Collaborative assessment of spiny population and responses to a marine reserve network Author(s): Matthew C. Kay, Hunter S. Lenihan, Carla M. Guenther, Jono R. Wilson, Christopher J. Miller and Samuel W. Shrout Source: Ecological Applications, Vol. 22, No. 1 (January 2012), pp. 322-335 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/41416762 Accessed: 22-09-2015 14:45 UTC

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This content downloaded from 152.3.102.242 on Tue, 22 Sep 2015 14:45:39 UTC All use subject to JSTOR Terms and Conditions EcologicalApplications , 22(1),2012, pp. 322-335 ©2012 by the Ecological Society ofAmerica

lobster Collaborative assessment of California spiny population and to a marine reserve network fishery responses MatthewC. Kay,1,3Hunter S. Lenihan,1Carla M. Guenther,1Jono R. Wilson,1Christopher J. Miller,2 and SamuelW. Shrout2 1 BrenSchool of Environmental Science and Management, University ofCalifornia, Santa Barbara, California 93106-5131 USA 2CaliforniaLobster* and Trap Fishermen's Association, P.O. Box 2294, Capo Beach, California 92624 USA

Abstract.Assessments of the conservationand fisherieseffects of marinereserves typicallyfocus on singlereserves where sampling occurs over narrow spatiotemporal scales. A strategyfor broadening the collection apd interpretationof data is collaborativefisheries research(CFR). Herewe reportresults of a CFR programformed in partto testwhether reservesat theSanta Barbara Channel Islands, USA, influencedlobster size and trapyield, and whetherabundance changes in reservesled to spilloverthat influenced trap yield and effortdistribution near reserve borders. Industry training of scientistsallowed us to sample reserveswith fishery relevant metrics that we comparedwith pre-reserve records, a concurrentport sampling program, fishery effort patterns, the local ecologicalknowledge (LEK) of fishermen,and fishery-independentvisual surveys of lobsterabundance. After six yearsof reserveprotection, there was a four-to eightfoldincrease in trapyield, a 5-10% increasein themean size ( length) of legal sized , and largersize structure of lobsterstrapped inside vs. outsideof threereplicate reserves. Patterns in trapdata were corroboratedby visualscuba surveysthat indicated a four-to sixfoldincrease in lobster densityinside reserves. Population increases within reserves did not lead to increasedtrap yieldsor effortconcentrations (fishing the line)immediately outside reserve borders. The absenceof thesecatch and efforttrends, which are indicativeof spillover,may be due to moderatetotal mortality (Z = 0.59 forlegal sized lobstersoutside reserves), which was estimatedfrom analysis of growthand lengthfrequency data collectedas partof our CFR program.Spillover at theChannel Islands reserves may be occurringbut at levelsthat are insufficientto influence the fishery dynamics that we measured.Future increases in fishing effort(outside reserves) and lobsterbiomass (inside reserves) are likelyand maylead to increasedspillover, and CFR providesan idealplatform for continued assessment of fishery- reserveinteractions. Keywords: California ; collaborative research; ecosystem-based management; effort; fishery dependence; fishing the line; LEK (localecological knowledge); mortality; interruptus;spillover; yield.

Introduction Murawskiet al. 2007,Sethi et al. 2010,Branch et al. thereis that Humansdepend substantially on the proteinand 2011), widespreadperception management mustembrace new for economicrevenues generated by marinefishing. Con- strategies improvedstewardship ofhuman and natural Lubchencoet cernfor the sustainability of fishing industries and the systems(UN 2002, al. FAO ecosystemsupon whichthey depend has increasedin 2003, 2007). recentdecades due to depletionof fishstocks (Pauly et Marinereserves that prohibit consumptive activities al. 2002,Hilborn et al. 2003,Myers and Worm2003), are common globallyand have the potentialto evidenceof resource collapse (Myers et al. 1997,Mullon simultaneouslyprotect ecosystems and fisheries (UNEP- et al. 2005), and perceivedmanagement failure (Pew WCMC 2008). Empiricalstudies indicate that marine 2003). These suboptimalhuman-resource interactions reserves are generallyeffective conservation tools that impactsocioeconomics (Hamilton and Otterstad1998, increasethe abundanceand mean size of organisms Milich1999), as wellas marineecosystems (Pauly et al. withinreserve borders, especially those organisms 1998,Jackson et al. 2001,Lotze et al. 2006).Although targetedby local fisheries(reviews by Côtéet al. 2001, thescale of thefisheries problem is subjectto debate Halpern2003, Lester et al. 2009).However, most studies (Caddyet al. 1998,Walters 2003, Hampton et al. 2005, proceed with considerableand often unaddressed uncertaintydue to lack of replication(at the reserve level),the absence of data to reserve Manuscriptreceived 27 January2011; revised 5 July 2011; prior implementa- accepted4 August 201 1. Corresponding Editor: K. Stokesbury.tion, and thecollection of data oversmall spatial scales 3E-mail: [email protected] (Osenberget al. 2006). Theseshortcomings are under- 322

This content downloaded from 152.3.102.242 on Tue, 22 Sep 2015 14:45:39 UTC All use subject to JSTOR Terms and Conditions January2012 COLLABORATIVELOBSTER RESEARCH 323 standablebecause many reserves are designated as single studythat attributesimproved catches to a nearby areas and/oron politicaltimetables that preclude reserve(Roberts et al. 2001) has been questioned scientificsampling prior to establishment.Regardless, (Hilborn2006). Such uncertaintyis a challengefor manyassessments do not controlfor generallyhigh marinescientists and contributesto stakeholderskepti- spatiotemporalvariability in ecologicalprocesses, and cismregarding the development of reservesas fishery the environmentaldrivers, that contribute to real or managementtools, especially where spillover effects are perceivedreserve effects (Willis et al. 2003,Sale et al. predictedby simplisticmodels that do notaccount for 2005).Reserve studies focused on spinylobster indicate uncertaintyand site-specificfactors (Agardy et al. 2003, thatpopulation increases are commoninside reserves Sale et al. 2005). For marinereserves to reachtheir (MacDiarmidand Breen1993, Edgar and Barrett1997, potentialas conservationand managementtools, there 1999,Kelly et al. 2000, Goñi et al. 2001), but such is a needfor monitoring and assessmentstrategies that increasesare not ubiquitous(MacDiarmid 1991, Mac- fosterstakeholder support and limituncertainty in Diarmidand Breen1993, Acosta 2001, Lipcius et al. measurementsofconservation and fisheryeffects. 2001,Mari et al. 2002,Mayfield et al. 2005),and they One potentialstrategy for limiting uncertainty and providea cautionaryexample regarding the generality of fosteringstakeholder support is the inclusionof for reserveeffects and theneed spatiotemporalcoverage fishermenin reservemonitoring and fisheryresearch. in assessments. Collaborativefisheries research (CFR), in whichfisher- An mechanism whichreserves influence important by men workwith scientists in some or all phases of fisheriesis themovement of adultanimals from within research,is an effectivemeans of increasingthe quality reservesto fishedareas adjacent (spillover).Theory and quantityof data collectedfor managementand that fishedareas to predicts immediatelyadjacent policy assessments(NRC 2004). Additionally,the reserveborders should receive ratesof highest spillover scientificbenefits of CFR are complementedby social of and adults,such that catch rates are juveniles highest benefitsthat often include greater buy-in for manage- nearborders etal. effort (Hilborn 2006).However, high ment and Jentoft1996, and to reserves the (McCay Conway Pomeroy immediatelyadjacent (fishing line)may 2006, and Robertson2009). Due to thesocial localabundance such that catch rates Hartley ultimatelydepress andscientific benefits of CFR, thereare calls nearborders are lowerthan sites farther from reserves widespread to expandthis practice (Pew 2003, U.S. Commissionon (Kellneret al. 2007). Catchrates near reserve borders Policy2004). Collaborativefisheries research is thatare significantlyhigher than catch rates at sitesfar wellsuited to thestudy of interactions between fisheries fromreserves, or thatare lowerbut accompaniedby and marinereserves. Compared with traditional ecolog- higheffort, are ofteninterpreted as reservespillover ical samplingtechniques (e.g., visualscuba surveys), effects.Such patterns have been observed for reef fishes CFR may be superiorfor studying spillover because in thePhilippines (Russ et al. 2004,Abesamis and Russ catchrates at reserveborders are readily to 2005,Abesamis et al. 2006) and Kenya(McClanahan compared effortdistribution, catch therange of and Mangi 2000, Kaunda-Araraand Rose 2004), fishing throughout the historicalcatch records, and other artisanalfisheries in the Mediterranean(Harmelin- fishery, fishery relevantdata sets. CFR can enhance Vivienet al. 2008,Forcada et al. 2009,Stobart et al. Additionally, assessmentof conservationeffects and 2009),spiny lobster in theMediterranean (Goñi et al. population insidevs. outsideof reservesbecause catch 2008),and a trawlfishery in the northeasternUnited changes rates abundance for States(Murawski et al. 2005). Taggingstudies also provide proxies cryptic,nocturnal, or taxa thatare difficultto provideimportant insight into movement across reserve deepwater surveyvisually. The of CFR for both the borders(e.g., Kelly et al. 2002,Kelly and MacDiarmid advantages measuring marinereserves 2003, Goñi et al. 2006), but such studiesare less conservationand fisherieseffects of toolfor assessmentand commondue to highcosts and logisticalconstraints. make it a promising improving Marinereserves benefit conservation through increased stakeholderparticipation. abundanceand/or size of adulttarget organisms inside Herewe report the results of a CFR programdesigned borders,and suchincreases can impactfisheries through in partto testthree research questions concerning the spilloverthat is detectablein catchand effortdynamics influenceof a networkof marine reserves on an actively at reserveborders. fishedmarine , the Potentialfishery benefits of marine reserves are likely ( Panulirusinterruptus). First, we testedwhether over a to be highlyvariable among fisheries (Parrish 1999, relativelyshort period of time(six yearsafter reserve Hilbornet al. 2004) and individualreserves where establishment)reserves influenced trap yield (a proxyfor habitatfeatures are heterogeneouslydistributed and lobsterabundance) and lobstersize structure in reserves influencespillover (Tupper 2007, Goñi et al. 2008, usinga beforevs. aftercomparison. Second, we tested Freemanet al. 2009). As a consequence,the spillover whetherspillover occurred and influenced trap yield and potentialof a givenreserve may be difficultto predict. meanlobster size immediatelyoutside reserve borders. Even whereempirical evidence exists, scientists may Finally,we tested whether effort near interpretdata differentlyand at leastone high-profilereserve borders was higherthan at moredistant sites,

This content downloaded from 152.3.102.242 on Tue, 22 Sep 2015 14:45:39 UTC All use subject to JSTOR Terms and Conditions 324 MATTHEWС. KAY ET AL. EcologicalApplications Vol.22, No. 1 indicatingthat lobster fishermen respond to reserves criteria:(1) historicaltrap yield; (2) historicalpopulation throughfishing the line. size structure;(3) depthand surroundingbathymetry; (4) habitatcharacteristics; and (5) weather Materialsand Methods physical exposureand oceanographicconditions. This informa- Studysites tionwas generatedthrough discussion, scuba surveys, Researchwas conductedat Santa Cruz and Santa and comparisonof pre-reservetrap yield. We selected four sitesfor the reserve Rosa Islands,part of the northernSanta Barbara trapping Scorpion (twoIn, one ChannelIslands (SBCI) located-30 km offshorein Near,one Far), and five(three In, one Near,one Far) forthe Gull and reserves the westernportion of the SouthernCalifornia Bight Carrington (Fig. 1). The is a (Fig. 1). archipelago productivefishing ground Effectsof reserves on trapyield forPanulirus interruptus and manyother and fishesthat inhabit nearshore rocky reefs. The state We testedwhether three reserves (Gull, Scorpion, ofCalifornia designated a network of 10no-take marine Carrington)influenced the spatial patterns of trap yield reservesand twomarine conservation areas in theSBCI in and around reservesby comparingtrap yields in April2003. The reservesencompass 21% of state recordedby fishermenin commercialfishing logbooks waters(high line to 4.8 kmoffshore) surrounding before reserves were established (1998-2002) with trap the SBCI, while the other 79% remainsopen to yieldsthat we generatedin a collaborativetrapping commercialand recreationalfishing (CDFG 2008).We programafter reserves were established (2007 and 2008); sampledat sitesassociated with three marine reserves: A beforevs. after comparison of logbook data alone was Scorpion,Gull (Santa Cruz Island),and Carrington notpossible because commercial fishing is prohibitedin (SantaRosa Island;Fig. 1).A regionalassessment of the reserves.As requiredby law, logbook data record effort effectsof SBCI reserveson lobstercatch, using a before- and catch as the numberof trapspulled and legal aftercontrol-impact paired-series (BACIPS) assessment lobstersretained, respectively, in catch areas defined by of fishery-dependentlandings data reportedfrom the specificgeographic landmarks selected by individual SBCI and nearbymainland, where reserves are notyet fishermen.Detailed calculations of pre-reserve trap yield located,found that total lobster catch and revenueof are providedby Guenther(2010). Briefly,we were fishermenthat fishedwithin the reservenetwork grantedaccess to hard-copylogbooks through collabo- decreasedin the five-yearperiod afterthe reserve rationwith partners in the CaliforniaDepartment of networkwas established (Guenther 2010). That BACIPS Fish and Game (DFG) and the commercialfishery. analysisalso found that total catch and revenue were not Guentherdigitized this data set as a GIS layer, decliningfurther but were increasing in thesixth year conductedfisherman interviews and GIS mappingto afterreserve establishment. Here we examinemore definethe spatial overlap of our research trapping areas localizedresponses in yield, lobster populations, and the and the trappingareas associatedwith fisherman- responseof fishermenaround a subsetof thereserves definedlandmarks in logbooks,and thencalculated withinthe network. the dailyaverage trap yield (i.e., numberof lobsters Individualtrapping sites inside and outsideof each caughtper trap per day) by the fishery in theimmediate reservewere selected in collaboration with five commer- vicinityof ourIn, Near,and Far samplingsites. cialfishermen with a combinedtotal of >60 yearsfishing We measuredtrap yield as wellas thelength of legal- at each site priorto the 2003 reserveestablishment. sized lobsters in the After period from traps placed at In, Collaborationduring site selection and other activities is Near,and Far sitesassociated with each of thethree beneficialbecause fishermen spend more time at sea than reserves(total traps = 15-20replicate traps per In, Near, scientistsand thereforehave enhanced opportunity to andFar site X 3 trappingsites = 45-60traps per reserve). observeand understandthe biologicaland physical Trapswere sampled every two to fourdays at eachsite processesthat influence resource dynamics. This under- duringAugust-October in both2007 and 2008.Across- standingis commonlyreferred to as fisherknowledge or sitecomparisons of research trap yield are based on data local ecologicalknowledge (LEK). Whenincorporated that were collectedprior to the commercialfishing intoecological studies, LEK can enhancehypothesis season,which begins in earlyOctober every year. We formation,sampling efficiency, and the interpretation of constrainedanalysis of trap yielddata to this time results(Hartley and Robertson2009). Accessingthe periodbecause fishery effort can influencecatch rates, LEK of fishermenallowed us to identifyreefs with such that samplingamidst variable effort (i.e., high similarhistorical (i.e., pre-reserve)catch dynamics, effortat Near and Far sitesbut low effortat In sites) physical/biologicalhabitat characteristics, and was mighthave biasedour results.Data collectedduring essentialin guiding selection of individual trapping sites periods when our samplingoverlapped with the locatedinside (referred to in thisreport as In), adjacent commercialfishing season were used in length frequency outside(Near), and -2-6 kmfarther away from (Far) analyses.Traps were deployed haphazardly at 2-20 m reserveborders (Fig. 1). Fishermenworked with waterdepth within areas stratifiedby reefboundaries scientiststo identifytwo to fourreefs inside and outside (i.e., extentof hard bottomsubstrate) that were eachreserve that were similar according to thefollowing delineatedprior to samplingbased on qualitativescuba

This content downloaded from 152.3.102.242 on Tue, 22 Sep 2015 14:45:39 UTC All use subject to JSTOR Terms and Conditions January2012 COLLABORATIVELOBSTER RESEARCH 325

Fig.1 . Mapof sites where collaborative lobster trapping surveys took place (blue circles) at SantaCruz and Santa Rosa Islands inthe Southern California Bight, USA (inset). Also shown are marine reserves (black rectangles) and polygons representing area- specificpre-reserve yields (mean number oflegal sized lobsters/trap) during the period from 1998 to 2002, as calculated fromanalysis of commercial lobster logbooks. surveys,LEK ofcollaborative fishery partners, and the informationabout each sitehad beencommunicated, distributionof giantkelp ( Macrocystispy rif er a). As thebiologist possessed the skills to sampleand re-deploy such,the exact position of each trap on theseafloor was trapsfrom a university-ownedvessel retrofitted with a not controlled,and replicatetraps were separated by commercial-gradetrap hauler. -30 m toavoid nonindependence ofsampling units. The Trapsused in this study were identical to thoseused in distanceof 30 m was identifieda prioriby fishery thefishery for P. interruptusat the SBCI (91.5X 122X partnersas a distancethat would not cause trapsto 45.7cm tall;constructed of Riverdale2X4 inch[5.1 X competeagainst each other,and individuallobstermen 10.2cm] mesh wire; attached at theirbase to a single oftenset theirown trapsmuch closer together. We 91.5X 122cm rectangular frame constructed from 1 inch recordedthe depth, time, date, and GPS coordinatesfor [2.5 cm] diametersteel rod; and coated with a each trapwhen sampling, as wellas thetotal number, hydrocarbonasphalt sealant used to preventcorrosion). sex, carapace'length (to the nearestmillimeter using The onlydifference between research and commercial verniercalipers), injuries (e.g., missing legs or antennae), trapsis thatthe former did not haveescape ports for and reproductivecondition of all lobstersin thetrap. sublegaladult lobsters(~ 70-82.5 mm). Traps were We minimizedstress to lobsterson deck by shading baited with ~500 g of Pacificmackerel ( Scomber themwith wet burlap sacks, placing them in standing japonicus)that was placedin 1-L plasticbait capsules seawater,and returningthem to theocean as quicklyas (oneper trap) after each sampling event. Each trapwas possible.Lobsters were returned to theexact location of connectedto a 3/8inch (0.95 cm) polypropylene line and capture(using GPS coordinates)and releasedby hand. surfacebuoy that allowed for rapid locationand Trappingwas conductedin a two-stageprocess retrievalas in thecommercial fishery. consistingof a trainingperiod conductedaboard The validityof comparing logbook data (1998-2002) commercialvessels followedby trappingfrom a andresearch trapping data (2007-2008) hinges upon two universityboat for much of the remainderof the assumptions:(1) researchtrapping was notbiased (i.e., program.A uniqueaspect of our collaborative program caughtmore or fewerlobsters per trap) relativeto wasthe transfer of LEK fromfishermen to M. Kay,who commercialtrapping, and; (2) pre-reservetrap yield receivedextensive training from a veteranlobsterman across sitesinside/outside reserves did not converge (C. Miller)prior to the project.During this training, upona commonvalue due to uneveneffort across sites. Kay workedas crewduring commercial To clarifythe second assumption, trap yield can be a tripsin and aroundthe study sites. Additionally, other confoundedmeasure of area-specificproductivity in lobstermenon the fishinggrounds provided support fisherieswhere effort is spatiallyheterogeneous and duringthe project, such that the biological sampling was causescatch per unit effort to equilibrateacross space in facilitatedby a collectiveand community-supportedaccord with the equal gainspredictions of ideal free LEK transferfrom the fisheryto the biologist. distribution(e.g., Swain and Wade 2003). To ensurethat Consequently,after traps were initially deployed from our pre-reservetrap yield estimates were reliable for commercialvessels and criticalsafety and fishery analysisand notconfounded by spatially varying effort

This content downloaded from 152.3.102.242 on Tue, 22 Sep 2015 14:45:39 UTC All use subject to JSTOR Terms and Conditions 326 MATTHEWС. KAY ET AL. EcologicalApplications Vol.22, No. 1 trends,we measuredeffort levels in each research Augustto October 2008, we conducted 80 scubatransect trappingarea prior to reserveestablishment using surveysacross 13 of our 14 trappingsites inside and LEK ofour fishery partners. Specifically, we interviewed outside reserves. At each site, we conducted a minimum fishermento determinethe density of trapspresent at ofsix transect surveys on transectsthat were 45 m x 10 each trappingsite for the five-year period immediately m (450m2). Thus, we surveyed >2700 m2of reef at each precedingreserve implementation. Fishermen were sitein Fig. 1. We recordedthe total number of legal- provideda map of trappingareas and askedto report sizedlobsters observed on replicatetransects and then the averagenumber of totaltraps that they recalled calculateda meanfor each location(In, Near,Far) at seeingin each area duringOctober and November each reserve. (timingof commercialseason time most closely corre- The additionof visualsurvey data allowedus to test spondingto oursurveys) from 1998 to 2002.Estimates whetherour trapdata werebiased by unknowntrap withineach area were averaged from all respondents(N performancefactors that might vary across space and = 2-5) andwere used to testfor effort differences across time.Such factors include differential catchability (i.e., sites(In, Near,Far) at eachreserve. theprobability that lobsters at a givensite will enter a Duringthe 2007 and 2008field seasons we conducted trap)and fishingeffort that was lowerduring scientific two activitiesto testthe assumptionthat yields from surveys(After) than during the Before period, when trap researchtrapping and logbookswere unbiasedand yielddata were taken from commercial logbooks. Visual comparable:(1) a comparisonof yields from commercial surveydata also providedan additionaland direct fisherytrapping (estimated from logbook data) and measureof lobsterresponses to SBCI reserves. researchtrapping that took place simultaneously at the ' data to test ScorpionNear and Far sites,and; (2) portsampling. Tagging further forspillover Port samplingconsisted of meetingfishermen at the Althoughwe emphasizespatially explicit trap yield dock and measuringthe size structureof lobsters andeffort patterns to detect spillover of lobsters, we also harvestedfrom relatively large regionsoutside of conducteda companiontag-recapture study to detect reservesthat encompassedour Near and Far sites. movementof lobstersacross reserve borders. During Collectionof these fishery-dependent dataexpanded the trappingevents at In, Near,and Far sites,all lobsters spatialcoverage of our samplingoutside reserves, and weretagged with an individuallynumbered T-bar tag helpedensure that our trapping data were representative (TBA-2 standard;Hallprint Tags, HindmarshValley, of commercialcatches. In total,we portsampled 19 Australia).Tags were applied through a thinmembrane timesfor lobsters caught at SantaCruz Islandand 27 on theventral surface between the tail and carapace, timesfor those caught at SantaRosa Island. suchthat the "T" portionof thetag was anchoredin muscleand persisted Visualscuba lobster throughmolting. Tag-recapture surveysof density studieswere conducted prior to the 2007 and 2008 We comparedour trap yield results inside vs. outside fishingseasons to reducepotential bias caused by of reservesin the beforeand aftertime periods with commercialfishing (October to March)and theunre- lobsterabundance data collectedby NationalPark portedcapture of taggedanimals. Service(NPS) scuba diversin the NPS aroundreserves monitoringprogram. The NPS data were collected Fishingeffort beforeand afterreserve implementation (April 2003) To testwhether fishermen aggregated effort along from1 1 sitesdistributed across Santa Cruz (N= 5 sites), reserveborders (fishing the line) we mapped the SantaRosa (N = 3), and SantaBarbara (N = 3) Islands. distributionof commercialeffort (trap buoys) at Near Threeof thesites were located inside existing reserve and Far trappingsites at each of the threereplicate boundaries,and twosites were located inside the Gull reserves.Effort was mappedfrom a researchvessel by and Scorpionreserves on reefswhere we trapped.The recordingthe GPS coordinatesof individualbuoys on othereight NPS sitesdid notoverlap with our trapping four dates duringthe 2008-2009fishing season: 1 areas.To ensuretemporal consistency with our trapping October(Carrington and Gull), 1 Novemberand 3 data,NPS dataused to comparelobster densities before December2008 (Scorpion,Carrington, Gull), and 19 vs.after reserves were constrained to the1997-2003 and January2009 (Gull and Scorpion). 2007-2010summer field seasons, respectively. NPS Data diverscount lobsters at each siteonce peryear on 12 analysis replicate20 x 3 m transects(12 transectsx 60 m2= 640 The numberof legal sized lobsters(>82.5 mm) m2 sampled at each site) as part of a broader capturedin researchtraps at In, Near,and Far sites community-levelkelp forestmonitoring protocol de- was comparedwith a two-wayANOVA in whichtime scribedby Davis et al. (1997). We estimatedannual (beforevs. afterreserves) and sitelocation (In, Near, meanlobster density for each site from these data. Far)were crossed, fixed factors. Data usedfor the before BecauseNPS data providetemporal coverage but do period(1999-2003) were from logbook analysis, and not align spatiallywith our trappingareas, we also data usedfor the after period were from collaborative reportdata fromour own visualscuba surveys. From researchtrapping (2007-2008). Logbook catch data

This content downloaded from 152.3.102.242 on Tue, 22 Sep 2015 14:45:39 UTC All use subject to JSTOR Terms and Conditions January2012 COLLABORATIVELOBSTER RESEARCH 327 reportthe total number of legallobsters caught from a 50-msegment. Buoy data werepooled from all surveys knownnumber of traps in a givenarea, and from this we (N = 3 surveysat Carringtonand Scorpion,N = 4 calculatedthe average number of legal lobsters per trap. surveysat Gull) at each Near site and separate To standardizelogbook and researchdata, our research regressionswere run for each Nearsite. trappingdata werealso averagedacross all trapsat an Data of lobsterabundance estimated from scuba individualsite for each dailysampling event. We then surveyswere analyzed in two ways.First, a two-way calculateda grandmean of trapyield from all daily ANOVA was used to testwhether lobster abundance fishingand samplingevents at each In, Near,and Far fromNPS scuba surveysvaried as a functionof time sitebefore and afterreserve implementation, such that (beforevs. after;fixed factor), location (In vs. Out of thestandardized unit of replicationin theanalysis was reserves;fixed factor), and theirinteraction. Second, a theaverage trap yield for each of theIn, Near,and Far one-wayANOVA was used to comparelobster abun- sitesat each reserve(total N = threereplicate reserves danceestimated from our own scuba surveys conducted [Gull,Scorpion, Carrington] x threesite locations [In, at trappingsites during 2008 (i.e., in theafter period). Near,Far] x twotime periods [before vs. after]= 18). For all ANOVA analyses,data werelog-transformed The grandmeans for each sitewere pooled from >13 (1п[У4- 1]) to homogenizevariances. Homogeneity of dailysampling events conducted in theafter period of varianceafter transformation was confirmedwith 2007and 2008. Prior to ANOVA,grand mean data were Cochran's test. Only data describingscientific vs. log-transformed(1п[У + 1]) to homogenizevariances. commercialtrap yield in activefishing grounds (used AfterANOVA we comparedmean effects of differentto testassumption that research trapping was not biased treatmentswith Tukey's hsd post hoc tests. Data relativeto commercialtrap yield) failed to meet gatheredto testthe two assumptions that we identifiedstandards for parametric analysis, and in thosecases (comparabilityofcommercial fishing vs. scientific survey we reportresults from Welch's ANOVAs (Zar 1999). trappingand heterogeneouseffort distribution) were Significancelevel in all testswas a = 0.05.Results tables analyzedwith separate one-way ANOVAs. forall ANOVAsare presentedin theAppendix. Mean size of all sized lobstersin was legal traps Results comparedusing a one-wayANOVA in whichsite location(In, Near,Far) was thefixed factor (total N = Trapyield, mean size, visual surveys, threereplicate reserves [Gull, Scorpion, Carrington] X andmovement of lobsters = threesite locations [In, Near,Far] 9). The carapace Analysisof fisherylogbook data fromthe five-year lengthsof all legalsized (>82.5 mm)lobsters trapped at periodprior to reserveimplementation indicated that a givensite were averaged for each samplingday, and catchranged from 0.59 to 0.99 legallobsters/trap at In, fromthese daily means we calculatedgrand means at Near, and Far sites.Trap yieldson Santa Cruz and each site for use in our analysis.Before vs. after Santa Rosa islandsin generalwere spatially heteroge- comparisonswere not possible in thisanalysis because neousand rangedfrom 0.06 to 3.12 legallobsters/trap sizedata are not recorded in logbooks. After ANOVA, a duringthe same period (Fig. 1). Trapyields around the Tukey'shsd post hoc testwas usedto comparemeans threereplicate reserves in theperiods before and after acrosssites (In, Near,Far). reserveestablishment varied with the interaction of time Lengthfrequency data fromeach site location(In, and trappinglocation (two-way ANOVA, time [before Near, Far) were comparedwithin (but not across) vs. after]X location[In, Near, Far], F2, 12 = 15.99,P < individualreserves using Kolmogorov-Smirnov(KS) 0.001; Fig. 2; Appendix:Table Al). A significant tests.Similarly, survey trapping data at theGull and interactionwas generatedbecause trap yield at In sites Scorpionsites (In, Near, Far forboth reserves) were afterreserve establishment (henceforth: In-after) was thencompared with port sampling data fromSanta significantlyhigher than all othertime X location CruzIsland, and survey data from Carrington sites were treatments(Tukey's, P < 0.05; Fig. 2), all of which comparedwith port sampling data fromSanta Rosa were statisticallyindistinguishable from each other Islandwith KS tests. (Tukey's,P > 0.05). Althoughthe mean trap yield at To testwhether reserves influenced fishery dynamics In-aftersites was uniformlyhigher than all other throughfishing the line,the locationand densityof treatments,trap yields at ScorpionIn-after were about commerciallobster trap buoys in all Nearand Far sites one-halfthe yields at Gulland CarringtonIn-after sites. wereexamined using ArcGIS 9 (ESRI 2009).Density of The numberof lobsters per square meter recorded on commercialeffort in the Near vs. Far sites was NationalPark Service (NPS) sciibasurveys varied with comparedwith a one-wayANOVA usingthe commer- theinteraction of timeand location(two-way ANOVA, cial buoydata collectedin 2008-2009.The distributiontime [before vs. after]x sitelocation [In, Out], Fj, ц7 = of trapswithin the Near sites was examinedwith linear 14.13,P < 0.001;Fig. ЗА; Appendix: Table A2a). Mean regression,where distance from reserve border (mea- lobsterdensities at In-aftersites were 4.31-5.60 times suredat the midpointof sequential50-m alongshore higherthan at anyother time X locationtreatments, and segments)was the independentvariable and the thedifferences were statistically significant (Tukey's, P dependentvariable was the numberof trapsin each < 0.05; Fig. ЗА). Mean lobsterdensities measured on

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sitesranged from 89.7 ± 0.60mm (Gull Near) to 96.0± 0.81mm (Carrington Near). Allof the 499 lobsters that were tagged and recaptured acrossIn, Near, and Far sitesat the threereplicate reserveswere recaptured nearest to the reserve where they weretagged. A totalof 310lobsters originally tagged at In siteswere recaptured. Of these, 94.5% were recaptured withinthe In sitewhere they were tagged, while 5.2% and 0.3%exited the reserve and were recaptured in Nearand Far sites,respectively (Fig. 4). Similarly,97% oflobsters (N= 127)tagged at Far siteswere later recaptured at that samesite, while 1.5% were recaptured at boththe Near andIn sites.In contrast,of the 62 lobsterstagged in Near sites,only 70% were recaptured within the same Near site, whereas24% wererecaptured at In sitesand 6% were recapturedat Far sites. Fig.2. Number(mean + SE) oflegal sized (>82.5 mm) The size structureof lobsterpopulations at all three lobsters in at siteswithin immedi- caught replicatetraps (In), reserveshad a greaterproportion of largelobsters In atelyadjacent to (Near), and 2-6 km distant from (Far) three replicateChannel Island marine reserves. Data describingreserves than at Nearor Far sites(Fig. 5; Kolmogorov- conditionsbefore and after reserve implementation arefrom Smirnov[KS], P < 0.05).There was no differenceinsize analysisof commerciallogbooks and collaborative trapping structure between Near and Far sitesfor both Scorpion Letters resultsof surveys,respectively. represent Tukey'spost (KS, P = 0.13)and Gull (KS, />= 0.18)reserves, but size hoctest (a > b at P < 0.05). structurewas significantlydifferent at Carrington Near and Far sites(KS, P < 0.05). Size frequencydata from scubatransects conducted by our researchteam varied significantlybylocation (one-way ANOVA, ^,6= 10.56, P = 0.011;Fig. 3B; Appendix:Table A2b). Mean lobster densitiesat trappingsites In reserveswere 4.23 and 5.38 timeshigher than mean densities at our Near and Far sites,respectively, and thedifferences were significant (Tukey's,P < 0.05). Researchtraps and commercialtraps (reported throughlogbooks) that were deployed in thesame area duringthe 2006-2007 and 2007-2008 fishing seasons did notdiffer in yieldper trap (one-way ANOVAs: 2006- 2007Welch's FXA9 = 0.007,/> = 0.93;2007-2008 Welch's ^1,53= 1.75, P = 0.19; Appendix:Table A3), thus indicatingthat research trapping was not biased relative to commercialtrap yield. With regard to thepotential foreffort heterogeneity toconfound our use of trap yield as a metricfor pre-reserveconditions, fisherman interviewssuggest no statisticallysignificant pre-reserve effortheterogeneity across our surveysites during the 1998-2002fishing seasons (one-way ANOVA, F2¿5= 1.43,P = 0.26; Appendix:Table A4). Thusour use of logbookand researchtrapping data to comparetrap yieldbefore vs. afterreserves is justified. The size of legalsized lobsters caught in trapsafter reserveimplementation varied significantly by location (one-wayANOVA, F2¿ = 8.94,P = 0.016;Appendix: Table A5) and was statisticallygreater at In sites(all threereserves = 100.4± 1.20mm, mean ± SE) thanat theNear or Far sites(Tukey's, P < 0.05),while size at Fig.3. Thedensity of lobsters (mean + SE) observedon Near(92.8 ± 1.85mm) and Far (93.4 ± 0.92mm) sites visualscuba surveys conducted by (A) theNational Park did notdiffer P > Sizesof lobstersIn Service(NPS) kelpforest monitoring program and (B) our (Tukey's, 0.05). researchteam. NPS data include all sizes of lobsters the and Gullreserves were 102.2 observed, Scorpion,Carrington, whiledata from our research team include only legal sized ± 0.67,100.7 ± 0.47,and 98.13 ± 0.36mm, respectively. lobsters. Letters represent results of Tukey's post hoc test (a > Outsidereserves, the size of lobstersat Near and Far b at P < 0.05).

This content downloaded from 152.3.102.242 on Tue, 22 Sep 2015 14:45:39 UTC All use subject to JSTOR Terms and Conditions January2012 COLLABORATIVELOBSTER RESEARCH 329 portsampling (fishery dependent and collectedin the afterperiod) showed similar patterns to data fromtrap surveysat Near and Far sites(Fig. 5). However,port samplingdata from Santa Cruz and SantaRosa Islands had significantlymore large lobsters than our scientific trapsampling when we pooledNear and Far dataat all threereserves (KS, P < 0.001).Such resultswere not surprisingbecause the port sampling data setconsisted of an orderof magnitudemore lobsters than the trap samplingdata. Commercialeffort distribution Wemapped the location of 617 total buoys at all Near and Far siteson fourdates during the 2008-2009 fishing season. Trap densitieswithin these sites were not statisticallydifferent (one-way ANOVA, FU8 = 1.61, Fig.4. Thepercentage oflobsters that were tagged, and P = 0.22; Appendix:Table A6). Trapswere generally subsequentlyrecaptured, ineach of the three survey locations distributed thetotal number and (In,Near, Far). Data for each survey location are pooled from alongshore,although all threereserves. The legendindicates original tagging positionchanged with time, and we did not observea locationsand the number of lobstersrecaptured from each concentrationofcommercial traps near reserve borders tagginglocation (not the total number tagged in each location). (Fig. 6). At Scorpionand Carringtonreserves, traps Dataare from lobsters tagged and recaptured during research were absent trappingsurveys prior to the 2007-2008 and 2008-2009 fishing consistently immediatelyadjacent reserves, seasons. andregression analysis revealed no relationshipbetween distancefrom reserve border (predictor) and the number the effectsand fisheries oftraps (response) within any of our three Near sites (P 1998).Understanding ecological of marinereserves the > 0.05 forall tests).Qualitative comparison of pre- managementpotential against of baselinesfurther underscores the need reserveeffort (fishermen interviews) and post-reservebackdrop sliding to collectrobust data. effort(buoy surveys) indicates that effort at eachsite has spatiotemporal increasesthat we observedacross time notdrastically shifted since reserve implementation (Fig. The trapyield In have been 6, insets). (beforevs. after)at sites reservesmay partiallydue to thefact that effort was lower during our Discussion researchsurveys (after) than during fishing seasons from Thenumber and mean size of legal (>82.5 mm)lobsters whichlogbook data were estimated in the before period. capturedinside reserves were greater than in traps placed However,it is unlikelythat this effortdifference is outsidein fishedareas, and we thereforeconclude that primarilyresponsible for the large increasesinside Santa BarbaraChannel Island (SBCI) reserveshave reserves:if thiswere the case, then similar increases at significantconservation benefits for spiny lobster. These the Near and Far siteswould have been observed. benefitsdeveloped within 5-6 yearsof reserveestablish- Furthermore,the magnitudeof trap yieldincreases ment,and included larger mean size, shifts in population inside reservesis remarkablysimilar to increases structurestoward larger size classes, and approximatelyobserved in the two independentscuba surveys(NPS fourto eighttimes greater trap yield (lobster/trap) inside data and our own surveys).Although mean lobster thanoutside of reserves.Similarly rapid responses to densitiesfrom our scuba surveyswere approximately reserveprotection have been observed across many taxa fivetimes greater than those from NPS surveys,relative (Halpernand Warner2002) and havebeen reported for increasesinside vs. outside were nearly identical (Fig. 3). otherspiny lobster (MacDiarmid and Breen 1993, Our densityestimates may have been higher than those Goñiet al. 2001,Follesa et al. 2008,Pande et al. 2008). of NPS becausewe workedin stratifiedareas of high Parnellet al. (2005) used fishery-independenthistorical lobsterabundance and/or because our transectswere datafrom scuba surveys to examinetemporal changes in bothlarger (very few with zero lobsters)and focused thedensity of P. interruptusat sites inside a southern explicitlyon lobstercounts. The disproportionately Californiareserve, and they observed an eightfold decrease largeincrease in trapyield inside vs. outsidereserves, from1979 to 2002 (reserveimplemented in 1971). andthe consistency between trap and scuba survey data, However,surveys conducted in 2002reported by Parnell strengthensour conclusion that the observed trap yields etal. (2005)did not reveal significantly higher densities of insidereserves were indeedpopulation level reserve legalsize P. interruptusinside vs. outside the same reserve. effectsand not an artifactof confoundedtrap perfor- Thisdisparity in temporalvs. spatialdifferences may be mancedue to spatiotemporaldifferences infishing effort explainedby a temporaldecline in lobsterabundance, or catchability.The extent to whichdifferential effort in bothinside and outside the reserve, that reflects an overall thebefore vs. afterperiods might have influenced trap decreasein reefproductivity in the region (Dayton et al. yieldis illustratedby the yield differences across time at

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Fig.5. Lengthfrequency histograms forlobsters caught during collaborative trapping surveys and concurrent port sampling of commercialcatch from Santa Cruz and Santa Rosa Islands. Fishery data for the Scorpion and Gull reserves are identical because portsampling was conducted for the entirety ofSanta Cruz Island, but catches were not segregated ata finerresolution. the replicateNear and Far sites(Fig. 2), but other populationsinside reserves(Guénette et al. 1998, temporallydynamic factors might also contributeto Botsfordet al. 2009). However,reproductive output thesedifferences. and lifetimeegg* production from reserves is influenced Althoughmany studiesdocument spiny lobster by the abundanceand populationstructure of target populationincreases inside marine reserves, relatively organisms(Tetreault and Ambrose2007, Taylor and few reportaggregate data collectedacross replicate Mcllwain2010), which we foundto varyacross reserves reserves(but see Edgarand Barrett1997, 1999, Kelly et in thisstudy (Fig. 5). Additionally,there is growing al. 2000,Mayfield et al. 2005),and to our knowledge interestin use of marinereserve populations as proxies onlytwo include data priorto reserveimplementation for unfished stocks in fisheryassessments (e.g., Morgan (Shearset al. 2006,Follesa et al. 2008).We knowof no et al. 2000,Willis and Millar2005, Wilson et al. 2010). studythat has combinedbefore vs. afterdata across Our resultsimply that spatial variation in population replicatereserves, even though spatiotemporal variabil- size structureand trapyield should be consideredand ity is an importantconsideration when measuring measuredwhen selecting reserve sites as ecologicaland reserveeffects. Spatial variability in lobsterabundance fisherybaselines. and populationstructure inside reserves has obvious We did notobserve higher trap yield or effortat sites pertinencefor conservation and biodiversityprotection, Near vs. Far fromreserve borders, and therefore butsuch patterns also haveimportant implications for concludethat spillover did not significantlyinfluence fisheries.Specifically, the potentialfor reservesto trapyield or effort distribution outside reserves. A similar increasefisheries yield throughexport of larvae is resultindicating that reserves did notenhance trap yield dependentupon increasedlifetime egg productionof outsidereserves was estimatedby Guenther(2010) at a

This content downloaded from 152.3.102.242 on Tue, 22 Sep 2015 14:45:39 UTC All use subject to JSTOR Terms and Conditions January2012 COLLABORATIVELOBSTER RESEARCH 331 geographicscale of the wholereserve network using logbookdata only.The absenceof catchand effort patternsindicative of spilloveris corroboratedby tag- recapturedata (Fig. 4) and can be explainedby several factors.The mostlikely explanation is thatthe SBCI reserveswere established only six years prior to our study, andhad not yet experienced population biomass increas- es sufficientenough to cause resourcelimitations that initiatedensity-dependent emigration (e.g., Sánchez- Lizaso et al. 2000,Shears et al. 2006).This hypothesis is supportedby recent work suggesting that, unlike rapid populationincreases observed for fished organisms inside reserves,indirect effects such as density-dependent spillovertypically develop over decadal time scales (Babcocket al. 2010).Additionally, lobster habitat at Nearsites has lower topographic relief and is structurally lesscomplex than habitat at In sites(M. Kay,unpublished data),which might restrict spillover for reserve popula- tionsthat are not critically resource limited. Finally, the spilloverof lobsterfrom reserves is enhancedby contiguousreef habitatthat connectsareas within reservesto thoselocated outside (Freeman et al. 2009). Fishingthe line for P. interruptusthat was first observed by Parnellet al. (2006) at an older(established 1971) reservein La Jolla,California, developed apd intensified in thelatter stages of the2007-2008 fishing season and was associatedwith complex habitat features near the reserveboundary (Parnell et al. 2010).We observedno suchspatial configuration of reefs at theSBCI reserve network,at least forhabitat considered exceptionally productivefor lobsterfishing. In fact,most réserve bordersin theSBCI networkwere placed in sandyareas or at considerabledistance from historically productive reefsinside reserves.Consequently, the absence of evidencefor spilloverand/or fishing the line is not surprising,especially for such young reserves. Anotherpossible explanation for the absence of spillover-drivencatch and effortpatterns, which may interactwith the timeand habitatfactors described previously,is a moderateexploitation rate for lobster outsidereserves. Spillover effects such as increasedyield and effortnear bordersare most pronouncedfor fisheriesin whichpopulations outside reserves are heavilyexploited (e.g., Goñi et al. 2010). In such fisheries,very high total mortality rates are detectable in lengthfrequency data whenthe data are truncated andcontain relatively few legal sized (or larger)lobsters andBarrett Iaccheiet al. Barrett (e.g.,Edgar 1999, 2005, Fig. 6. Commercialeffort distributionat each data (buoy) et al. 2009,Goñi et al. 2010).Our portsampling reservesite on four(Gull) or three(Scorpion, Carrington) (Fig. 5) do not indicatesuch extreme truncation and datesduring the 2008-2009 fishing season. Blue polygons are suggestthat exploitation at theChannel Islands may be areaswhere collaborative trapping took place. Commercial lowerthan in other lobster even effortwas not sampled between the Near and Far polygons spiny fishinggrounds, similar.Insets show effort withinCalifornia et al. Bevertonand (sites),but was qualitatively patterns (Iacchei 2005). (mean+ SE) beforeand afterreserve implementation, as Holt (1956) establisheda formalrelationship between measuredfrom fisherman interviews and buoysurveys, total mortality,growth rates, and catch data that respectively.MPA is themarine protected area. estimatestotal mortality (Z; naturalmortality + fishing mortality)as a functionof length frequency data and the von Bertalanffygrowth parameters k and Loo(see also

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Sparreand Venema 1998). We appliedthe Beverton and for abundancegradients inside reserves are possible Holt(1956) formula to lengthfrequency data from sites approachesfor studying spillover not detectablewith outsidereserves (Fig. 5) and estimatesof к (0.105)and fishery-dependenttechniques alone, and mergingfish- Loс (121.5 mm) froma mark-recapturegrowth study ery-dependentand fishery-independentapproaches rep- (M. C. Kay, unpublisheddata), and we estimateda Z resentsan importantfrontier for CFR. value of 0.59 forfemale lobsters at Santa Cruz and Our studyis a valuablecontribution to studiesof Santa Rosa Islands(estimates for male lobstersnot reserve-fishinginteractions because we demonstratethe available).Our estimateof Z (0.59) for femaleP. potentialfor CFR to improveecological assessments interruptuswithin the Channel Island reserve network is that informpolicy. Fishery-dependent methods and low relativeto publishedvalues for a numberof other metricsenhanced this study by allowingus to perform spinylobster fisheries. For example, Lipcius et al. (2001) a beforevs. afteranalysis, accurately and precisely reportedZ = 2.01 and 2.28 forthe Caribbeanspiny measuretrapped lobsters to the nearest1 mm (not lobster() at two sitesin the Bahamas feasiblewith diving methods), reliably compare effort (basedon averagedannual data presented in their Table and catch patternsnear borders,perform a tag- 3); Kagwade(1993) reported Z = 0.93-1.24 forPanulirus recapturestudy, access fishermen LEK, compareresults polyphagusin ;and Caputiet al. (2008)reported Z withport sampling data, and estimatetotal mortality = 1.42-2.12for Panuliruscygnus in threezones off (Z) for fishedareas. Additionally,we established WesternAustralia (we convertedfrom their harvest community-basedcapacity for monitoring future chang- ratesof 70-85% and naturalmortality = 0.22). The es to this coupled human-naturalsystem and the relativelymoderate Z valuethat we estimated for female broaderfishery. Our collaborativeapproach not only P. interruptusat Channel Islands, and the length improvedthe ecological assessment, but our ecological frequencydata thatare not completelytruncated at findingsfeedback into the humancomponent of the thelegal size limit(Fig. 5), are similarto conditionsin system.For example,fishery partners in thisstudy view fisheriesfor edwarsii in SouthAustralia (Linnane the work as usefulbecause it has increasedtheir et al. 2009я,b). Mortalityestimates for Channel Islands awarenessand trustfor science-based management. As lobsterare presented solely to accountfor the absence of a consequence,the California Lobster and Trap Fish- catch and effortincreases at borders,but are not ermen'sAssociation supports continued research at intendedas an assessmentof thefishery. SBCI, as well as expansionof our collaborative The absenceof fisherycatch and effortpatterns approachthroughout the U.S. rangeof the P. inter- indicativeof spilloveris consistentwith localized ruptusfishery, in an effortto engageits membersin movementpatterns observed for tagged and recaptured stakeholder-basedreserve monitoring, data collection lobstersat our studysites. Among lobsters initially forstock assessment, and a third-partysustainability captured,tagged, and releasedat each of theIn, Near, certification.This is a directimpact of our innovative and Far sitesat the Scorpion,Gull, or Carrington partnershipat SBCI and exemplifiesthe stewardship reserves,a vast majority were later recaptured within the thatoften arises from collaborative resource manage- originaltagging site (Fig. 4). Suchlocalized movement ment(Gutiérrez et al. 2011). supportsour conclusion that spillover was not operative Continuedspiny lobster CFR at SBCI marinereserves on a scale that influencedfishery dynamics. An is importantbecause neitherthe fisherynor lobster alternativeinterpretation of our spillover results is that populationsinside reserves are likelyto be at equilibri- lobstersmight emigrate from reserves on timescales not um.In particular,LEK of seniorfishermen at Channel coveredby our sampling,and we thereforefailed to Islandssuggests that recently increased effort is likelyto detectspillover that might indeed occur (i.e., Type II intensifyas ex-vesselprices for California spiny lobster error).For example,LEK of our fisherypartners trendupward (from ~$9 to $17 (US$) perpound from suggeststhat movementof P. interruptusincreases the 2006-2007to 2010-2011seasons) and effortis duringwinter storm events in California,whereas most concentratedas fishermen along the California mainland ofour trapping surveys were conducted in latesummer are displacedby an imminentnetwork of marine and fall.Although this is possible,commercial effort reserves.With regard to temporalchanges in lobster surveyswere conducted later in theseason and showed populations,research from older reservesin New no indicationof fishingthe line. Furthermore,we Zealandsuggests that lobster biomass will continue to explicitlytested the predictions of spilloveras a process increasein ChannelIsland reserves (Kelly et al. 2000, drivenby nonseasonalmovement due to density Shearset al. 2006), and this increasemay enhance dependence(Polacheck 1990, DeMartini1993, Sán- spillover.Due to this temporaldynamism, future chez-Lizasoet al. 2000),diffusion (Hilborn et al. 2006, monitoringat ChannelIslands should address lobster Kellneret al. 2007,Walters et al. 2007),or homeranges populationchanges inside and outside reserves, spatially that cross reserveborders (Moffitt et al. 2009). explicitcatch rates,effort distribution, and fishery- Explorationof temporallydynamic (e.g., seasonal, reserveinteractions, and a CFR approachsuch as we ontogenetic)emigration from reserves is newlydevelop- presenthere is an importanttool. CFR hasthe potential ing(Botsford et al. 2009).Expanded tagging or sampling toenhance many aspects of fisheries research and enable

This content downloaded from 152.3.102.242 on Tue, 22 Sep 2015 14:45:39 UTC All use subject to JSTOR Terms and Conditions January2012 COLLABORATIVELOBSTER RESEARCH 333 the adaptivemanagement of California'snearshore bridgingthe gap between conventional fisheries management fisheries.This is certainlytrue for spiny lobster, for andmarine protected areas. Reviews in FishBiology and whichthe California of Fishand Game is Fisheries19:69-95. Department Branch,Т. A.,O. P. Jensen,D. Ricard,Y. Ye,and R. Hilborn. developinga stockassessment and an adaptiveman- 2011.Contrasting global trends in marinefishery status agementplan. The abilityto gatherinformation and obtainedfrom catches and from stock assessments. Conser- manageadaptively will be criticalas we reach (or vationBiology 25:777-786. J. J. M. surpass)sustainable yields for most fisheries (Hilborn et Caddy, F., Csirke,S. Garcia,and R. J.R. Grainger. 1998.How pervasive is "fishing down marine food webs"? al. 2003,Mullon et al. 2005). Science282:1383a. Acknowledgments Caputi,N., R. Melville-Smith,S. de Lestang,J. How,A. Thomson,P. Stephenson, 1.Wright, and K. Donohue.2008. Thisstudy was made possible by the involvement ofmany Draftstock assessment for the West Coast rock lobster fishermenwho did not participate as authors, in particular:S. fishery. Fisheries and MarineResearch Davis,R. Kennedy,M. Becker,K. Bortolazzo,J.Peters, R. Laboratories,North Beach, Western Australia, Australia. Ellis,M. Brubaker,and broker/ombudsman T. Wahab. The CDFG (CaliforniaDepartment of Fishand Game).2008. authorsthank R. Parrish(NOAA fisheries) and K. Barsky ChannelIslands marine protected areas: first 5 yearsof (CDFG) fortheir guidance. D. Hall at HallprintTags monitoring:2003-2008. www.dfg.ca.gov/marine (Australia)salvaged numerous tagging events with expedited Conway, F. D. L., andC. P. Pomeroy.2006. Evaluating the andpersonal service. 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SupplementalMaterial Appendix ANOVAtables describing effects ofreserves on lobstertrap yield, size, and density on the seafloor, comparisons ofresearch vs. fishingtrap yields, and effort distributions before and after reserves (Ecological Archives A022-020-A1).

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