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Aquatic Functional Biodiversity An Ecological and Evolutionary Perspective Edited by Andrea Belgrano Department of Aquatic Resources, Institute of Marine Research, Swedish University of Agricultural Sciences, Lysekil; Swedish Institute for the Marine Environment (SIME), Gothenburg, Sweden Guy Woodward Department of Life Sciences, Imperial College London, Ascot, Berkshire, United Kingdom Ute Jacob Institute for Hydrobiology and Fisheries Science, University of Hamburg, Hamburg, Germany AMSTERDAMlBOSTONlHEIDELBERGlLONDON NEWYORKlOXFORDlPARISlSANDIEGO SANFRANCISCOlSINGAPORElSYDNEYlTOKYO AcademicPressisanimprintofElsevier AcademicPressisanimprintofElsevier 125LondonWall,LondonEC2Y5AS,UK 525BStreet,Suite1800,SanDiego,CA92101-4495,USA 225WymanStreet,Waltham,MA02451,USA TheBoulevard,LangfordLane,Kidlington,OxfordOX51GB,UK Copyright(cid:1)2015ElsevierInc.Allrightsreserved. CoverImage,Lower(cid:1)2010:createdusingthesoftwareFoodweb3D,whichwasprovidedby RichWilliams,J.A.DunneandN.D.Martinez(Williams,R.J.Network3DSoftware.Microsoft Research,Cambridge,UK) Nopartofthispublicationmaybereproducedortransmittedinanyformorbyanymeans, electronicormechanical,includingphotocopying,recording,oranyinformationstorageand retrievalsystem,withoutpermissioninwritingfromthepublisher.Detailsonhowtoseek permission,furtherinformationaboutthePublisher’spermissionspoliciesandourarrangements withorganizationssuchastheCopyrightClearanceCenterandtheCopyrightLicensingAgency, canbefoundatourwebsite:www.elsevier.com/permissions. Thisbookandtheindividualcontributionscontainedinitareprotectedundercopyrightbythe Publisher(otherthanasmaybenotedherein). Notices Knowledgeandbestpracticeinthisfieldareconstantlychanging.Asnewresearchandexperience broadenourunderstanding,changesinresearchmethods,professionalpractices,ormedical treatmentmaybecomenecessary. Practitionersandresearchersmustalwaysrelyontheirownexperienceandknowledgein evaluatingandusinganyinformation,methods,compounds,orexperimentsdescribedherein. Inusingsuchinformationormethodstheyshouldbemindfuloftheirownsafetyandthesafety ofothers,includingpartiesforwhomtheyhaveaprofessionalresponsibility. Tothefullestextentofthelaw,neitherthePublishernortheauthors,contributors,oreditors, assumeanyliabilityforanyinjuryand/ordamagetopersonsorpropertyasamatterof productsliability,negligenceorotherwise,orfromanyuseoroperationofanymethods,products, instructions,orideascontainedinthematerialherein. ISBN:978-0-12-417015-5 BritishLibraryCataloguing-in-PublicationData AcataloguerecordforthisbookisavailablefromtheBritishLibrary LibraryofCongressCataloging-in-PublicationData AcatalogrecordforthisbookisavailablefromtheLibraryofCongress ForinformationonallAcademicPresspublications visitourwebsiteathttp://store.elsevier.com/ Publisher:JaniceAudet SeniorAcquisitionsEditor:KristiA.S.Gomez SeniorEditorialProjectManager:PatGonzalez ProductionProjectManager:Luc´ıaPe´rez Designer:MatthewLimbert TypesetbyTNQBooksandJournals www.tnq.co.in Contributors Mat´ıasArimDepartamentodeEcolog´ıayEvolucio´n,FacultaddeCienciasandCentro Universitario Regional Este (CURE), Universidad de la Repu´blica, Montevideo, Uruguay Andrea Belgrano Department of Aquatic Resources, Institute of Marine Research, Swedish University of Agricultural Sciences, Lysekil, Sweden; Swedish Institute fortheMarineEnvironment(SIME),Go¨teborg,Sweden Mauro Berazategui Departamento de Ecolog´ıa y Evolucio´n, Facultad de Ciencias and Centro Universitario Regional Este (CURE), Universidad de la Repu´blica, Montevideo,Uruguay SofiaBerg EnviroPlanningAB,Go¨teborg,Sweden Iliana Bista Molecular Ecology and Fisheries Genetics Laboratory, School of BiologicalSciences,EnvironmentCentreWales,BangorUniversity,Gwynedd,UK AnaIne´sBorthagarayDepartamentodeEcolog´ıayEvolucio´n,FacultaddeCiencias and Centro Universitario Regional Este (CURE), Universidad de la Repu´blica, Montevideo,Uruguay ThomasBreyAlfredWegenerInstituteforPolarandMarineResearch,Bremerhaven, Germany Joachim Claudet National Centre for Scientific Research, CRIOBE, CNRS-EPHE, Perpignan, France Simon Creer Molecular Ecology and Fisheries Genetics Laboratory, School of BiologicalSciences,EnvironmentCentreWales,BangorUniversity,Gwynedd,UK AnthonyI.DellNationalGreatRiversResearchandEducationCenter,Alton,IL,USA BenoitO.L. DemarsTheJamesHuttonInstitute, Aberdeen,Scotland,UK Anna Eklo¨f Department of Physics, Chemistry and Biology (IFM), Linko¨ping University,Linko¨ping,Sweden Francesco Falciani Institute of Integrative Biology, University of Liverpool, Liverpool, UK JonathanA.D.FisherCentreforFisheriesEcosystemsResearch,FisheriesandMarine InstituteofMemorialUniversityofNewfoundland,St.John’s,NL,Canada Charles W.FowlerBiologyDepartment,Seattle University,Seattle,WA,USA Kenneth T. Frank Department of Fisheries and Oceans, Bedford Institute of Oceanography,Dartmouth,NS,Canada xi xii Contributors David C. Fryxell Department of Ecology and Evolutionary Biology, University of California,SantaCruz,CA,USA Clare Gray School of Biological and Chemical Sciences, Queen Mary University of London, London, UK; Department of Life Sciences, Imperial College London, Ascot,Berkshire,UK Ute Jacob Institute for Hydrobiology and Fisheries Science, University of Hamburg, Hamburg,Germany TomasJonssonPopulationEcologyUnit,InstituteforEcology,Uppsala, Sweden PierreLeenhardt CRIOBE,CNRS-EPHE, Perpignan,France NatalieLowHopkinsMarineStation,Stanford University,PacificGrove,CA,USA Susanna M. Messinger Department of Ecology and Evolutionary Biology, Yale University,NewHaven,CT,USA Fiorenza Micheli Hopkins Marine Station, Stanford University, Pacific Grove, CA,USA Katja Mintenbeck Alfred Wegener Institute for Polar and Marine Research, Bremerhaven,Germany Christian Mo¨llmann Institute for Hydrobiology and Fisheries Science, University of Hamburg,Hamburg,Germany Don T. Monteith Centre for Ecology & Hydrology, Lancaster Environment Centre, Lancaster,UK CamiloMoraDepartment ofGeography,UniversityofHawaii,Honolulu,HI,USA LyneMorissette M-ExpertiseMarine,Sainte-Luce,Canada Eric P. Palkovacs Department of Ecology and Evolutionary Biology, University of California,SantaCruz,CA,USA NicolasPascalEcolePratiquedesHautesEtudes,CRIOBE,CNRS-EPHE,Perpignan, France Samraat Pawar Department of Life Sciences, Imperial College London, Ascot, Berkshire,UK Daniel M. Perkins Department of Life Sciences, Imperial College London, Ascot, Berkshire,UK Owen Petchey Institute of Evolutionary Biology and Environmental Studies, UniversityofZurich,Zurich Vero´nica Pinelli Departamento de Ecolog´ıa y Evolucio´n, Facultad de Ciencias and Centro Universitario Regional Este (CURE), Universidad de la Repu´blica, Montevideo,Uruguay David M. Post Department of Ecology and Evolutionary Biology, Yale University, NewHaven,CT,USA DavidRaffaelli EnvironmentDepartment, UniversityofYork,York,UK AndreaRauJohannHeinrichvonThu¨nenInstituteforBalticSeaFisheries,Rostock, Germany Contributors xiii Luc´ıaRodr´ıguez-TricotDepartamentodeEcolog´ıayEvolucio´n,FacultaddeCiencias and Centro Universitario Regional Este (CURE), Universidad de la Repu´blica, Montevideo,Uruguay Van M. Savage Department of Biomathematics, David Geffen School of Medicine, University of California, Los Angeles, CA, USA; Department of Ecology and EvolutionaryBiology, University of California, Los Angeles, CA, USA; Santa Fe Institute, SantaFe,NM, USA Xiaoliang Sun Department of Molecular Systems Biology, University of Vienna, Vienna,Austria Nash E. Turley Department of Biology, University of Toronto at Mississauga, Mississauga, ON,Canada DavidA.VasseurDepartmentofEcologyandEvolutionaryBiology,YaleUniversity, NewHaven,CT,USA Guy Woodward Department of Life Sciences, Imperial College London, Ascot, Berkshire,UK Perspective: Functional Biodiversity during the Anthropocene Andrea Belgrano, Ute Jacob, Charles Fowler, and Guy Woodward We are living through a new temporal epoch, recently described as the Anthropocene (Latour, 2014), in which human activities are increasingly shapingthebiotawithinandamongEarth’secosystems.These anthropogenic forcesbringavariety ofmajor consequences, includingthesignificant lossof biodiversityinallitsforms,andnotjustthemorefamiliarmeasureofspecies richness (Mora et al., 2011; Cardinale, 2013). Biodiversity, in the broader sense of the term that we use here, includes patterns in the links between species assemblages and their functional organization within a web of in- teractions under environmental constraints. This shift of focus from the traditional emphasis on the “nodes” to the “links” represents an important philosophical change: it forces us to recognize that multispecies systems are not simply passively mapped onto an environmental template, but that their own internal dynamics influence higher level phenomena. Biodiversity is linked to ecosystem functioning and, by extension, to the socioeconomically valuable services they provide (Millennium Ecosystem Assessment (MA), 2005), yet the strength and form of these relationships are still surprisingly poorlyunderstood.Thisemphasizestheurgentneedtodevelopstrategiesthat promotesustainableuseofecosystemsatlocalscales,andofthebiosphereata global scale, in order to preserve a “safe operating space for humanity” (Naeem et al., 2012; Perrings et al., 2010; Dobson, 2009; Mace et al., 2015). If we are to do this, first we need to understand how structure and func- tioning are connected. Functional diversity (FD) is a key component of biodiversity(Perringsetal.,2010;Mouillotetal.,2014)andprovidesadirect link between biodiversity and ecosystem processes (Naeem, 2006), yet com- parisons of FD across systems has largely ignored perspectives that combine ecological and evolutionary principles and understanding (Perrings et al., 2010). It is imperative that a more comprehensive approach is developed for effectively conserving species and their functional roles that considers both ecological and evolutionary principles (Stouffer et al., 2012). xv xvi Perspective:FunctionalBiodiversityduringtheAnthropocene MacroecologicalpatternsintheFDcanbemanifestedacrossawiderange of taxa and systems, as well as across spatial and temporal scales, and orga- nizational levels. This is often achieved by focusing on the explicit links be- tween biodiversity and ecosystem functioning in experimental manipulations, afieldthathasgainedhugetractioninthepastcoupleofdecades(Reissetal., 2009).The(usuallypositive)relationshipbetweenspeciesrichnessandFDhas profoundimplicationsforconservationandmanagement(especiallyofouruse of natural resources) and is also increasingly being linked to food web structure and ecosystem services. Combining macroecological studies of system properties with detailed analyses of community functional diversity patternsandfoodwebstructure(PetcheyandGaston,2007)couldprovideone means of understanding which ecosystem services (Dobson, 2009) are particularlyimportantforsustainingoverallhealthyenvironmentalstatus,and alsowhich might be most vulnerable to perturbations. This is only one of the manychallenges that we need to resolveand understand (Figure 1), if we are EVOLUTIONARY PERSPECTIVE FFuunncc(cid:415)(cid:415)oonnaall DDiivveerrssiittyy FFoooodd WWeebb ssttrruuccttuurree EEccoossyysstteemm SSeerrvviicceess WWhhiicchh eeccoossyysstteemm WWhhaatt aarree tthhee eeffffeeccttss ooff sseerrvviicceess aarree aalltteerreedd bbyy hhuummaann iimmppaaccttss oonn cchhaannggeess iinn ffoooodd wweebb ffuunncc(cid:415)(cid:415)oonnaall ddiivveerrssiittyy ssttrruuccttuurree?? aanndd bbiiooddiivveerrssiittyy lloossss?? HHooww ddoo ffuunncc(cid:415)(cid:415)oonnaall ttrraaiittss rreessppoonndd ttoo cchhaannggeess iinn WWhhiicchh ffuunncc(cid:415)(cid:415)oonnaall WWhhiicchh iimmppaaccttss aarree ffoooodd wweebb ssttrruuccttuurree aatt ttrraaiittss ccaann bbee uusseedd aass mmoosstt iinnflfluueenn(cid:415)(cid:415)aall iinn tthhee ssppeecciieess aanndd iinnddiiccaattoorrss ffoorr ddeetteerrmmiinniinngg cchhaannggeess ccoommmmuunniittyy lleevveell?? eeccoossyysstteemm sseerrvviicceess iinn ffuunncc(cid:415)(cid:415)oonnaall ttrraaiittss?? tthhaatt rreeflfleecctt cchhaannggeess HHooww iiss ffoooodd wweebb iinn ffoooodd wweebb ssttrruuccttuurree bbootthh ssttrruuccttuurree?? ttooppoollooggiiccaallllyy aanndd WWhhiicchh ffuunncc(cid:415)(cid:415)oonnaall ttrraaiittss ddyynnaammiiccaallllyy lliinnkkeedd ttoo WWhhiicchh eeccoossyysstteemm aarree mmoosstt rreessiilliieenntt ttoo eeccoossyysstteemm sseerrvviicceess?? sseerrvviicceess aarree bbootthh aanntthhrrooppooggeenniicc cchhaannggee eesssseenn(cid:415)(cid:415)aall aanndd --iinncclluuddiinngg cclliimmaattee vvuullnneerraabbllee iinn tthhee cchhaannggee?? ccoonntteexxtt ooff tthhee AAnntthhrrooppoocceennee?? FIGURE 1 Challenges and questions at the time of the Anthropocene: linking functional diversity,foodwebstructure,andecosystemservices. Perspective:FunctionalBiodiversityduringtheAnthropocene xvii tomovetowardafunctionallybasedframeworkthatcanincludetheecological and evolutionary roles that species (including humans) play within the local ecosystems on a global scale. Thebiodiversityofecosystemsworldwideisundergoingmajorchange,via human-induced extinctions set against gains via intentional and accidental introductions, with the balance increasingly on the red side of the ledger. These combined effects are altering the structure and function of ecosystems on a global scale, and we are now experiencing what has been widely described as the 6th Great Extinction. An especially challenging task is to understandhowindividualspeciescombinetodeliverecosystemservicesina changing world, yet a rigorous, systematic methodology for doing so still eludes us. This is, at least partially, because the dynamics of ecosystem ser- vices remain poorly characterized at local-to-regional scales, and the role of species in contributing to services is not fully understood. A priori, we know that functional traits, food web structure, and ecosystem services are inter- linked and need to be considered in conjunction with the recognition that ecosystems change andrequire adaptivegovernance toensure their long-term sustainability (Folke et al., 2005; Folke, 2007; Dietz et al., 2003). Recently there has been an explosion of interest in employing network theory to disentangle and explore the relationships between biodiversity and ecosystem functioning (Reiss et al., 2009; Ke´fi et al., 2015). This volume Aquatic Functional Biodiversity: An Ecological and Evolu- tionaryApproachisanattemptbysomeofthemostprominentinvestigatorsin thefieldtoprovideamoregeneralconceptualframeworkthatcanincludenew ecological and/or evolutionary approaches to the understanding of functional diversity. This is accomplished in a way that holds the promise of leading to effectiveandrealisticconservation,especiallyindoingourbestatensuringthe sustainability of ecosystem services in aquatic systems. The collection of chapters in this book represents a substantive contribution to: (1) defining commongroundintermsofterminologyandconceptualissues,(2)connecting conceptualframeworksfromthe ecological and/or evolutionary scienceswith those from classical biodiversity theory, to make progress toward better practical application, and (3) providing examples of how biodiversity and ecosystem services might be conserved more effectively in the real world. TERMINOLOGY AND CONCEPTUAL ISSUES IN ECOLOGICAL AND EVOLUTIONARY PERSPECTIVES Understanding and predicting interactions between ecological and evolu- tionary processes are extremely challenging tasks. Pawar and colleagues (Chapter 1) provide a summary of recent theoretical and empirical advances for developing a mechanistic understanding of trophic interactions, and identifykeymethodsandchallengesforunderstandingandpredictingtheeco- evolutionary dynamics of aquatic ecosystems. These authors introduce a xviii Perspective:FunctionalBiodiversityduringtheAnthropocene theoretical approach to understanding how the metabolic and biomechanical bases of trophic interactions can enhance general predictions regarding the eco-evolutionarydynamicsandfunctioningofaquaticecosystems.Theirideas will be applicable to other types of ecological interactions that involve metabolism and biomechanics (e.g., pollination, parasitism, and competitive interactions). Numerous relationships in which biodiversity contributes posi- tively to ecosystem function have been identified across a wide variety of ecological communities in aquatic and terrestrial ecosystems. Even though these relationships are acknowledged in multiple ways, the underlying mechanisms are poorly understood and are often questioned. In their chapter, Vasseur and Messinger (Chapter 3) introduce and use two models that depict autotrophic and heterotrophic competitors. Their work includes a case study involving a series of experiments using in silico biodiversity and ecosystem function. They show that the “ancestry” of species, which is defined in terms of whether or not they have coevolved in the presence of competitors or in monocultures,isanimportantdeterminantintheextenttowhichtransgressive overyielding can occur. Their research has important implications for the interpretation of previous meta-analyses, and may provide insight of impor- tancetothemanagementofhumaninfluenceonaquaticcommunitiesandtheir recovery from abnormal human influence. Borthagarayandcoworkers(Chapter4)presentthestateoftheartofeffects ofmetacommunitynetworksonlocalcommunitystructures;theirchaptercovers theoretical predictions and empirical evaluations. By using metapopulation models they point outthewide range of patterns predicted by different mech- anisms.Theystressthatrelativeeffectsofdispersalondominantandsubordinate speciesdeterminetheweakeningorstrengtheningofpatchdynamicsimportant tofullyunderstandingcommunitystructure. CONCEPTUAL FRAMEWORKS IN ECOLOGICAL AND EVOLUTIONARY SCIENCES In Chapter 6, Woodward and Perkins focus on the impacts and consequences ofdifferentdriversofchangeonfreshwaterecosystems,whichareparticularly vulnerable to environmental stressors in general, and climate change in particular.Areviewofthecurrentstateoftheserelationshipsispresenteddas a “jigsaw puzzle of our understanding of ecological and evolutionary re- sponses to climate change.” Freshwater ecology has a long history and is arguably much better understood than that of many marine systems: many of the biotic and abiotic constraints (i.e., chemical or physical properties) on species coexistence and food webs are now relatively well established. Although knowledge about how individuals and ecosystem processes are likely to respond to temperature change has improved over the years, the understandingof the community-levelresponses towarming, however, is still in its infancy. Woodward and Perkins conclude that obvious current gaps in Perspective:FunctionalBiodiversityduringtheAnthropocene xix freshwaterecologyareincreasinglybeingfilledbyawiderangeofestablished and novel techniques to enable a better understanding of how freshwater ecosystemsfunction,howtheyarevalued,andhowbesttomanagethemmore sustainably in the future. The monitoring of freshwater ecosystems is crucial forassessingtheiroverallhealthandparticularlyformaintainingthesupplyof ecosystem services. However, the biomonitoring and conservation of fresh waters have failed historically to incorporate a fully ecological and evolu- tionary perspective. Clearly, the predictive capacity and outcomes of current biomonitoring in freshwater ecosystems will therefore be limited in their abilitytoadaptinthefaceofrapidandglobalhabitatmodificationandchange, especiallyasthereferenceconditionsweusedtogaugethestrengthofimpacts shiftawayfromtheirincreasingly“obsolete”baselinesaswemovedeeperinto the Anthropocene. Gray and coworkers (Chapter 10) outline a list of limita- tions in the current state of biomonitoring, and suggest how these problems might be overcome to develop a more holistic ecological and evolutionary approachthatcouldunderpinnewandmoreeffectiveoperationalframeworks. BIODIVERSITY AND ECOSYSTEM SERVICE CONSERVATION Palkovacs and coworkers (Chapter 2) emphasize consumer diversity and the related intra- and interspecific impacts on aquatic communities and ecosys- tems. They provide a synthesis of the state of the art of work focused on the effects of biodiversity among consumer species in regard to the effects of removing species, compared to replacing species (biodiversity loss versus gain). The results of their meta-analysis reveal that, while the effects species have in their influence on communities are stronger on average, the more in- direct elements of intraspecific effects can be important, suggesting that biodiversity within consumer species has important consequences for the ecologyofaquaticsystems.Biodiversitylossposesaglobalthreattoecosystem structureandfunctionandsubsequentlyathreattothegoodsandservicesthey provide (Cardinale et al., 2012); such losses were recently ranked among the dominant drivers of ecosystem change and the loss of ecosystem services (Hooper et al., 2012; Reich et al., 2012). We are now faced with the identifi- cationofthecomponentsofbiodiversity,andthespecificationofwhichspecies in a system are most responsible for providing ecosystem services. Then we need to determine how vulnerable those species are. Jacob and coworkers (Chapter 8) introduce a new approach to the study offunctional diversity and trophic structure within communities in relation to providing ecosystem ser- vices. This approach complements previous topological and dynamical ana- lysesofcommunityrobustnessandmaybeusedtopredicttheresponseoffood webs to the loss of species and resulting degradation of ecosystem services. The chapter by Fisher and coworkers (Chapter 7) focuses on Integrated Ecosystem Assessments (IEAs) and the need to include full consideration of theecologicalandevolutionaryeffectsofselectivefisheries.Theyproposethat

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Aquatic Functional Biodiversity: An Ecological and Evolutionary Perspective provides a general conceptual framework by some of the most prominent investigators in the field for how to link eco-evolutionary approaches with functional diversity to understand and conserve the provisioning of ecosystem
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