-
7/29/2019 1-s2.0-S0165783612001051-main BUDIMAN
1/8
Fisheries Research 125126 (2012)
190197ContentslistsavailableatSciVerseScienceDirect
FisheriesResearchjournal
homepage:www.elsevier.com/locate/fishres
ThereproductivecycleofOctopuscyaneainsouthwestMadagascarandimplicationsforfisheriesmanagementD.Raberinarya,b,1,S.Benbowa,aBlueVentures
Conservation, Level 2Annex, Omnibus Business Centre, 39-41North
Road, LondonN7 9DP, UKbInstitut Halieutique et des Sciences
Marines,Avenue de France, Mahavatse I, Toliara 601, Madagascara r t
i c l e i n f o a b s t r a c tArticle history:
TheOctopuscyaneafisheryisthemosteconomicallyimportantfisheryinsouthwestMadagascar.Grow-Received
25August 2011 ingconcernsoverthesustainability ofexploitation
havepromotedanumberofconservation effortstoReceived in revised form
21 February 2012Accepted 22 February 2012 improvemanagement
ofthefishery.Weanalyseoneyearofcatchdatatoidentifyseasonalvariations
insexualmaturityandkeyreproductive periodsofthespecies,
usingmicroscopic analysisofgonadtis-Keywords:Octopus
cyaneaFisheries managementReproductive cycleRecruitment
suestovalidatefieldassessmentsofmaturity.Datashowseasonalvariability
inmaturityandsizeatfirstmaturityforbothsexes,aswellastemporalchangesinthesexratioofthespecies.Maturityoccurredataminimummeanweightof2246gforfemalesand643gformales.Aclearrelationshipbetweengonadweightandtotalweightinmaleoctopusindicatesthattotalweightcanbeusedasaproxyforsexualmaturityinmales.Conversely,femalesshowhighvariability
inweightatfirstmaturityandno
Depth-refuge clearrelationship
betweentotalweightandmaturitystage.Fullysexually
maturefemaleswereverySouthwest Madagascar rare,constituting
lessthan1%ofthetotalsample.Wehypothesise
thattheartisanalfisherymaynotbe
currentlyexploitingmaturefemaleindividualsbecausefemalesretreattodeeperwaterspriortorepro-duction,thusremainingbeyondthereachofthefishery.AnabundanceofjuvenileindividualsinthecatchfromJune,andagainfromOctobertoNovember,indicatesrecruitmentpeaksatthesetwotimes.Inrecentyears,managementofthisspeciesinsouthwestMadagascarhasfocusedonshort-termclosurestofishingwithinspecifictidalreefflatareas.Identificationofthekeyphasesofthereproductive
cycleofO.cyaneainsouthwestMadagascar
mayprovidemanagerswithbiologicalevidencetosupportseasonalclosuresdesignedtoprotectkeylifestagesofthespecies.
2012ElsevierB.V.Allrightsreserved.1. Introduction Exploitation
ofoctopusstockshasincreasedrapidlysince2003
asaresultofexpandingcommercialisationofMadagascarsocto-The reef
octopus Octopus cyanea is thought to be the most
pusfishery,responsible for6070%ofthevalueofmarineresources
commonoctopusspeciesonreefsworldwideandisthedominant
purchasedbycollection
andexportcompaniesinsouthwestMada-octopusinmanycoralreefhabitatsintheIndianOceanregion(Van
gascar(LHaridon,2006).Inrecentyearslocalcommunities
haveHeukelem,1983;RoperandHochberg,1988).Howeverdespiteits
expressedconcernsabouttheover-exploitation
ofoctopusstocksabundance,researchintothebiologyofthespecieshasbeenlimited
andassociateddirectreefdamage,bothofwhichthreaten
thetra-toafewkeystudies(GuardandMgaya,2002;VanHeukelem,1973,
ditionallivelihoods
ofthelocalVezocommunitiesthatinhabitthe1983;Caverivire,2006).Thespeciesisofconsiderablecommercial
southwestcoastofMadagascar(Langley,2006;Epps,2007;Astuti,valuetoartisanalfisheriesincoastalEastAfricaandWesternIndian
1995).Fishingpressureisdepth-limited
bythenatureoftheglean-Oceanislandstates,wherecatchesaregenerallysoldthrougha
ingandspearfishingtechniques andoctopusaregenerally
onlynetworkofcollectorstoreachnationalandinternationalexport
exploitedonreefflatsinshallowintertidal
andsubtidalzones.markets(LHaridon,2006),andtherearefewalternativestomarine
Effectively managing the fishery requires a thorough
under-resourceextraction,asthearidclimatepreventsanyformoflarge-
standingofO.cyaneareproductivebiologysothatmanagementscaleagriculture.
measurescanbetargetedatkeyperiodsinthelifecycletoenhance
effectiveness.Inparticular,temporaryclosuresofindividualfishingsiteshavebeentrialledinsouthwestMadagascarsince2004andaretimedtocoincidewiththeoreticalrecruitmentandspawning
Corresponding author. Tel.: +261 34 29 100 18.E-mail addresses:
[email protected] (D.Raberinary),
[email protected] (S. Benbow).1Tel.: +44 020 7359
1287.
peaks.Todate,thetimingsoftheseclosureshavelargelybeenbasedonanecdotalreportsfromfishersandscientifictheories.Quan-titativedatatosupportthetimingoftheseclosures
willgreatlycontributetocontinued managementofthefishery.
0165-7836/$ see front matter 2012 Elsevier B.V.All rights
reserved.doi:10.1016/j.fishres.2012.02.025
-
7/29/2019 1-s2.0-S0165783612001051-main BUDIMAN
2/8
D.Raberinary, S. Benbow/ Fisheries Research 125126 (2012) 190197
191O.cyaneaaredioeceousandfemaleslaybetween150,000and
700,000eggsinasingleclutch(Caverivire,2006;VanHeukelem,1973).Maturefemaleindividualsoftenmigratefromshallowreefflats
into deeper subtidal areas for spawning (Oosthuizen
andSmale,2003;SmaleandBuchan,1981;Whitakeretal.,1991).Uponhatching,planktoniclarvaemoveintothewatercolumnforonetotwomonths,anddispersalisthoughttobewiderangingwithlar-vaetravellinguptoseveralhundredkilometresinoceancurrents(Murphyetal.,2002;Casuetal.,2002).Thusitislikelythatspawn-ingfemalescontributetothemaintenanceandreplenishmentofstocksataregionallevel,notsimplyinthevicinityofthespawn-inglocality.Priorstudiesofthereproductive
cycleofO.cyaneaintheIndianOceanarelimited.ReproductionofO.cyaneahasbeendocumentedtooccurthroughouttheyearinbothTanzaniaandMadagascarwithreproductivepeaksinJuneandDecember(GuardandMgaya,2002;Caverivire,2006)suggestingthisspeciesutilisesan
intermittent spawning strategy occurring over an
extendedperiodoftimeasdescribedbyRochaetal.(2001).
Thisstudypresentsdatafrom12monthsofmonitoringpat-ternsofsexualmaturityandrecruitmentofO.cyaneafromfisherylandingsinsouthwestMadagascartoprovideinsightintotherepro-ductivepatternsofthespeciesinthisregion,withaviewtoadvisingmanagers
on recommending timings for temporary fishery clo-sures.2.
Materialsandmethods2.1. Studyarea
ThestudyareaislocatedintheMozambique Channelonthesouthwest
coastofMadagascar,approximately 200km north
oftheregionalcityofToliara(Fig.1).Octopuscatchdataoriginatesfrom
fishers in villages along approximate 30km of
coastlinefromAmpasilava in the south toAndragnombala in the
north(Fig.1).Withinthisregion,octopusfisherspredominantlytargetthereefflatalongthenearshorefringingreefs,theshallowlagoonalreefflatsandtheoffshorebarrierreefs.Fishersselltheircatchesto
one of a number of buyers representing seafood
collectioncompanies.2.2. Samplingmethods
SamplingwasconductedfromFebruary2005toFebruary2006inthevillageofAndavadoaka.Datawerecollectedeverydaythatoctopuseswerelandedinthevillage,andfisherssurveyedwereselectedthroughhaphazard,conveniencesamplingbycommunitydatacollectorswhentheyarrivedatthelandingcollectionpoint.Itwasnotpossibletosamplealloctopusescaughtbyallfishers.Within2hoflandingthefollowinginformationwasrecordedforeachsampledoctopus:octopusweight(includinggonads),weightandlengthofextractedgonads,octopussex,andin-fieldvisualassessmentofgonadmaturity.Forarandomlyselectedsubsetofindividuals,totaloctopuslengthanddorsalmantlelengthwererecordedforonedayperspringtide.
Octopusweightwasmeasuredusingaspringbalancegradu-atedin50gintervalstoamaximumof4kg,andgonadweightwasmeasuredusinganelectronicscaleaccurateto1g.Sexwasdeter-minedbyidentificationoftheheterocotylusandspermatophoricgrooveonthethirdrightarmofmales(Mangold,1983b).Gonadlength,totaloctopuslengthanddorsalmantlelengthweremea-suredusinga1mmgraduatedtapemeasure.Gonadmaturitywasassessedthroughobservationofthemaximumsizeandcolourofthegonadsforbothsexes,thepresenceofawhiteborderatthebeginningoftheoviductglandsinfemales,andtheappearanceofspermatophores
inmales(Table1).Gonadswereextractedprior
Fig. 1. Regional map and local area map showing study location,
villages and
reefareas.torecordingofmeasurementsandfishers,whosellindividuals
perunitwetweightofcatch,weregivenasmallfeetocompensateforthereductionintotalweight.
For the randomly selected subset, gonad samples were
pre-paredforsecondarylaboratoryassessmentusingBouinsolution,andtreatedwith70%alcoholsolution
toremoveresidualstain.Sampleswereviewedusingamicroscopeat100magnification,todeterminenumber,arrangementanddiameterofspermatophores
formales,andthemaximumdiameterandlengthofoocytesforTable 1Maturity
indices formale and femaleOctopus cyanea.
Maturity stage IdentificationGonadmass Gonad appearance
MalesI Immature
-
7/29/2019 1-s2.0-S0165783612001051-main BUDIMAN
3/8
192 D.Raberinary, S. Benbow/ Fisheries Research 125126 (2012)
190197
Fig.2. Monthly gonosomatic index for field sampled O. cyanea.
Error bars represent standard error around the
monthlymean.females,usinganocularmicrometre.Thesemeasurementswere
100gisgenerallyconsidered
tobenewlyrecruited(VanHeukelem,usedtodeterminegonadmaturitystagebasedonthecharacteris-
1983),butindividuals
thissmallwererarelycapturedusingtheticsdescribedinTable1.
fishingtechniques employedinthisregion.Thereforeforthepur-
NodatawereavailableforJanuary2006duetotheregionalclo-
posesofthisanalysisoctopuscaughtweighinglessthan200gweresureoftheoctopusfisheryfrom15thDecemberto1stFebruary,
consideredtobenewlyrecruited,sincethesizedifferencebetweenthustheresultsfromFebruary2006maynotreflecttheexpected
100and200gisrepresentedbyapproximately
2weeksofgrowthseasonalfluctuations
inoctopusiffishingpressurehadbeenmain-
(Caverivire,2006).Thus,theproportion
ofthemonthlycatchthattainedduringthatperiod.DataforFebruary2005wereremoved
wassmallerthan200gwasconsidered
aproxyforoctopusrecruit-fortheseanalyses,becausealowsamplesizereducedthestatistical
menttothereef.powerofthesedata.
Inordertoassessthesexualmaturityofindividuals inrelationto
Datawerecollectedfromatotalof3253octopuses(1578female,
gonaddevelopment,gonosomatic index(GSI)wascalculated
using1675male)overthestudyperiod,gatheredfromcatchesderived
theformulaGSI=Mo/(MtMo)100,whereMoistheweightoffrom738fishersand28fishingsites.Totaloctopusanddorsalman-
theovaryandMtisthetotalwetweightoftheindividual(Guardtlelengthweremeasuredforasub-sample
of812(532male,280 andMgaya,2002).Monthlyvariations incalculated
gonosomaticfemale)andafurther131(68female,63male)octopuseswere
indicesweretestedusingat-testandKruskalWallisnonpara-randomlyselectedforlaboratoryanalysis.Allfield-sampledocto-
metricanalysis.puseswerere-assignedamaturitystagebasedonthelaboratory
Weightfrequency distributions
wereplottedtoshowdistribu-confirmationdrawnfromthesubsample.
tionofoctopusweightbysexanddifferences inthemeanwere
testedusingWelchst-test.2.3.
Dataanalysesandcalculationofsexratio,biometricandlife-historyparameters
3. Results
Sexratioswerecalculatedforeachmonthandtestedforsignifi-cantfluctuationsfromtheexpected50:50ratiousingPearsonsChiSquaredtest.
Regression analyses were carried out on the random
subsetoftheindividualsselectedforlaboratoryanalysistoinvestigatebiometric
relationships between a variety ofweight and
lengthmeasurements,andtodeterminewhethertotalweightorlength
Secondarylaboratoryassessmentsshowedthatin-fieldassess-mentofmaturitywasbetween72100%accurateforfemalesand48100%formales,dependingonmaturitystage.Overalldiffer-ences
between in-field and laboratory maturity estimates
weresignificant(t=1.69,df=117,p=
-
7/29/2019 1-s2.0-S0165783612001051-main BUDIMAN
4/8
D.Raberinary, S. Benbow/ Fisheries Research 125126 (2012) 190197
193Table 3Biometric relationships for male and female Octopus
cyanea (total length (TL), dorsalmantle length (DML), total weight
(TW), gonad weight (GW)).
f(x)M r2 p n EquationMaleTL(DML) 0.78 0.000 523 y=7.2824x1.1975
(linear)TW(TL) 0.83 0.000 523 y=0.057x2.2052 (power)TW(DML) 0.87
0.000 523 y=1.858x2.5745 (power)TW(GW) 0.87 0.000 1675
y=78.275x+185.01 (linear)FemaleTL(DML) 0.82 0.000 380
y=7.2756x0.6324 (linear)TW(TL) 0.87 0.000 380 y=0.0379x2.3004
(power)TW(DML) 0.89 0.000 380 y=1.602x2.6364 (power)TW(GW) 0.15
0.000 1578 y=14.422x+770.52 (linear)
furthermaturitystages.ItwasnotpossibletoclassifyfemalesashigherthanstageIIIinthefield.Thematuritystageofmaleswasfrequently
overestimated in in-field measurements, with
malescommonlyclassedasstageIIwhenlaboratorystudiesshowedtheywereinfactstageI.Itwasnotpossibletounderestimate
stageI,oroverestimatestageIIIindividualsastheserepresentthelowestandhigheststageswhichcanbeidentifiedinthefield,sothesevaluesarerecordedasnotapplicable(n/a)(Table2).
Wefoundstrongcorrelationsbetweentotallength(TL),dor-salmantlelength(DML)andtotalweight(TM)forbothfemalesandmales(Table3).Similarly,gonadweight(GM)inmaleswasalsostronglycorrelatedtototalweight(r2=0.87).Therelationshipbetweentotalweightandgonadweightwasverymuchweakerforfemales(r2=0.15).
The gonosomatic index (GSI) generally showed very
littlevariation throughout the study period for females
(range=0.23,variance=0.004)withtheexceptionofadistinctpeakinSeptember(Fig.
2).This peakwas very slightly significantwhen comparedwith grouped
samples from all other months (t=1.79, p=0.04).In contrast, theGSI
formales varied more throughout the
year(range=0.38,variance=0.017)andbetweenmonthvariationwasstatisticallysignificant(X2=93.57,df=9,p=
-
7/29/2019 1-s2.0-S0165783612001051-main BUDIMAN
5/8
194 D.Raberinary, S. Benbow/ Fisheries Research 125126 (2012)
190197
Fig.4. Weight distributions for male and female octopuses
sampled.Theresultsshowedtwosignificantpeaksinrecruitment,whena
greaternumberofindividualsweighinglessthan200gwerecaught(females:t=3.06,p=
-
7/29/2019 1-s2.0-S0165783612001051-main BUDIMAN
6/8
D.Raberinary, S. Benbow/ Fisheries Research 125126 (2012) 190197
195
Fig. 6. Number and percentage change in sequential stages of
maturity (assessed in the field) of male octopuses during the 12
months of sampling (total n=1677). Noindividuals are recorded for
January 2006 due to the fishery closure during this time.
Fig.7. Numberandpercentage of individualsatdifferentstages
ofmaturity (basedon fieldobservations,verified through laboratory
subset)of female octopusesduring the12 months of sampling (total
n=1581).
We found that 50% maturity of male and female
octopusesoccurredat643gand2246grespectively.Thisissimilartofind-ingsforO.vulgarisinSpain(903gand1788g;Oteroetal.,2007)butmuchhigherthanpreviousresearchonO.cyaneainTanzania(320gand600g;GuardandMgaya,2002),indicatingthatthevastmajor-ityoffemaleoctopusescaughtinAndavadoakasartisanalfisheryarelikelytobeimmature.Just8%(n=130)offemaleindividualswereheavierthan2246gwhereas67%(n=1123)ofmalescaughtwereheavierthanthe643gaveragesizeatfirstmaturity.
Fig.8. Monthly variation in O. cyanea recruitment index
(percentage of total catch
-
7/29/2019 1-s2.0-S0165783612001051-main BUDIMAN
7/8
-
7/29/2019 1-s2.0-S0165783612001051-main BUDIMAN
8/8
D.Raberinary, S. Benbow/ Fisheries Research 125126 (2012) 190197
197Katsanevakis, S.,Verriopoulos, G., 2006. Seasonal population
dynamics of Octopus
vulgaris in the eastern Mediterranean. Journal of Marine Science
63, 151160.Khallahi, O.M.F., 2001. Etude de la gamtogense chez le
poulpe Octopus vul-
garis (Cuvier, 1797). Bulletin Scientifique du Centre National
de RechercheOcanographiques et des Pches vol. XXVIII 8, 4553.
Langley, J., 2006. Vezo knowledge: traditional ecological
knowledge inAndavadoaka,
southwestMadagascar.BlueVenturesConservationReport,p.73.http://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdf.
Leite,T.,Haimovici,M.,Mather,M.,LinsOliveira,J.E.,2009.Habitat,distribution,andabundance
of the commercial octopus (Octopus insularis) in a tropical
oceanicisland, Brazil: information for management of an artisanal
fishery inside amarine protected area. Fisheries Research 98,
8591.Leporati,S.,Pecl,G.T.,Semmens,J.M.,2007.Cephalopodhatchlinggrowth:theeffectsof
initial size and seasonal temperatures. Marine Biology 151,
13751383.
LHaridon, L., 2006. Evolution de la collecte de poulpe sur la
cte Sud-Ouest deMadagascar: lments de reflexion pour une meilleure
gestion des ressources.Blue Ventures Conservation Report.
http://blueventures.org/downloads/bv-
research-report-2006-haridon-copefrito-report fr.pdf.
Mangold, K., 1983a. Food, feeding and growth in cephalopods.
Memoirs of theNational Museum ofVictoria 44, 8193.
Mangold, K., 1983b. Octopus vulgaris. In: Boyle, P.R. (Ed.),
Cephalopod Life Cycles,vol. I.Academic Press, London, pp.
335364.
Mangold, K., Boletzky, S. von., 1973.New data on reproductive
biology and growthof Octopus vulgaris. Marine Biology 19, 712.
Murphy, J.M., Balguerias, E., Key, L.N., Boyle, P.R., 2002.
Microsatellite DNA markersdiscriminate between two Octopus vulgaris
(cephalopoda: octopoda) fisheriesalong the northwestAfrican coast.
Bulletin of Marine Science 7, 545553.
Navarte, M., Gonzalez, R., Fernandez, M., 2006. Comparison of
Tehuelche octopus(Octopus tehuelchus) abundance between an
open-access fishing ground and a
marine protected area: evidence from a direct development
species. FisheriesResearch79, 112119.
Oosthuizen, A., Smale, M.J., 2003. Population biology of Octopus
vulgaris on thetemperate south-eastern coast of SouthAfrica.
Journal of the MarineBiologicalAssociation of theUnited Kingdom83,
7.
Otero,J.,Gonzlez,A.F.,Sieiro,P.M.,Guerra,A.,2007.Reproductivecycleandenergyallocation
ofOctopusvulgaris inGalicianwaters,NEAtlantic.FisheriesResearch85,
122129.
Perry, R., Walters, C.J., Boutillier, J.A., 1999. A framework
for providing scientificadvice for the management of new and
developing fisheries. Reviews in FishBiology and Fisheries9,
125150.
Rocha, F., Guerra, A., Gonzlez, A., 2001. A review of
reproductive strategies incephalopods. Biological Reviews 76,
291304.Roper,C.F.E.,Hochberg, F.G., 1988.Behaviour and systematics
ofCephalopods from
Lizard Island, Australia, based on colour and body patterns.
Malacalogia 29,153193.
Sanchez, P., Obarti, R., 1993. The biology and fishery of
Octopus vulgariscaught with clay pots on the Spanish Mediterranean
coast. In: Okutani, T.(Ed.), Recent Advances in Fisheries Biology.
Tokai University Press, Tokyo,pp. 477487.
Smale,M.J.,Buchan,P.R.,1981.BiologyofOctopusvulgarisofftheeastcoastofSouthAfrica.
MarineBiology 65, 112.
Van Heukelem, W.F., 1973. Growth and life-span of Octopus cyanea
(Mollusca:Cephalopoda). JournalofZoology 169, 299315.
Van Heukelem, W.F., 1983. Octopus cyanea. In: Boyle, P.R. (Ed.),
Cephalopod LifeCyclesVolume I.SpeciesAccounts.Academic
Press,London, pp. 267276.
Whitaker, J.D., Delancey, L.B., Jenkins, J.E., 1991.Aspects of
the biology and fisherypotential for Octopus vulgaris off the coast
of South Carolina. Bulletin of MarineScience 49, 482493.
http://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-haridon-copefrito-report_fr.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdfhttp://blueventures.org/downloads/bv-research-report-2006-langley-tek.pdf
