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DeterminantsofDistribu/on 1) TheNiche FundamentalNiche:totalrangeofabio/cenvironmentalcondi/onsinwhicha taxoncansurviveandreproduce FromHutchinson(1957)then-dimensionalhypervolume(ormul/dimensional space)thatdescribestherangeofabio/cenvironmentalcondi/onsinwhicha taxoncansurviveandreproduce(eachabio/cfactorisasingledimension) 1 DeterminantsofDistribu/on 1) TheNiche 0 GrowthRate Temperature Temp(C) 0 GrowthRate 10 -10 20 Salinity(ppt) 10 -10 GrowthRate NicheSpace GrowthRa te GrowthRate 10 Salinity 2 DeterminantsofDistribu/on 1) TheNiche Whendorangeboundariescoincidewiththefundamentalniche? EasternPhoebe Northernwintering rangelimit hOp://upload.wikimedia.org/ wikipedia/commons/c/c7/ Eastern_Phoebe1.jpg -4oCJanuaryminimum isotherm (fromRoot1988) Northernrangelimitiscorrelatedwithmetaboliccostsofthermoregula/on foratleast14species(inmul/plesofbasalmetabolicrate:MR=~2.5xBMR) 3 DeterminantsofDistribu/on 1) TheNiche Whendorangeboundariescoincidewiththefundamentalniche? Northernrangelimitiscorrelatedwithmetaboliccostsof thermoregula/onforatleast14species(MR=~2.5xBMR) EasternPhoebe Northernwintering rangelimit (fromRoot1988) hOp://upload.wikimedia.org/ wikipedia/commons/c/c7/ Eastern_Phoebe1.jpg http://ebird.org/plone/ebird/news/patterns-from-ebird-eastern-phoebe 4 DeterminantsofDistribu/on 1) TheNiche RealizedNiche:Asubsetofthefundamentalnichecomprisingtheactual environmentalcondi/onsinwhichataxonsurvivesandreproducesin nature,includingbio/cfactors(compe//on,preda/on,mutualism,etc). Balanus:realizedniche~fundamentalniche Chthamalus:constrainedbycompe//ve interac/onstonarrowerrealizedniche Howcouldwetestthisexperimentally? DeterminantsofDistribu/on 1) TheNiche RealizedNiche:Asubsetofthefundamentalnichecomprisingtheactual environmentalcondi/onsinwhichataxonsurvivesandreproducesin nature,includingbio/cfactors(compe//on,preda/on,mutualism,etc). Anaturalexperiment...withbirds SpeciesAandBarecloselyrelated~similarniche CordilleraVilcabamba MainAndes InthemainAndes: -bothspeciespresent -rangesdonotoverlap Terborgh&Weske1975,Ecology DeterminantsofDistribu/on 1) TheNiche RealizedNiche:Asubsetofthefundamentalnichecomprisingtheactual environmentalcondi/onsinwhichataxonsurvivesandreproducesin nature,includingbio/cfactors(compe//on,preda/on,mutualism,etc). InarangeisolatedfromtheAndes: -higheleva/onspeciesabsent -loweleva/onspeciesexpands rangeupward CordilleraVilcabamba MainAndes CerrosdelSira 100kmfromAndes Ecologicalrelease:expansionoftherealizednicheofaspecieswherefewcompe/torsexist butanundiminishedrangeofresourcesandhabitatsispresent Terborgh&Weske1975,Ecology DeterminantsofDistribu/on 1) TheNiche Importantcaveatstothenicheasthemaindeterminantofaspecies’range: a) Speciesmayoccupyunfavourableareas b) Speciesmaybeabsentinfavourableareas 8 DeterminantsofDistribu/on 1) TheNiche Importantcaveatstothenicheasthemaindeterminantofaspecies’range: a) Speciesmayoccupyunfavourableareas • r=b+i–d–e • popula/onscanbeasource(birthrateexceedsdeathrate;b>d) orsink(deathrateexceedsbirthrate;d>b) • sinkpopula/onsdependonimmigra/on(i)fromsourcepopula/ons • peripheralpopula/onsofaspeciesrangeareogensinkpopula/ons b) Speciesmaybeabsentinfavourableareas • duetogeographicbarriersorisola/on c)Metapopula/onstructurewhensuitablenichespaceispatchy, somepatchesmaybeoccupiedintermiOently 9 DeterminantsofDistribu/on 2) Metapopula?onstructure Metapopula?on:apopula/onconsis/ngofasetofsubpopula/onslinked byacycleofalterna/ngcoloniza/onandex/nc/on(Levins1970) metapopula/on subpopula/ons 10 DeterminantsofDistribu/on 2) Metapopula?onstructure Habitat1(sourcepopula/on) Representa/onofsourceandsinkhabitatsand rela/vemagnitudeofthefourprocessesthat b1>d1 determinegrowthandpersistenceofpopula/ons “Excess”individualsinHabitat1disperse,resul/ng e1>i1 inhigheremigra/onthanimmigra/on Habitat2isabletopersistduetohigher i2>e2 immigra/onthanemigra/on,despitelowerbirths thandeaths b2<d2 Metapopula/onsource/sinkdynamicsmorelikely tooccurtowardtheperipheryofaspecies’range Habitat2(sinkpopula/on) (basedontheabundance-centerhypothesis) Figfromch.4,Lomolinoetal.2006 DeterminantsofDistribu/on 2) Metapopula?onstructure ÅlandIslands,Finland Glanvillefri/llarybuOerfly (Melitaeacinxia) PhotographcourtesyofHannuAarnio. ~4000drymeadowsin2012 Red=occupied Blue=unoccupied Ojanen,S.Petal.2013EcologyandEvolu>on 12 DeterminantsofDistribu/on 2) Metapopula?onstructure 1993-1994 1994-1995 Indexofsynchrony Glanvillefri/llaryontheÅlandIslandsbecamemoresynchronousover/me, possiblyduetoincreasingfrequencyofextremeweatherevents: 2005-2006 Year 2006-2007 Reddown-poin/ngtriangles=decline;Greenup-poin/ngtriangles=increase Sizeofthetriangle~magnitudeofpercapitachange Ojanen,S.Petal.2013EcologyandEvolu>on 13 DeterminantsofDistribu/on 3) Disturbance Habitatdisturbancecancauseabruptrangelimits. Countlessexamples,butoneinourresearchisthehighAndeantreeline: Peruvianfarmersin highlandcommuni/esset firestomaintainfreshgrass sproutsforcaOle Anthropogenictreelineat ~3400m,upperlimitsof forestspecies High-eleva/onspeciesthat moveupslopewithclimate changehit“grassceiling” ManuNa/onalPark,Peru(photosM.Chappell,Z.Peterson) 14 DeterminantsofDistribu/on 3) Disturbance Disturbance-adaptedanimals?Urbanadapters? AmericanCrow (Corvusbrachyrhynchos) CommonStarling (Sturnusvulgaris) 15 DeterminantsofDistribu/on 4) Bio?cInterac?ons:DirectCompe??on Directcompe//on–tes/ngspeciesreplacements Monteverde,TilaránMountains,CostaRica1100-1800m Wood-Wrens (Henicorhina) Wood-Wrens (Henicorhina) White-breasted Wood-Wren Gray-breasted Wood-Wren White-breasted Wood-Wren Gray-breasted Wood-Wren Nightingale-Thrushes (Catharus) Legend Orange-billed Nightingale-Thrush Orange-billed Nightingale-Thrush Black-headed Nightingale-Thrush Black-headed Nightingale-Thrush Slaty-backed Nightingale-Thrush Slaty-backed Nightingale-Thrush 16 DeterminantsofDistribu/on 4) Bio?cInterac?ons:DirectCompe??on Targetspecies:Wood-Wrens Gray-breasted Wood-Wren White-breasted Wood-Wren DeterminantsofDistribu/on 4) Bio?cInterac?ons:DirectCompe??on Targetspecies:Nigh/ngale-Thrushes Slaty-backed Nigh/ngale-Thrush Black-headed Nigh/ngale-Thrush Orange-billed Nigh/ngale-Thrush DeterminantsofDistribu/on 4) Bio?cInterac?ons:DirectCompe??on Ourpreviousexampleofspeciesreplacementsandcompe//onwasa “naturalexperiment”withobserva/onalevidenceofecologicalrelease. Forbirdsthatdefendterritories usingsong... wecanuseterritorydefenseasa behaviouralmetricofcompe//ve interac/ons ...anddesignanexperiment Doesinterspecificcompe//onreinforcerangeboundariesofspeciesalong eleva/onalgradients? Terborgh&Weske1975,Ecology Jankowskietal.2010,Ecology DeterminantsofDistribu/on 4) Bio?cInterac?ons:DirectCompe??on Ourpreviousexampleofspeciesreplacementsandcompe//onwasa “naturalexperiment”withobserva/onalevidenceofecologicalrelease. Forbirdsthatdefendterritories usingsong... wecanuseterritorydefenseasa behaviouralmetricofcompe//ve interac/ons ...anddesignanexperiment Doesinterspecificaggressionreinforcerangeboundariesofspeciesalong eleva/onalgradients? Terborgh&Weske1975,Ecology Jankowskietal.2010,Ecology Behavioralresponsesrecorded: • Closestapproachtospeaker(meters) • Latencytoapproachspeaker(seconds) • Averagelengthofinter-songintervals #@*!? Behavioralresponsesrecorded: • Closestapproachtospeaker(meters) • Latencytoapproachspeaker(seconds) • Averagelengthofinter-songintervals #@*!? DeterminantsofDistribu/on 4) Bio?cInterac?ons:DirectCompe??on Ifspeciesareinterspecificallyaggressiveatcontactzones, thissupportsthehypothesisofdirectcompe//on Playbackprotocoltotestspeciesaggressiveresponses: Trial1 Trial2 Observa/on (Control) Playback (Congener/ Conspecific) Observa/on (Control) Playback (Congener/ Conspecific) 8min.obs 3min.song, 5min.obs 8min.obs 3min.song, 5min.obs DeterminantsofDistribu/on 4) Bio?cInterac?ons:DirectCompe??on ClosestApproach(m) Wood-Wrens 20 15 Gray-breasted W hite-breasted Songplayedtobird: a a Control Congener(otherspecies) 10 b b 5 b b Conspecific(samespecies) 0 Gray-breasted (s/mulus) White-breasted (response) Graybreasted White-breasted Jankowskietal.2010,Ecology DeterminantsofDistribu/on Closest Approach to Speaker (m) ClosestApproach(m) 4) Bio?cInterac?ons:DirectCompe??on Nigh/ngale-Thrushes 20 Slaty-backed Black-headed a a 15 Songplayedtobird: a Control b 10 c Congener(otherspecies) b 5 Conspecific(samespecies) 0 20 15 Black-headed Orange-billed a a b Slatybacked X 10 X Black-headed c b 5 0 CNTR CNGN CNSP b CNTR CNGN CNSP Orange-billed DeterminantsofDistribu/on 4) Bio?cInterac?ons:DiffuseCompe??on Diffusecompe??on:thecombinedeffectofcompe//onwithmanyother species–onespeciesisnega/velyaffectedbynumerousotherspeciesthat collec/velycausesignificantdeple/onofsharedresources(MacArthur1972). Addmorenestboxes(i.e.,cavi/es)morecavitynesterslessopencupnesters Westernbluebird(Sialiamexicana) Americanrobin(Turdusmigratorius) (Bocketal.1992) 26 20 40 60 %NestDepredated 80 100 Nestpreda/on Fortropicalbirds,nestpreda/onisa majorsourceofmortality InManu,Peru,wearegatheringdata tounderstand: 1)hownestpreda/onchangeswith eleva/on? 2)howdoesthenestpredator communitychangewitheleva/on? 0 5) Bio?cInterac?ons:Preda?on DailyNestPreda/onRate DeterminantsofDistribu/on 350$600& 750$1100& 1250$1600& Eleva/on(m) 2500$3000& 27 pearance did not decrease with elevation (linear regression, t = 0.8, P = 0.445, R2 = 0.100). The highest incidence of canary egg disappearance was at 1,650 m. I found no relationship between elevation and incidence of unknown 5) Bio?cInterac?ons:Preda?on marks (linear regression, t = ¡0.4, P = 0.681, R2 = 0.030). Burhans 2 by snakes factors mu lower-elev DiVeren predation Meta-analysisandar/ficialneststudyshowlineardecrease imposed b innestpreda/onratewitheleva/on(CentralAmerica) change alo that at low redate mo nest crypsi elevations dominate, ment in sit This stu standing o tropical fo studies, da 0 500 1000 1500 2000 2500 3000 (Faaborg 2 Skutch'1985,'Boyle'2008' Eleva/on(m) multiple s Fig. 3 Relationship between elevation and proportion of nests depreimpractica Propor/onofnestsdepredated DeterminantsofDistribu/on DeterminantsofDistribu/on 5) Bio?cInterac?ons:Preda?on 40 60 80 100 %NestDepredated DailyNestPreda/onRate Howdoratesofnestpreda/on changewitheleva/onin ManuNa/onalPark,Peru? 0 20 Eleva/on(m) 350$600& 750$1100& 1250$1600& 2500$3000& 29 DeterminantsofDistribu/on 5) Bio?cInterac?ons:Preda?on 40 60 80 100 %NestDepredated DailyNestPreda/onRate Howdoratesofnestpreda/on changewitheleva/onin ManuNa/onalPark,Peru? 0 20 Eleva/on(m) 350$600& 750$1100& 1250$1600& 2500$3000& 30 20 40 60 80 100 Howdoesthenestpredator communitychangewitheleva/on inManuNa/onalPark,Peru? 0 5) Bio?cInterac?ons:Preda?on %NestDepredated DailyNestPreda/onRate DeterminantsofDistribu/on 350$600& 750$1100& 1250$1600& Eleva/on(m) 2500$3000& 31 20 40 60 %NestDepredated 80 100 Catchingtheculprits... 0 5) Bio?cInterac?ons:Preda?on DailyNestPreda/onRate DeterminantsofDistribu/on 350$600& 750$1100& 1250$1600& Eleva/on(m) 2500$3000& 32 DeterminantsofDistribu/on 5) Bio?cInterac?ons:Preda?on Learninglotsaboutthenaturalhistoryof tropicalbirds...includingwhoeatswhom Cock-of-the-Rocknestlingandadult (Rupicolaperuviana) hOp://cdn-7.itsnature.org/wp-content/uploads/2010/04/Cock-of-the-Rock-antbird.jpg DeterminantsofDistribu/on -0.02 1000 1000 1500 2000 2500 1500 2000 2500 Eleva.on&(meters)& Eleva/on(m) elevation 3000 3000 Figure&2.&Rela.onship&between&bird&species&turnover&and& Daily&nest&preda.on&& Changeindailynestpreda/onrates 0.06 0.04 0.00 0.45 0.02 0.55 Species&Turnover& Daily&Preda.on& 0.35 turnover Speciesturnoverishigh=manyspeciesinthecommunityhave rangeboundariesinthesameplace 0.25 Speciesturnover Species&turnover&(Simpson&Index)& 5) Bio?cInterac?ons:Preda?on DeterminantsofDistribu/on -0.02 1000 1000 1500 2000 2500 1500 2000 2500 Eleva.on&(meters)& Eleva/on(m) elevation 3000 3000 Figure&2.&Rela.onship&between&bird&species&turnover&and& Daily&nest&preda.on&& Changeindailynestpreda/onrates 0.06 0.04 0.00 0.45 0.02 0.55 Species&Turnover& Daily&Preda.on& 0.35 turnover Highestchangeinspeciescomposi/on(speciesturnover) correspondswiththehighestchangeinratesofdailynestpreda/on 0.25 Speciesturnover Species&turnover&(Simpson&Index)& 5) Bio?cInterac?ons:Preda?on DeterminantsofDistribu/on 5) Bio?cInterac?ons:Preda?on Browntreesnake(Boigairregularis) SpreadofbrowntreesnakeonGuam (arrows)andsubsequentdeclinesin numbersofna/vebirdsacrossyears Savidge1987Ecology 36 DeterminantsofDistribu/on 5) Bio?cInterac?ons:Preda?on Browntreesnake(Boigairregularis) USGSTimelineoftheBrowntree snaketravelacrossGuam hOp://www.fort.usgs.gov/resources/educa/on/bts/invasion/colonize.asp#dispersal DeterminantsofDistribu/on 6) Bio?cInterac?ons:Mutualism Clark’snutcracker(Nucifragacolumbiana)andwhitebarkpine(Pinusalbicaulis) Whitebark,limber,Colorado pinyon,single-leafpinyon andsouthwesternwhite pinesdependonnutcrackers forseeddispersal Thisinterac/onhaschanged thetrees’seeds,cones,and evenoverallshapein comparisonwithother wind-dispersedpinespecies hOp://www.allaboutbirds.org/guide/clarks_nutcracker/lifehistory 38 DeterminantsofDistribu/on 6) Bio?cInterac?ons:Mutualism Indo-Pacificclownfish(Amphiprionclarkii)andtwoseaanemones (Heterac>smaluandMacrodactyladoreensis) Mutualismsareseldomperfectpredictorsofdistribu/on. 39 DeterminantsofDistribu/on 6) Bio?cInterac?ons:Mutualism Timingofclownfishradia/onwithinterac/ngseaanemonespeciesshownfor eachclownfishspecies 40 DeterminantsofDistribu/on 6) Bio?cInterac?ons:Mutualism LeOersA-Frepresent Indo-Pacificprovinces 26pairsoftropicalclownfish occupy~10speciesofsea anemones Clownfishhaveamucus layertoavoids/ngsfrom anemones Anemonesprovideshelter andprotec/onfrom predatorstoclownfish 41 DeterminantsofDistribu/on 6) Bio?cInterac?ons:Mutualism Radia/onofclownfishhastwogeographicalreplicates.Phylogenyoftheeast Africanclade(GenusAmphiprion)andpiechartscolouredaccordingto mutualis/cseaanemonespecies(withseaanemonespeciesshownbelow) 42 DeterminantsofDistribu/on 7) HistoricalFactors Sapsuckers (Sphyrapicusspp.) ! WoodWarblers (Vermivoraspp.) ! Vireos (Vireospp.) ! ! WoodWarblers (Oporornisspp.) SeveralspeciesofNA birdshavegeographical dividesintherockies WoodWarblers (Setophegaspp.) ! Weir&Schluter2004 ! Sparrows (Passerellaspp.) millionyearsago ! ! ! ! Pacific! Interior! Boreal! Flycatchers (Empidonaxspp.) Chickadees (Poecilespp.) DeterminantsofDistribu/on 7) HistoricalFactors Severalareaswithhigh concentra/onofspecieswith secondarycontactasranges trackedglacialretreat Manyofthese“suturezones” clusteraroundtheRockyMountains ofBC icecover! 1 - 4 ! refguia! secondarycontact! Cannings&Cannings2004 A-C! DeterminantsofDistribu/on mourning winterwren ! Birdspecieswith contactzonesin theBCrockies ! pacificwren townsend’s ! macgillivray’s ! ! myrtle ! ! audubon’s ! hermit ! black-throatedgreen Toewsetal.2011 ! black-fronted goldman’s !