectevidences,immunologicalstudy,and"evenplausibleguesses".AllthesemaysupplementpublishedlinkagedataandevenserveasasubstituteforfreshGCA,especiallywhenGCAcannotbecarriedout.PerformingGCAforamajorityofspeciesintheimmenselyspeciosetropicalfoodwebs,mostofwhichexistintheThirdWorId,seemsimpossibletocarryout;apartfromtheunwieldynumberofspecies,thepaucityofskilledresearchersand/ortherarityand"endan-gered"statusofcertainspeciesthatshouldnotbesacrificedforGCA,and/orthelackofinfrastructureandfundsforfoodwebresearcharethemajorimpedi-ments.Insuchcases,directobservationsofpredationactandindirectevidenceswillbeuseful(e.g.LiebermanandLieberman1987,DinnersteinandWemmer1988).Havensetal.(1996)haveusedamixtureofdirectmeasurementsincludingbirdnestlingregurgitantsam-pling,andinferencesfrompublisheddata.IndigenouspeoplesintheThirdWorIdseemtohaveawealthofempiricalknowledgeaboutthefoodhabitsofalargenumberofspecies.OneeffectivewayoflearningaboutthetropicalsystemsintheThirdWorIdcountrieswouldbetoverifytheindigenouspeople'sexperiencesaboutthefoodhabitsofdifferentspeciesandtheirlifehistory.Ofcourse,onemustbecautiousinrecordingsuchanecdotes,becausemanyofthemarearrantsuperstitions(forexample,apopularbeliefinIndiaisthatsnakesingestbananaandmilk).Ontheotherhand,numerousitemsoffolkknowledge,thougheasilyverifiable,arenoteveninvestigated,becausetheyarepresumedbymanytobescientificallyunusableatthebest,andgrosslyunscientificattheworst.Inthelightofpost-modernistcritique,thismaybeexplainedasalegacyoftheWesternprejudicesagainstindigenousknowledgebaseingeneral.Thus,suchfactsasthewhite-breastedkingfisherpreyingonrats,andthegreateradjutantstorkconsumingcrowanddomesticchickenarehardlyreportedinscientificliterature.Thequestionastowhetheraparticularlinkageistooinfrequenttodeserveestimationinfoodwebstatisticsseemssecondarytotheproblemofhighqualityfoodwebdescription,forwhichidentificationofasmanyfoodlinksaspossibleistheprimaryprerequisite.Thiscouldbefacilitatedbyinvolvinganarmyoflocalindigenouspeopletrainedas"parataxonomists"(NRC1992)forinventoryinglocalbiodiversityaswellasforfoodlinkages.Thefolkknowledge,filteredthroughtrainedobservation,islikelytosupplysufficientlyde-193

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