The Impact of Environmental Variability on Ecological Systems THE PETER YODZIS FUNDAMENTAL ECOLOGY SERIES VOLUME2 Series Editor K.S.McCANN The Impact of Environmental Variability on Ecological Systems Editedby D. A. VASSEUR UniversityofCalgary, AB, Canada K. S. McCANN UniversityofGuelph,ON, Canada AC.I.P.CataloguerecordforthisbookisavailablefromtheLibraryofCongress. ISBN-13 978-1-4020-5850-9(HB) ISBN-13 978-1-4020-5851-6(e-book) PublishedbySpringer, P.O.Box17,3300AADordrecht,TheNetherlands. www.springer.com Printedonacid-freepaper AllRightsReserved (cid:1)2007Springer Nopartofthisworkmaybereproduced,storedinaretrievalsystem,ortransmitted inanyformorbyanymeans,electronic,mechanical,photocopying,microfilming, recordingorotherwise,withoutwrittenpermissionfromthePublisher,withthe exceptionofanymaterialsuppliedspecificallyforthepurposeofbeingenteredand executedonacomputersystem,forexclusiveusebythepurchaserofthework. TABLEOFCONTENTS ListofContributors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi 1 ElNin˜oandLaNin˜a:PhysicalMechanismsandClimateImpacts . . . 1 MichaelJ.Mcphaden 2 HowdoScaleandSamplingResolutionAffectPerceived EcologicalVariabilityandRedness? . . . . . . . . . . . . . . . . . . . 17 JohnM.Halley 3 AssessingtheImpactofEnvironmentalVariabilityonTrophic SystemsusinganAllometricFrequency-resolvedApproach . . . . . . 41 DavidA.Vasseur 4 FilteringEnvironmentalVariability:ActivityOptimization, ThermalRefuges,andtheEnergeticResponsesofEndotherms toTemperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 MurrayM.HumphriesandJamesUmbanhowar 5 EnvironmentForcingPopulations . . . . . . . . . . . . . . . . . . . . 89 EsaRanta,VeijoKaitala,MikeS.FowlerandJanLindstro¨m 6 InteractionAssessmentsinCorrelatedandAutocorrelated Environments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111 Jo¨rgenRipaandAnthonyR.Ives 7 Specialist–GeneralistCompetitioninVariableEnvironments; theConsequencesofCompetitionbetweenResources . . . . . . . . . 133 PeterA.Abrams 8 EnvironmentalVariabilityModulatestheInsuranceEffects ofDiversityinNon-equilibriumCommunities . . . . . . . . . . . . . 159 AndrewGonzalezandOscarDeFeo v vi TableofContents 9 EffectsofEnvironmentalVariabilityonEcologicalCommunities: TestingtheInsuranceHypothesisofBiodiversityinAquatic Microcosms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179 OwenL.Petchey 10 EnvironmentalVariabilityandtheAntarcticMarineEcosystem . . . . 197 ValerieLoeb Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227 ListofContributors PeterA.Abrams MurrayM.Humphries DepartmentofEcologyand DepartmentofNaturalResource EvolutionaryBiology Sciences UniversityofToronto MacdonaldCampus 25HarbordSt. McGillUniversity, Toronto,OntarioM5S3G5,Canada Ste-Anne-de-Bellevue [email protected] QuebecH9X3V9,Canada [email protected] OscarDeFeo DepartmentofMicroelectronic AnthonyR.Ives Engineering DepartmentofZoology UniversityCollegeCork UW-Madison ButlerBuilding Madison,WI53706,USA NorthMall,Cork,Ireland VeijoKaitala MikeS.Fowler IntegrativeEcologyUnit IntegrativeEcologyUnit DepartmentofBiological DepartmentofBiological andEnvironmentalSciences andEnvironmentalSciences P.O.Box65(Viikinkaari1),FIN-00014 P.O.Box65(Viikinkaari1),FIN-00014 UniversityofHelsinki,Finland UniversityofHelsinki,Finland JanLindstro¨m GrahamKerrBuilding AndrewGonzalez DivisionofEnvironmental DepartmentofBiology andEvolutionaryBiology McGillUniversity InstituteofBiomedical 1205aveDocteurPenfield andLifeSciences Montreal,QCH3A1B1,Canada UniversityofGlasgow,Glasgow [email protected] G128QQ,Scotland JohnM.Halley ValerieLoeb DepartmentofEcology MossLandingMarineLaboratories SchoolofBiology 8272MossLandingRoad AristotleUniversityofThessaloniki MossLanding,California U.P.B.119, USA,95039 54124Thessaloniki,Greece [email protected] [email protected] vii viii ListofContributors MichaelJ.McPhaden Jo¨rgenRipa NOAA/PacificMarineEnvironmental DepartmentofTheoreticalEcology Laboratory EcologyBuilding,LundUniversity 7600SandPointWayNE SE-22362Lund,Sweden Seattle,WA [email protected] 98115USA [email protected] JamesUmbanhowar DepartmentofZoology OwenL.Petchey UniversityofGuelph DepartmentofAnimalandPlant GuelphOntario Sciences N1G2W1,Canada UniversityofSheffield Sheffield,S102TN DavidA.Vasseur O.Petchey@Sheffield.ac.uk DepartmentofBiologicalSciences UniversityofCalgary EsaRanta Calgary,Alberta IntegrativeEcologyUnit CanadaT2N1N4 DepartmentofBiologicaland [email protected] EnvironmentalSciences P.O.Box65(Viikinkaari1),FIN-00014 UniversityofHelsinki,Finland esa.ranta@helsinki.fi Preface Thisbook,andtheseriesinwhichitispublished,arosefromaseriesofongoing colloquiaaptlytitled“FundamentalEcology”andheldinhonourofthecontributions of Peter Yodzis to the field of ecology. The impact of environmental variability on ecological systems was a topic selected by Peter for the 2005 colloquium. Peter succumbed to ALS (Amyotrophic Lateral Sclerosis) one month prior to the conference. ix Introduction Fluctuationsinenvironmentalconditionsareubiquitousacrosstheglobe;within terrestrialandaquaticenvironments,polarandequatorialenvironments,andalpine and ocean-deep environments, the physical conditions that influence life are continuously changing in time and space. The action of these fluctuations on an individualofanyspeciescaninducefluctuationsinphysiology,behaviour,phenol- ogy,andmorphology(tonameonlyafew),butitistheirultimateimpactupontraits at the level of populations, communities, and ecosystems which ecologists most often seek to describe. Establishing links between the environment, the proximate, andtheultimateimpactsofchangingconditionswillundoubtedlybetterourability tounderstandandpredicttheoutcomeofnaturalandanthropogenicenvironmental changes. One of the largest challenges for ecologists, when faced with the goal of under- standing the impacts of environmental variability, is to determine which environ- mentalcharacteristicstomeasureormodel.Whilethereareafewperhapsglobally relevantenvironmentalcharacterssuchastemperature,differenthabitatsarelimited tovaryingextentsbydifferentenvironmentalcharacters.Acrossmuchoftheplanet primaryproductionisinfluencedbysolarirradiance,butthereareindeedecosystems existing deep in the oceans and in caves where light does not permeate; primary production in terrestrial systems can be limited by nitrogen and water availability, whileaquaticsystemsprimaryproductionmaybepredominantlyinfluencedbythe availabilityofphosphorus,iron,andother“rare”elements.Heterotrophicorganisms too experience environmental fluctuations which can directly affect their ability to acquire organic and inorganic resources: reductions in stream flow may limit the amount ofsuitablehabitatforacertainfishspecies;waveactionmaysimplywash away the residents of marine intertidal zones; and drought may desiccate even the sturdiestofterrestrialorganisms.Insomecasestheindirecteffectsofchangingenvi- ronmentalcharacteristicsmaybemoreimportantthanthedirecteffectsandinmost casestheirimpactcertainlycannotbeneglected.Whenoneconsidershowanenvi- ronmentally driven change in primary production would impact the herbivores in anecosystembyalteringtheavailabilityoffoodorhowachangeinenvironmental conditions may facilitate or hamper the transmission of disease, it becomes imme- diately clear that a myriad of environmental characteristics may influence a single ecosystem. Fluctuationsinenvironmentalcharactersoccurupontwoaxeswhichareofequal and utmost importance for ecological processes, space, and time. At the regional level, environmental indices such as the North Atlantic Oscillation (NAO) and El xi