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Defining Rarity and Determining the Mechanisms of Rarity for North

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Defining Rarity and Determining the Mechanisms of Rarity for North Definingrarityanddeterminingthemechanismsofrarityfor NorthAmericanfreshwaterfishes JeremyJ.Pritt ThesissubmittedtotheFacultyofVirginiaPolytechnicInstituteandState Universityinpartialfulfillmentoftherequirementsforthedegreeof MasterofScience In FisheriesandWildlifeSciences EmmanuelA.Frimpong,Chair PaulL.Angermeier C.Andrew Dolloff 1/6/2010 Blacksburg,Virginia Keywords:rarespecies,numerical rarity,proportionalrarity,imperiledspecies list,sampling intensity,habitattemplates,speciestraits Copyright2010,JeremyJ.Pritt Definingrarityanddeterminingthemechanismsofrarityfor NorthAmericanfreshwaterfishes JeremyJ.Pritt Abstract Conservingrarespecies andprotectingbiodiversitydepends onsoundinformationonthe natureofrarity.Rarityismultidimensional,presentingtheneedforaquantitativeclassification scheme bywhichtolabelspeciesasrareorcommon.Idefinedrarityforfreshwater fishes based ontherangeextents,habitat breadths,andsiteabundanceandexaminedtherelationship between thesedimensionsofrarityandimperilment.Imperiledfishes weremostoftenrare byall three dimensions,whereasundesignatedspecieswere mostoftencommonbyallthreedimensions. Next,Iexaminedtheeffectofsamplingintensityonobservedrarityofstreamfishusingdifferent numerical andproportionalraritycriteriaandfoundthat increasingsamplingintensityincreased thenumberofspecieslabelledasrarewithproportionalcriteria butdidnotaffectthenumberof species labelledasrarewithnumericalcriteria.Additionalelectrofishingpasseswithinafixed reachincreasesthe likelihoodofdetectingrare andendemicspecies.Atradeoff between informationcollectedandsamplingresources shouldbecarefullyconsideredinthecontextof objectiveswhensamplingforrarespecies.Finally,Iexaminedtheeffect ofregionaland watershedhabitatvariables,biotic interactionvariables,andinstreamhabitatvariables,onthe rareorcommonstatuson23NorthAmericanfreshwaterfishes.Ialsocomparedbiologicaland reproductivetraitsamongspecies classifiedintotherarityframework.Raritywassuccessfully explainedin19ofthe23species andIfoundthat regionalandwatershedhabitatvariableswere themostimportant predictorsofrarity.Ialsofoundthatspecies large bodysize,highfecundity, andlongageatmaturityweregenerallymorecommonbyrangeextentandsite abundancewhile thosespeciesthatdidnotguardnestsweremorefrequentlyrare bysiteabundance. Acknowledgements Thisthesis wouldnot be possiblewithout thehelpandsupportofmanypeople.First,Iwould like tothank myadvisor,EmmanuelFrimpongforhisguidance.Withouthis patience,help,and knowledge,thisthesiswouldnothave beenpossible.Thankstomyadvisorcommittee,Paul AngermeierandAndyDolloff,whohave beeninstrumentalindevelopingtheideasforthis work.IwouldliketothankBrandonPeoplesfor participatinginfrequent ramblingsonall things fishy(scientificandotherwise).Ialsothankthe membersof teamFrimpong:RichPendleton, MollyTainer,AprilGeisler,Victor Wooten,JasonHerrala,JeffersonDeweber,Michael Henebry,andYawAnsah,fortheir assistance inthefield.Finally,special thankstomywife Andreaforherunwaveringsupportoverthe past twoyears. iii Table of Contents Abstract ……………………………………………………………………………………...…ii Acknowledgements ……………………………………………………………………………iii List of Tables …………………………………………………………………………………..vi List of Figures ……………………………………………………………………………...….vii Attribution ……………………………………………………………………………………..ix General Introduction ……………………………………………………………………..……1 Chapter 1 .AquantitativeclassificationofrarityforfreshwaterfishesoftheUnitedStatesand implicationsforimperiledspeciesdesignations……………………………………………….….5 Abstract ………………………………………………………………………………….………..5 Introduction ………………………………………………………………………………………6 Methods ………………………………………………………………………………………….11 Data sources ……………………………………………………………………………..11 Data analysis …………………………………………………………………………….12 Results …………………………………………………………………………………………...15 Discussion ……………………………………………………………………………………….18 Patterns of rarity in freshwater fishes. ………….……………………………………….18 Importance to conservation ……………………………………………………………...19 References ……………………………………………………………………………………….24 Chapter 2 .Theeffectofsamplingintensityontheoccurrenceofrarespecies andcommunity assessmentmetricsinstreamfishsamples………………..……………………………………..38 Abstract …………………………………………………………………………………………38 Introduction ……………………………………………………………………………………..39 Methods ………………………………………………………………………………………….44 Study area ………………………………………………………………………………..44 Sampling methods. ……………………………………………………………………….45 Rarity criteria and assessment metrics …………………………………………………..47 Data analyses …………………………………………………………………………….48 Results …………………………………………………………………………………………...49 Discussion ……………………………………………………………………………………….51 References ………………………………………………………………………………………54 Chapter 3.Macroecological andhabitatfactors determiningrarityinfishesandbiological traits ofrarefishspecies……………………………………………………………………………….66 Abstract ………………………………………………………………………………………….66 iv Introduction ……………………………………………………………………………………..67 Methods ………………………………………………………………………………………….73 Spatial, habitat, and biotic predictions of site-level rarity ………………………………73 Traits as mechanisms for rarity…………………… ……………………………………….75 Results …………………………………………………………………………………………...76 Spatial, habitat, and biotic predictions of site-level rarity ………………………………76 Traits as mechanisms for rarity…………………… ………………………………………78 Discussion ……………………………………………………………………………………….79 References ………………………………………………………………………………………84 Summary and Conclusions …………………………………………………………………100 Additional References………………………………………………………………………102 Appendix ……………………………………………………………………………….……105 v List of Tables Table 1.1. Eight permutationsofthe rarityclassificationframeworkofRabinowitz (1982) brokendownbythreedimensionsof rarity: extent,habitat breadth,andlocal populationsize. Sevenoftheeightgroupsdisplayrarityonatleastonedimensionwhileonegroup(A) is commonacrossall three dimensions………………………………………………..…………..30 Table 1.2. The19habitattypesrepresentingsixbroadcategoriesusedtodetermineindexof habitat breadth(IHB)scoresfor678species intheFishTraits database(Firmpongand Angermeier2009).Fine substrate preferenceindicates anassociationwithmaterialssuchas clay,silt,andsandwhilecoarsesubstrate preferenceindicates anassociationwithgravel,cobble, boulder,or bedrock………………………………………………………………………………30 Table 1.3. Theeightgroupsinthe rarityclassificationschemeandthecriteriacutoffsforeach group………………………………………………………………………………………….….31 Table 1.4. Species that wererarealongall threedimensionsofrarity(groupH) butwerenot designatedasimperiledbyJelksetal.(2008).Stateoccupancyandimperilmentinformationwas obtainedusingthe NatureServeExplorer………………………………………………………..32 Table 2.1. Communityassessment metricsforone-,two,andthree-pass electrofishing. Differentletters indicate significantdifferenceswithBonferroniconfidencelimits.…..………61 Table 2.2. Assessmentoffour NewRiverendemicspecies forone-,two,andthree-pass electrofishing………………………………………………………………………………….…61 Table 3.1. Eight permutationsofthe rarityclassificationframeworkofRabinowitz (1982) brokendownbythreedimensionsof rarity: extent,habitat breadth,andlocal populationsize. Sevenoftheeightgroupsdisplayrarityonatleastonedimensionwhileonegroup(A) is commonacrossall three dimensions……………………………………………………………91 Table 3.2 .Descriptionofregional,watershed,andinstream habitatvariablesusedtomodelthe rarityor commonnessoffishes…………………………………………………………………91 Table 3.3. Modeldetailsofspecies successful predictedas rareor commonbylogisticregression models.…………………………………………………………………………………………92 Table 3.4. Significantexplanatoryvariablesoftherareorcommonstatusof19fishes………..93 vi List of Figures Figure 1.1. Stepsfordistinguishingbetweenrareandcommonspecies basedonsite abundance. Thefirst partofthe processis todefineasmallsiteabundancewhilethesecondis todefinea consistentlysmallsiteabundance………………………………………………….………….33 Figure 1.2. a)Histogramofadjustedspecies rangeareas,b)costfunctionfor breakingadjusted rangeintotwoclasses (wherecost isdefinedas thetotalsumsofsquareddeviations ofspecies valuesfrom theirrespectivegroupmeans),c)histogramofIHBscores,andd)histogramof species at proportionsof sites thattheyoccupyandmeetatleasttwooutofthree localrarity criteria.Verticallinesrepresent thecutoff betweenrare (leftofline)andcommon(rightofline) foreachdimension……………………………………………………………….…………….34 Figure 1.3. a)Quantileregressionfornumerical criteriaandb)relativeabundancecriteriafor sites.Verticallinesrepresent thecriterionforwhichthenumberofspeciesclassifiedasrare is approximately80%oftheasymptotic(stable) rarity…………………………..………………35 Figure 1.4. Numberoftotalspecies andlistedspecies ineachoftheeight finalrarity classificationgroups.Groupsaredefinedas:A) large range extent,widehabitat preferences,and largesiteabundance,B) large extent ,widehabitat,smallsiteabundance,C)large extent,narrow habitat,smallsiteabundance,D)largeextent,narrowhabitat,smallsiteabundance,E) small extent,widehabitat,largesiteabundance,F)smallextent,widehabitat,smallsiteabundance,G) smallextent,narrowhabitat,large siteabundance,andH)smallextent,narrowhabitat,andsmall site abundance…………………………………………………………………………………..36 Figure 1.5. Pairwise bubbleplotmatriciesofspecies bya)Indexofhabitat breadth(IHB)scores andadjustedrangesizes, b)IHBscoresandproportionsofoccurrencesasrare,andc)adjusted rangesizesandproportionsofoccurrencesas rare…………………………………………….37 Figure 2.1 .Fourtheoreticaloutcomesofsamplingintensityonnumericallyrarespecies.Circles indicateacommonspeciesina populationwhiletrianglesrepresenta rare species.Boxa indicates thataspecies israre atlowlevelsofsamplingintensity(areawithintheellipse)and highlevelsofsampling(areawithintheroundedrectangle),andinthis casesamplinghasa neutraleffectonnumericalrarity.Boxbindicatesthatsamplingintensityhasa
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