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text:
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text:
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text:
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text:
he loss of t hese species and the overall loss of sea urchinpr edators e g s ea ott ersin n orthe rnC ali for niaandO regon hasin cr easedth evulner abil it y of kelp forests wo rldwide Krumhansl et al 2016 S D Ling et al 2009 R asher et al 2020 When suffici entl y pertu rbed kelp for ests exhibi t abrupt non li near movement Alan Hasti ngs Wysham 2 010 t o an alt ern ati ve st able state known as ur c hin barrens whe re sea urchinoverg razingli mi tskelprecov ery Fil bee De xter S cheibl ing 2014 S D Lingetal 2015 S cott D L ingetal 2019 Whil etheprecisem echa nism scontroll ingtheseshiftsvarylatit udinally generall y th ey are broug ht about when se a ur chin s exhibi t a s witch in b eha vior going from being tucked aw ay in cra cks an d crevi ces and persist ing on drift alg a e to acti vel y wande ring gra zing and consum ing li ve kelp Harrold R eed 1985 Vanderklift Kendrick 2005 Vanderklift Wernbe rg 2008 This s witch in beh avior

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text:
oft en manifests following loc a li zed dist urbanc e e g following a large w ave e vent that removes both drift algae and li ve kelp Ebeli ng et al 1985 Howeve rit isoftenun cle arwh etherth eur chin isamul ti yearperiodoftransientdynami csfollowedbykelprecovery Fuka mi Nakaji ma 2011 Alan Hasti ngs et al 201 8 Hill ebrand et al 2020 The key to diff er enti ati ng these two depends upon whether th e strengt h of fe edback proc esses stabili zing the kelp for est state are sufficientl y degraded by th e pe rturba ti on If so and in particul ar if thi s occu rs across an enti re coastl ine e g see Fi lbe e D exter et al 2 016 differ ent stabili zing feedba cks may incr ease in strength to provide resil iencetoth eurchin b arren state Forexampl e increas edsea urching raz ingreduces kelpcov er reducing spore producti on and incre ases the cov er of en crust ing alga e tha t facil it ates addit ional sett lementofse aur chinlarvae T aniguchiet al 1 994 Thedif fer enc ebet ween alt ernati ve

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text:
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text:
ng he alt hy kelp fo rests a re stru ctu red by di ffe rent regulatory p rocess es at different spati otemporal scales Cavan augh et al 2013 and pulse perturbati ons c an alt er t he pred omi nant spati al s cale of regulation Ed wa rds 2004 R eed et al 2011 Giant kelp Mac rocysti s pyrif era rele as e massive quanti ti es of spores that produce gametophytesdispersed ontheorde rof te nsofm e terstoafew kil ometers S antelices 1990 S chiel Foster 2006 and thu s meta population theory predicts synchronous po pulation dynami cs at thespati alsc aleof disper sal Fit ness F ewe r 19 8 6 A Hasti ngs H arriso n 1994 Inste ad even conti guous stretches of r eef exhibi t kelp subpopul ati ons that fluctuate asynchronously akin to patches withi n a broad e r network Cav anaugh e t al 2014 Whil e spor e dispersal is key for enabli ngrecoloni z ati on particularlyfordist anc edandsmallerpatches loc ali zedexti ncti onisoften a stronger determi nant of kelp dynami cs Castorani et al 2015 This is because local heterogen eit y in

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text:
the form of variati on in substrate and comm unit y compos it ion drive local processessuchasint ra a ndint er specificcomp eti ti onforspacetogrowalo ngthebenthos aswell as herbivory and pred atory consum er r esourc e int eracti ons T he str ength and resil ience of these processes e nco mpass ma ny of the stabi li zing feedbacks that provide resil ience to the kelp forest stat e henc e the need to captur e lo cal high res olut ion metri cs of spe cies habit at associati ons across a broad spat ial extent Howeve r the local scale of regulatory cont rol i s subj ect to forcing fro m larger s cale processessuc hasstorms discussedbelow andthepelagicdispers aloflarv ae Marin epropagule s frominvertebr atessucha sseaurchinsandfishesc antravelfo rtenstohundr edsofkil ometerswith coastal curr e nts Baetsch er et al 2019 da Anadón et al 2011 Kinlan Gaines 2003 Their dispersal proc esses ope r ate at a much b road er sp ati al scale r elative to kel p spore dispersal thus invertebrat e and fish connecti vit y can synchroniz e populations a long a coastl ine a phenomenon Gouhie r et al 2010

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text:
R anta et al 1997 This dynami c synchrony across spa ce can o ccur when dispersal enabled s ett lement overwh elm s local hete rogen eit y That is whenlocalr atesofrep lenishm entared ecoupled fromthelocalregulatory processesstru cturing adult indi viduals Karata yev B askett 2020 For exampl e urchin b arr e ns can persist despit e high adult urchin m ortality from densit y depende nt di sease outbreaks Be hrens Lafferty 2004 because urchin sett lemen t and rec ruitm ent are d ec oupled from both local l arval output and f rom the top down disease mortalit y affecti ng adult s In essence disp ersal m edia ted stabili zing ur chin 8 barren f eedba cks supers ede the densit y depende nt urchin mortali ty locall y structuring the adult population enabli ng persist ence of the urchin ba rr en state In contr ast widespread dispers al may not eli cit spati al synchrony if local scale fe edbac k pro cesses ar e sufficient ly resil ient e g if the local communi ty exerts h igh pos t sett lement m ortali ty upon urchins S tochasti c

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