A Framework for Community Ecology Species Pools, Filters and Traits This book addresses an important problem in ecology: how are communities assembled from species pools? This pressing question underlies a broad array of practical problems in ecology and environmental science, including restoration of damaged landscapes, management of protected areas and protection of threatened species. This book presents a simple, logical structure for ecological assembly and addresses key areas, including species pools, traits, environmental filtersandfunctionalgroups.Itdemonstratestheuseoftwopredictive models (CATS and Traitspace) and consists of many wide-ranging examples, including plants in deserts, wetlands and forests, and communitiesoffish,amphibians,birds,mammalsandfungi.Globalin scope,thisvolumerangesfromthearidlandsofNorthAfrica,toforests intheHimalayas,totheAmazonianfloodplains.Thereisastrongfocus on applications, particularly the twin challenges of conserving biodiversityandunderstandingcommunityresponsestoclimatechange. PaulA.Keddyisanindependentresearcherwhohastaughtcommunity ecology for more than 30 years. Paul has conducted award-winning research on environmental factors controlling plant communities and theirmanipulationtoenhancenativebiodiversity.Hedelightsinbring- ingsciencealiveforhisaudience.Theauthorofsixbookswinningthree scientific prizes, he also co-edited Ecological Assembly Rules: Perspectives,AdvancesandRetreats(CambridgeUniversityPress1999). Daniel C. Laughlin is a professor of plant ecology and ecological modelling in the Department of Botany at the University of Wyoming. Daniel’s research focuses on developing quantitative approachestounderstandandpredicthowplantspeciesandcommu- nities respond to global change. His lab develops trait-based models thattranslateecologicalprocessesintostatisticalframeworkstopre- dict how communities assemble along environmental gradients and howspeciesinteractatlocalscales. . . A Framework for Community Ecology Species Pools, Filters and Traits paul a. keddy Independentscholar daniel c. laughlin UniversityofWyoming . UniversityPrintingHouse,CambridgeCB28BS,UnitedKingdom OneLibertyPlaza,20thFloor,NewYork,NY10006,USA 477WilliamstownRoad,PortMelbourne,VIC3207,Australia 314–321,3rdFloor,Plot3,SplendorForum,JasolaDistrictCentre, NewDelhi–110025,India 103PenangRoad,#05–06/07,VisioncrestCommercial,Singapore238467 CambridgeUniversityPressispartoftheUniversityofCambridge. ItfurtherstheUniversity’smissionbydisseminatingknowledgeinthepursuitof education,learning,andresearchatthehighestinternationallevelsofexcellence. www.cambridge.org Informationonthistitle:www.cambridge.org/9781316512609 DOI:10.1017/9781009067881 ©PaulA.KeddyandDanielC.Laughlin2022 Thispublicationisincopyright.Subjecttostatutoryexception andtotheprovisionsofrelevantcollectivelicensingagreements, noreproductionofanypartmaytakeplacewithoutthewritten permissionofCambridgeUniversityPress. Firstpublished2022 PrintedintheUnitedKingdombyTJBooksLimited,PadstowCornwall AcataloguerecordforthispublicationisavailablefromtheBritishLibrary. 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Contents Preface pageix 1 AGeneralFrameworkforCommunityEcology 1 Communities 1 AGroundinginReality 3 CompositionandCausalFactorsCanBothBeMapped 8 TheLanguageofSamplesandSampleUnits 9 MoreGeneralPrinciples:AssemblyandResponse 10 TheStudyofPatternIsNottheStudyofCommunity Assembly 13 FoundationsofCommunityEcologyLaidbyRaunkiaer 15 AGeneralFramework 18 TheBasicElementsofCommunityEcology 19 Summary 34 KeyPointsoftheChapter 39 2 Filters 41 ThePowerofFilters 41 DroughtActsasaFilter 43 ColdActsasaFilter 47 HypoxiaActsasaFilter 49 SalinityActsasaFilter 55 HerbivoresActasFilters 57 WildfiresActasFilters 60 PredatorsasFiltersinFrogCommunities 61 PredationasaFilter 63 HowManyFiltersDoWeNeed? 65 MoreonExperimentsandFilters 69 v . vi contents Conclusion:TheUbiquityandPowerofFilters 74 KeyPointsoftheChapter 75 3 SpeciesPools 76 TheListofParts 76 TypesofPools 81 SomeTheoryaboutPoolsandCommunities 88 RatesofArrivalandDepartureintheCommunity 91 SomeEmpiricalRelationshipsObservedinReal Communities 97 ExperimentalManipulationofImmigrationand EstablishmentfromPools 102 InvasiveSpeciesandthePotentialSpeciesPool 104 ApproachestoMeasuringPools 105 MoreonOlderListsandTheirUtilityinDocumentingSpecies Pools 114 LookingAhead 120 KeyPointsoftheChapter 124 4 Traits 125 NamingSpecies 125 ABiologicalParadox:TwoSystemsforOneReality 131 FunctionalClassifications:SomeHistoricalContext 133 MorphologicalTraitsinBirds:BillsAreFundamental 135 PlantFunctionalTraits 139 TheFundamentalGlobalTrait:LeafEconomics 144 GettingtheWatertotheLeaves:TraitsforWater Conduction 147 TraitsandTraitMatrices 149 Traits,EnvironmentsandApplications 163 KeyPointsoftheChapter 164 5 Trait–EnvironmentInteractions 165 ThePowerofFilters 165 . contents vii Overview:QuantifyingTrait–Environment Relationships 167 StaticAbundance-BasedApproaches 169 MoreontheFourthCornerProblem 171 AnExamplefromaWetland 178 DynamicFitness-BasedApproaches 183 Conclusion 186 KeyPointsoftheChapter 189 6 FunctionalGroups 190 FurnitureAssembly 190 FunctionalClassificationforEcologicalPrediction 195 FunctionalGroupsinPlants 197 FunctionalGroupsinBirds 207 FunctionalGroupsinInsects 209 FunctionalTypesofFish 210 FunctionalTypesofMammals 218 FunctionalTypesofFungi 218 SomeMethodologicalIssues 221 OntheRealityofFunctionalGroups 224 TheUnderlyingCausesofFunctionalGroups 226 FunctionalGroups,TraitMatricesandEcological Assembly 228 KeyPointsoftheChapter 230 7 PredictiveModelsofCommunityAssembly 232 PredictioninCommunityEcology 232 CommunityAssemblybyTraitSelection 233 HowManyTraitsAreEnough? 243 TheTraitspaceModelofEnvironmentalFiltering 244 ApplyingtheTraitspaceModeltoConiferForests inSouthwesternNorthAmerica 253 ApplyingBothCATSandTraitspacetoaKettlehole Wetland 256 . viii contents RelaxingtheAssumptionofaSingleOptimal TraitValueUsingTraitspace 257 SomeConcludingRemarksonTwoModelsforCommunity Assembly 260 KeyPointsoftheChapter 263 8 ProspectsandPossibilities 265 LogicalStructureforCommunityEcology 265 Species,FunctionalGroupsandCommunityAssembly 266 TheCanonicalModelforCommonnessandRarity:Lessonsfor CommunityAssembly 283 ResponseRulesandInertiainCommunityAssembly(Time Dependence) 306 TheRoleofCausalFactorsinEcologicalCommunitiesand Restoration 311 KeyPointsoftheChapter 319 References 320 Index 350 . Preface Beforedivingintothefirstchapter,takeamomenttoenjoytheacrylic paintingRamblingRioGrandebyJohnathanHarris,reproducedon thecoverofthisbook.Thisiconicdepictionofthearidsouthwestern USAillustratesmanyofthethemesinthisbook. Let’sstartattheriverandworkourwayupinelevation.The riparianzoneattheriver’sedgeisdominatedbyfast-growingplant species.Eventhoughthesurroundingregionishighdesert,the communityofriparianplantsthrivesinthefluvialsedimentsthat havebeendepositedneartheedgeoftheRioGrande.Thisvegetation consistsofwetsedgemeadowsandcottonwoodforestswith understoriesofCoyoteWillow(Salixexigua),Seepwillow(Baccharis wrightii),andIndigoBush(Amorphafruticosa).Yet,outsidethe floodplain,soilmoistureplummets,andtheseriparianspeciesare replacedbyPinyon–Juniperwoodlands.PinyonPine(Pinusedulis)and One-seedJuniper(Juniperusmonosperma)areshorttreesthattolerate dryconditionsbecauseofthephysiologicalpropertiesoftheirleaves andwood–thatis,their“traits.”Climbingtheelevationgradient further,weenteranarrowbandofmixedforestdominatedby PonderosaPine(Pinusponderosa),DouglasFir(Pseudotsuga menziesii)andQuakingAspen(Populustremuloides).Thistallerand denserforestissupportedbysummerthunderstormsanddeeper snowpack.Beyondthetreelineinthepurplemountains,cushion plantslikeMossCampion(Sileneacaulis)andotheralpinespecies toleratethewind,thebittercoldwinters,andtheshortgrowing seasons.Thus,thispaintingdepictsdistinctcommunitiesthatarise alongoneelevationgradient. Thegoalofthisbookisnotmerelytodescribesuchpatterns,but tolayoutageneralcausalframeworkthatexplainshowtheyarise. Boththecausalframeworkforecologicalcommunities,andthe ix .