Exploring Relationships between Demersal Resources and Environmental Factors in the Ionian ...

13 Pages·2014·2.47 MB·English

HindawiPublishingCorporation
JournalofMarineBiology
Volume2012,ArticleID279406,12pages
doi:10.1155/2012/279406
Research Article
Exploring Relationships between Demersal Resources and
Environmental Factors in the Ionian Sea (Central Mediterranean)
G.D’Onghia,A.Giove,P.Maiorano,R.Carlucci,M.Minerva,F.Capezzuto,L.Sion,
andA.Tursi
DepartmentofBiology,UniversityofBari“AldoMoro”,ViaE.Orabona4,70125Bari,Italy
CorrespondenceshouldbeaddressedtoG.D’Onghia,[email protected]
Received21December2011;Accepted20February2012
AcademicEditor:JakovDulcˇic´
Copyright©2012G.D’Onghiaetal.ThisisanopenaccessarticledistributedundertheCreativeCommonsAttributionLicense,
whichpermitsunrestricteduse,distribution,andreproductioninanymedium,providedtheoriginalworkisproperlycited.
The relationships between the abundance of demersal resources, environmental variables, and fishing pressure in the north-
westernIonianSeainthelasttwodecadeswereevaluated.Dataonthedensitycollectedduringseventeentrawlsurveyscarried
out from 1985 to 2005 were used. The following species were considered: Aristaeomorpha foliacea, Nephrops norvegicus, and
Parapenaeus longirostris for crustaceans; Merluccius merluccius, Phycis blennoides, and Mullus barbatus for teleost fish. The
recruitmentindexwasalsoconsideredforN.norvegicus,P.longirostris,M.merlucciusandMullusbarbatus.Sixcandidatemodels
wereevaluatedforeachdensityandrecruitmentdataseteithercombiningfishingeffortwithglobal(NAO)andregional(SST
andprecipitation)climaticindices,ormodelsseparatelyinvolvingfishingeffort,NAO,orregionalclimaticindicesastheonly
predictivevariable.Modelselectionwascarriedoutusinganinformation-theoreticalapproachthatappliesAkaike’sInformation
Criterion(AIC).Highchangesovertimewereobservedforthedensitydataandrecruitmentindicesineachspecies.Apartfrom
hakeabundanceandrecruitmentdata,forwhichaclearpositiverelationshipwiththeNAOindexalonewasdetected,thechanges
observedintheotherspeciesseemtobetheconsequenceoftheinteractionbetweenbottom-upeffectslinkedtochangesinphysical
environmentandtop-downonesduetothefishingpressure.
1.Introduction Since 1985 demersal resources in Italian waters have
mostly been monitored by scientists in the context of the
Itiswellknownthattheabundanceoftheaquaticpopulation
GRUND project [5] and successively as part of MEDITS
fluctuatesoverbothspatialandtemporalscalesinrelationto
project[6].Inthisrespect,governmentsandstakeholdersask
thevariabilityofabioticandbioticfactorsintheecosystem
scientiststoprovidetheirbestadvice,intermsofindicators
as well as to human activities, such as fishing [1–3]. Larkin
andreferencepointsthatshouldtriggermanagementaction.
[4] reported that the management of marine fisheries
MostMediterraneandemersalresourcesshowhighfluctua-
according to ecosystem properties is essentially a question
tionsovertimeandareconsideredtobeparticularlysuscep-
of distinguishing the impacts of fishing (top-down effects)
tibletooverexploitation[7].Theirmanagementisbasedon
from those of fluctuations in the environment (bottom-up
effects),understandingthedynamicsofspeciesinteractions “effortcontrol”measures,includinglicencelimitation,time-
areaclosure,andgearrestrictions[8].
andappreciatingthewayinwhichfishingfleetswillrespond
to changes in the abundance of various stocks. However, Attheendofthe1980stheMediterraneanSeaunderwent
Larkin[4]himselfreportedthat,althoughchangesinmany amajorchangethatencompassedatmospheric,hydrological,
marineecosystemhavenodoubtoccurred,effortstomodel andecologicalsystemsandthatledtonewregimeconditions
their holistic dynamics have not yet been successful due to [9–11]. The most important and evident phenomenon
thenumberofvariablesinvolvedandthewayinwhichonly that involved the change in circulation and thermo-haline
small errors in their estimation may lead to large effects in properties of the water masses is the so-called Eastern
thesimulatedresults. MediterraneanTransient(EMT),correspondingtotheshift
2 JournalofMarineBiology
in deep water formation in the eastern basin from its usual toevaluatetherelationshipsbetweentheabundanceofdem-
sourceinthesouthernAdriatictoanewsourceintheAegean ersalresourcesandenvironmentalvariablesandfishingpres-
Sea[12–15].Eventhoughsuchachangeincludedthewhole sureinthenorth-westernIonianSeainthelasttwodecades.
Mediterranean,somebasinssuchastheIonian,Adriatic,and
Levantine basin were primarily involved. During the EMT
highly saline (>39.0psu), warmer (around 15◦C) and well- 2.MaterialandMethods
oxygenatedintermediatewatersflowedfromtheAegeanSea
throughtheWesternCretanArcStraits.Thisfluxinterrupted 2.1.DataCollection. Thegeographicareaconsideredinthis
thetraditionalpathofLevantineIntermediateWaters(LIW) work is the north-western Ionian Sea, along the Italian
spreading northwards into the Ionian Sea and eastwards coasts (Figure1). The time series data utilized cover the
into the Levantine basin and affecting the dynamics of period1985–2005.OnthebasisofConversietal.[19]who
the upper, intermediate, and deep layers. During the EMT report the significant change of several climatic indicators
an uprising of the nutrients close to the euphotic zone (SST, SLP, NAO index, and NHT) in the years 1986–
occurred in the northern and eastern Ionian [12, 15]. This 1988, the sea surface temperature (SST) of the Ionian Sea
large modification in the eastern Mediterranean circulation was selected as an indicator of environmental changes in
lasted approximately 10 years, until 1997 when the general this basin. Temperature is one of the primary factors in
Ioniancycloniccirculationwasrestored[15,16].Thischange determiningthedistributionpatternandabundancechanges
in circulation of the north Ionian gyre from cyclonic to in aquatic populations [31–36]. SST data were obtained
anticyclonicandviceversahasbeenrecentlysuggestedasan usingtheAVHRR(advancedveryhigh-resolutionradiome-
internalprocessoccurringbetweenAdriaticandIonianwater ter)(http://podaac.jpl.nasa.gov/).Sincethevariationsofthe
masses on a decadal scale called Bimodal Adriatic-Ionian North Atlantic Oscillation (NAO) influence the weather on
Oscillations(BiOS)[17]. a large scale over the North Atlantic and Europe and have
a strong impact on the oceanic conditions, including the
TheEMT-BiOS-relatedregimeshiftrecordedintheearly
Mediterraneanregion,thewinterNAOindexwastakenfrom
1990s involved sea surface temperature, sea level pressure,
http://www.cgd.ucar.edu/.Theprecipitationvalues(inmm),
surface and deep circulation of water masses, salinity and
nutrient changes, affecting the primary production, and oftencorrelatedtotheNAOapartfromlocalweathercondi-
tions,wererecordedfromhttp://www.ilmeteo.it/.Thevalues
pelagic community as recorded by long-term studies on
of SST, NAO index, and autumn precipitation recorded
mucilage, red tides, plankton, jellies, and anchovies in
between1985and2005arepresentedinFigure2.
the western and eastern Mediterranean basins [18–20]. In
Fishing effort is an important variable influencing the
particular, changes in the standing stock of the autotrophic
abundance of living resources [1–3]. However, this is a
biomass and zooplankton abundance with consequent
complex variable that is difficult to quantify because it is
changes in the pelagic community structure were detected
influenced by different factors. The main fisheries of the
in both the Ionian Sea [21–24] and Adriatic and Ligurian
north-westernIonianSeaareGallipoli,Taranto,andCrotone
seas[18,19,25].Theinversionofthesurfacecurrentsinthe
(Table1)(http://www.irepa.org/).Differentfishinggearsare
Ionian Sea was also thought to influence the occurrence of
used to exploit the demersalresources;however, trawlingis
species in the Adriatic Sea, including the first record of the
themaintechniqueintermsofcatchandgrosstonnage(GT).
scleractinian coral Astroides calycularis and the teleost fish
Thetrawlersareequippedwithottertrawlnets.Fishinggen-
Sparisoma cretense in this basin [26]. Civitarese et al. [18]
erallyoccurs fromMonday to Friday during daylight hours
reportedalistofbiologicalrecordsintheAdriaticindicating
only. Trawlers generally work on daily trips. Commercial
that the presence of Atlantic and western Mediterranean
haulsarecarriedoutatdifferentdepths,fromcoastalwaters
specieswasconcomitantwiththeanticyclonicnorthIonian
to700–800m.Themanagementofthefisheriesisbasedon
gyre and thus with advection of Modified Atlantic Waters effort control and mesh size regulation. Since 1988 a 30–
(MAW)intotheAdriatic,whiletheoccurrenceofLessepsian
45-day “closed season” for trawling has been implemented
species coincided with the cyclonic north Ionian gyre that
in late summer-early autumn, as a management measure
advectseasternMediterraneanwatersintotheAdriatic.
imposed by the Italian government [30, 37]. As in most
The first finding of the tropical fish Sphoeroides pachy- Mediterraneanfisheriesdataoneffortarealmostabsentand
gaster[27],Elatesransonnetii[28],andDysommabrevirostre there is a low level of enforcement of the legislation [8]. In
[29]intheIonianSeawassuggestedbyMaioranoetal.[30] thepresentstudythenumberofdays-at-seaperyearhasbeen
as a consequence of the water masses variations occurring usedasameasureoffishingeffort.Thismaybeafairmeasure
in the Ionian basin during EMT-BiOS. Moreover, these of fishing effort for demersal gears [38]. Nonworking days
authorsreviewedthechangesofdemersalspeciesabundance were made up of days closed by management, days with
and faunal assemblage structure over the last twenty years. wind speeds >15 knots, public holidays, Saturdays, and
The causes of these changes were suggested to be linked Sundays. The wind intensity values were obtained from
to the interaction between the bioecology of the species http://www.ilmeteo.it/.Thetrendofdays-at-seaovertimeis
and environmental factors as well as fishing pressure in presentedinFigure3.
the study areas. However, no direct relationships between Data on the demersal resources used in this work were
environmentalvariablesandspeciesabundancechangeswere collected during seventeen trawl surveys carried out in the
examined.Withthisregard,thisworkrepresentsanattempt north-western Ionian Sea, from 1985 to 2005, as part of
JournalofMarineBiology 3
Table1:Numberofvesselsandmeangrosstonnage(GT)fordifferentgearsrecordedinthemainfisheries(Gallipoli,Taranto,andCrotone)
ofthenorth-westernIonianSea.
Trawling Longline Gillnet Purseseine
Fisheries
N.vessels MeanGT N.vessels MeanGT N.vessels MeanGT N.vessels MeanGT
Gallipoli 75 11.61 16 8.22 313 3.58 — —
Taranto 53 9.27 2 9.43 118 2.65 6 8.92
Crotone 95 18.55 16 9.31 262 2.71 — —
a national study project on the assessment of demersal trade-off between bias and variance. Considering the small
resources [5]. All the trawl surveys were carried out in size of the sample with respect to the number of estimated
autumn.Atrawlnetwithastretchedmeshof40mminthe parameters in each model, the second-order information
codend was used and the experimental hauls were carried criterion (AICc) was applied and AIC differences (Δi) and
out according to a random-stratified sampling design at Akaikeweights(wi)werealsobasedonAICc estimates.The
depthsbetween10and800m[30,37].TheScanmarSonar best approximating model for each data set was selected by
System was applied on the trawl net in order to estimate rankingofAkaikeweights.Forfurthercomparisonbetween
the horizontal net opening during each experimental haul the candidate models in terms of variation explained,
[39].Abundancedatawerestandardisedtothesweptsurface adjustedR2valuesarealsoreported.Regressionanalyseswere
unit[40].Thus,density(N/km2)indiceswerecomputedfor performedusingSPSSversion18[46].
eachspeciesandsurvey.Thefollowingspecieswereconsid-
ered: Aristaeomorpha foliacea, Parapenaeus longirostris, and
Nephrops norvegicus for crustaceans; Merluccius merluccius, 3.Results
Phycisblennoides,andMullusbarbatusforteleostfish.These
species were chosen because they are of great commercial Fluctuatingchangesovertimewereobservedforthedensity
interestandcomprisethebulkofthecommercialtrawlcatch incrustacean(Figure5(a))andfishspecies(Figure5(b))and
inthenorth-westernIonianfisheries.ApartfromM.barba- recruitmentindices(Figure6).
tus and, to a lesser extent, M. merluccius and P. blennoides, A summary of the six candidate models applied to
they are exclusively captured by trawling (Figure4). They explain the relationships between the density index and RI
arealsoamongthemoststudiedcommercialspeciesinthis ofeachspeciesandtheenvironmentalvariablesandfishing
Mediterraneanbasin.SincetherecruitmentofP.longirostris, effortisreportedinTables2(a)and2(b),respectively.Table3
N.norvegicus,M.merluccius,andM.barbatusoccursinthe reportstheestimatedregressioncoefficientsandtherelative
study area during autumn, the recruitment index (RI) for standarderrorsinthemodelsselectedonthebasisofranking
these species was calculated as the abundance (N/km2) of ofΔiAICc.
specimensyoungerthan1year(0+group)[41,42]collected Ranking of Δi for A. foliacea density data led to the
duringeachsurvey.AlthoughtheshrimpAristeusantennatus selection of the model involving fishing effort and NAO as
isoneofthemostabundantdeep-waterspeciesinthenorth- predictivevariables,inverselyanddirectlyrelatedtodensity
westernIonianSea,ithasnotbeenconsideredinthisstudy indices,respectively.ComparablevaluesofAICcandwiwere
duetoitsverywidedepthdistributionthatoftenmakesboth also estimated for the model including the fishing effort as
adultstockandrecruitmentlessavailabletotrawling[43,44]. theonlypredictivevariable.
ForN.norvegicusdata,theselectedmodelhighlightsan
inverserelationshipwithfishingeffortandadependenceon
2.2. Data Processing. With the purpose of trying to gain theregionalclimaticfactors(inverselytoSSTanddirectlyto
insight into which combination of the environmental vari- precipitation).
ables considered and fishing effort could determine the The best approximating model for P. longirostris is that
observed changes in the abundance data, multiple linear including the fishing effort together with global (NAO)
regression analyses were carried out. For each density and and regional (SST and precipitation) climatic indices. This
recruitment data set (response variables), six candidate model indicated inverse relationships between density and
modelswereevaluated.Besidestheglobalmodel,weselected fishing effort and between density and SST and positive
models that combined fishing effort with global (NAO) or relationshipsbetweendensityandNAOandbetweendensity
regional(SSTandprecipitation)climaticindices,andmodels andprecipitation.
involving NAO or regional climatic indices, separately, as The best approximating model for M. merluccius was
theonlypredictivevariablefishingeffort.Inordertoreduce chosen to be that consisting of a direct regression with the
the effect of extreme abundance data, these were log- NAOindex.
transformed. ForP.blennoidesdensitydata,comparablevaluesofAICc
Model selection was carried out in the framework of and wi were estimated for the model including as the only
information-theoreticapproach[45],whichappliesAkaike’s predictive variable the fishing effort (inverse) or the NAO
InformationCriterion(AIC)toselectthemostparsimonious index (direct). However, in both cases very low R2 values
model, that is, the one that achieves the most proper wereobtained.
4 JournalofMarineBiology
Table2:(a)SummaryofthesixcandidatemodelsfordensitydataofthespeciesinvestigatedintheIonianSea,includingAICc,Δivalues
forAICc,Akaikeweights(wi),basedonAICc,andadjustedR2values.ModelsareorderedintermsofΔiforAICc.Predictivevariablesare:
1:fishingeffort,2:NAOindex,3:SSTand4:precipitation.(b)Summaryofthesixcandidatemodelsforrecruitmentindexofthespecies
investigatedintheIonianSea,includingAICc,ΔivaluesforAICc,Akaikeweights(wi),basedonAICc,andadjustedR2 values.Modelsare
orderedintermsofΔiforAICc.Predictivevariablesare:1:fishingeffort,2:NAOindex,3:SSTand4:precipitation.
(a)
Dependentvariable Model AlCc ΔiAlCc wi AdjustedR2
{1 2} 9.168 0.000 0.408 0.434
{1} 9.381 0.213 0.367 0.397
Densityindex(ln) {1 2 3 4} 11.728 2.560 0.113 0.389
Aristaeomrophafoliacea {1 3 4} 12.059 2.891 0.096 0.352
{3 4} 16.315 7.147 0.011 0.116
{2} 18.228 9.060 0.004 −0.048
{1 3 4} 167.605 0.000 0.442 0.498
{1} 169.025 1.420 0.217 0.402
DensityindexNephrops {1 2 3 4} 169.329 1.724 0.187 0.465
norvegicus {1 2} 170.446 2.841 0.107 0.380
{3 4} 172.140 4.535 0.046 0.316
{2} 178.854 11.249 0.002 −0.067
{1 2 3 4} 5.772 0.000 0.555 0.235
{1 2} 7.776 2.004 0.204 0.263
Densityindex(ln) {1 3 4} 8.510 2.738 0.141 0.171
Parapenaeuslongirostris {3 4} 10.470 4.698 0.053 0.136
{1} 11.131 5.359 0.038 0.162
{2} 13.832 8.060 0.010 0.017
{2} 203.555 0.000 0.454 0.306
{1 2 3 4} 204.465 0.910 0.288 0.357
DensityindexMerluccius {1 2} 205.453 1.898 0.176 0.261
merluccius {3 4} 208.339 4.784 0.042 0.125
{1 3 4} 209.139 5.584 0.028 0.121
{1} 210.826 7.271 0.012 −0.064
{1} 181.760 0.000 0.329 −0.060
{2} 181.827 0.067 0.318 −0.064
DensityindexPhycis {3 4} 183.444 1.684 0.142 −0.115
blennoides {1 2} 183.738 1.979 0.122 −0.134
{1 3 4} 184.990 3.230 0.065 −0.169
{1 2 3 4} 186.990 5.230 0.024 −0.266
{1} −1.962 0.000 0.455 0.098
{1 2} −0.847 1.115 0.260 0.083
Densityindex(ln) {2} 0.610 2.571 0.126 −0.049
Mullusbarbatus {1 3 4} 1.947 3.909 0.064 −0.035
{3 4} 2.121 4.082 0.059 −0.092
{1 2 3 4} 3.147 5.109 0.035 −0.070
(b)
Dependentvariable Model AlCc ΔiAlCc wi AdjustedR2
{1} 15.095 0.000 0.447 0.476
{1 3 4} 16.546 1.451 0.216 0.480
Recruitmentindex(ln) {1 2} 16.653 1.558 0.205 0.453
Parapenaeuslongirostris {1 2 3 4} 17.601 2.506 0.128 0.467
{3 4} 25.136 10.041 0.003 0.099
{2} 27.166 12.070 0.001 −0.066
JournalofMarineBiology 5
(b) Continued.
Dependentvariable Model AlCc ΔiAlCc wi AdjustedR2
{1 3 4} 151.903 0.000 0.465 0.481
{1 2 3 4} 153.319 1.416 0.229 0.457
Recruitmentindex {1} 154.145 2.242 0.152 0.350
Nephropsnorvegicus {3 4} 155.191 3.288 0.090 0.342
{1 2} 155.976 4.073 0.061 0.311
{2} 161.916 10.013 0.003 −0.026
{2} 184.842 0.000 0.539 0.293
{1 2} 186.792 1.950 0.203 0.245
Recruitmentindex {1 2 3 4} 187.136 2.294 0.171 0.290
Merlucciusmerluccius {3 4} 189.820 4.978 0.045 0.098
{1 3 4} 191.050 6.208 0.024 0.072
{1} 191.765 6.923 0.017 −0.062
{2} 8.891 0.000 0.324 −0.070
{1} 9.017 0.126 0.305 −0.079
Recruitmentindex(ln) {3 4} 10.371 1.480 0.155 −0.124
Mullusbarbatus {1 2} 10.674 1.783 0.133 −0.149
{1 3 4} 12.355 3.464 0.057 −0.235
{1 2 3 4} 13.961 5.070 0.026 −0.335
For M. barbatus, the best approximating model from encompassing more trophic levels and different Mediter-
ranking of Δi and wi was that involving the fishing effort raneanbasins(see[11,18],andreferencestherein[19,24,25,
as the only inverse predictive variable, but the value of R2 49, 50]). The environmental change observed in the Ionian
estimatedwasalsoverylowinthiscase. Sea at the end of 1980s could have affected recruitment
Withrespecttorecruitmentdata,thebestapproximating directly,throughitseffectsonthesurvivalrateofindividuals
modelforP.longirostriscontemplatesaninverserelationship from eggs to juveniles, or indirectly, through bottom-up
withfishingeffort. effects of the changes in zooplanktonic species [51]. The
For N. norvegicus recruits, the same dependence on fluctuationsobservedintheabundanceindicesofthewhole
fishing effort and regional climatic indices as the whole sampled populations most probably reflect the changes in
populationwasfound. recruitment,whichrepresent the bulkof the stock in many
ThebestapproximatingmodelforM.merlucciusrecruit- demersalspecies[8].
mentdatatestifiedadirectrelationshipwithNAO. In this respect, even though at the end of the 1980s
ForM.barbatusdata,similarresultswerefoundfortwo the abrupt changes of SST, salinity, and density as well as
models including only one predictive variable each (NAO nutrients of the Ionian water masses together with large
and fishing effort), but both showed very low R2 values, as scaleclimaticdrivers(NAO)couldhavehadbothdirectand
inthecaseofthewholepopulation. indirect effects on the recruitment and abundance of the
demersalresources,theroleplayedbythefishingpressurein
4.DiscussionandConclusions theareaseemstobeimportant.Infact,theanalysiscarried
outthroughthemodelselectioncriteriadetectedsignificant
At the end of the 1980s the Ionian Sea experienced a relationships between the density of some species and both
strongmodificationofitswatermasspropertiesandthermo- climatic factors and fishing effort. In all these species an
haline circulation [9, 12, 14, 15, 18]. The present results increaseintheirabundanceswassignificantlyrelatedtothe
highlightthatintheperiod1985–2005theSSTintheIonian decreaseintheworkingdaysrecordedoverthestudyperiod.
did not show the increasing trend that characterized the Although population mechanisms act in very-high-
Mediterraneanasawhole[47].Onthecontrary,attheend dimensionnaturalsystemsofgreatcomplexitywhichmake
ofthe1980s,thereversaloftheIoniangyrecausedagreater itdifficulttodisentangleenvironmentalfromfishingeffects
inputofcolderandlesssalineAtlanticwatersintothisbasin as well as the nature of their interactions [52–54], there
[18]. Only at the end of EMT did the SST values begin to is no doubt that a decrease in the fishing pressure can
matchthesurfacewatersofthewholeMediterranean[48]. favour an increase in the abundance of fish and shellfish
All the demersal species considered in this study exhib- stocks [2, 3]. However, the effect due to the reduction of
itedoscillatingchangesintheirpopulationandrecruitment fishing varies greatly between the different species. In this
abundance. respect,theeffectobservedinA.foliacea,P.longirostris,and
The regime shift recorded at the end of the 1980s has N. norvegicus seems to be realistically causal. In fact, these
had direct effects on several marine ecosystem components crustacean stocks show overfishing conditions in both the
6 JournalofMarineBiology
Table3:EstimatedregressioncoefficientsandrelativestandarderrorsinthemodelsselectedonthebasisofAICc,ΔiforAICcandAkaike
weights(wi)values.Predictivevariablesare:1:fishingeffort,2:NAOindex,3:SSTand4:precipitation.
Dependentvariable Predictivevariable β Standarderror
(constant) 13.552 2.293
Densityindex(ln)A.
1 −0.035 0.010
foliacea
2 0.214 0.154
(constant) 2127.840 660.323
DensityindexN. 1 −2.693 1.091
norvegicus 3 −58.976 30.678
4 0.355 0.495
(constant) 16.037 5.567
1 −0.015 0.009
Densityindex(ln)P.
2 0.183 0.126
longirostris
3 −0.253 0.258
4 0.002 0.004
Densityindex (constant) 548.557 93.731
M.merluccius 2 122.402 43.096
Densityindex (constant) 369.053 369.330
P.blennoides 1 −0.489 1.572
Densityindex(ln) (constant) 10.631 1.662
M.barbatus 1 −0.012 0.007
Recruitmentindex(ln) (constant) 16.073 2.745
P.longirostris 1 −0.046 0.012
(constant) 1434.513 416.091
RecruitmentindexN. 1 −1.497 0.687
norvegicus 3 −45.165 19.331
4 0.036 0.312
Recruitmentindex (constant) 232.864 54.057
M.merluccius 2 68.711 24.855
Recruitmentindex(ln) (constant) 6.916 0.352
M.barbatus 2 −0.058 0.148
IonianSea[30,37,55–58]andItalianwaters[7].Duetothe has also been proved for other Mediterranean demersal
highvulnerabilitytotrawlingofbothadultsandjuvenilesin species[62,63],probablytemperaturealonecannotexplain
allthreespecies,areductionintheworkingdaysofthistype the change in the N. norvegicus abundance observed in
offishingcouldreliablyfavourstockrecoveryand,thus,an the Ionian Sea. Maynou and Sarda` [64] found significant
abundanceincrease.However,Capezzutoetal.[59]revealed relationships between the catch rate of this crustacean and
that the increase in the abundance of A. foliacea verified some environmental factors, such as atmospheric pressure,
mostlyintheperiod2000–2004correlatedsignificantlywith cloud cover, and sea-weather conditions. The precipitation
theincreaseintemperatureandsalinitydetectedfrom1995 effect detected in this work could be related to cloud
to2005between200and800mintheIonianSea.Thisfact cover observed by Maynou and Sarda` [64]. These authors
can be explained according to Ghidalia and Bourgois [60] hypothesizedthatthelightintensityreachingtheseabedhas
andBombace[61]byahydrologicalhypothesis:A.foliaceais aprimaryinfluenceontheactivityratesofN.norvegicusand
preferentiallydistributedinwarmerandhigh-salinitywaters. thusonitsvulnerabilitytotrawling.UngaroandGramolini
Temperature and salinity between 200 and 800m were not [65] found a possible influence of the water temperatureat
consideredinthepresentwork. sea bottoms on the spatial distribution of P. longirostris in
InN.norvegicusaninverserelationshipbetweendensity thesouthernAdriatic.However,theseauthors,inagreement
and both fishing effort and SST was detected while a direct with Abello´ et al. [66], reported that spatial differences in
relationship with precipitation came out. With reference to abundance could also be due to the fishery exploitation.
thefirstdriver,sincethecatchofNorwaylobsterintheIonian Recently, Ligas et al. [67] reported an increasing trend
Sea mostly consists of juveniles [58], this relationship also of P. longirostris abundance correlated to a rise in SST,
seems to be realistic for recruitment. In respect to the SST, a corresponding decrease of wind circulation and to the
althoughtheinfluenceofthetemperatureontherecruitment reductionoffishingeffort.
JournalofMarineBiology 7
16◦1 17◦2 18◦3
40◦5
40◦5
39◦4
39◦4
38◦3
38◦3
37◦2
37◦2
16◦1 17◦2 18◦3
10 0 10 Miles
100 m
200 m
500 m
1000 m
Figure1:Mapofthestudyarea.
InM.merluccius,bothforthewholesampledpopulation [19, and references therein] [51, 68–70]). The changes in
and recruits, a positive relationship with the NAO index environmental factors (atmospheric pressure, temperature,
wasdetected.Theexistenceofpositiverelationshipsbetween wind,precipitation,etc.)linkedtotheNAOcancausedeep
ecological processes in the sea and NAO was observed modifications to the ecosystem influencing, both directly
in many studies, even though the underlying mechanisms andindirectly,individualorganismsandpopulationsaswell
often remain unexplained ([18, and references therein], as interspecies relationships and trophic webs [51, 70, 71].
8 JournalofMarineBiology
8 26
4 24
O C)
NA 0 ◦T ( 22
S
S
−4 20
−8 18
1985 1986 1988 1990 1991 1992 1994 1995 1996 1997 1998 2000 2001 2002 2003 2004 2005 1985 1986 1988 1990 1991 1992 1994 1995 1996 1997 1998 2000 2001 2002 2003 2004 2005
m) 400
m
on ( 300
ati
cipit 200
e
pr
mn 100
u
ut
A 0
5 6 8 0 1 2 4 5 6 7 8 0 1 2 3 4 5
8 8 8 9 9 9 9 9 9 9 9 0 0 0 0 0 0
9 9 9 9 9 9 9 9 9 9 9 0 0 0 0 0 0
1 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2
Figure2:NAOindexandSeaSurfaceTemperature(SST)andautumnprecipitationrecordedintheperiod1985–2005forthenorth-western
IonianSea.
a) 350
e
ys-at-s 300
a
d
ort ( 250
eff
g 200
n
hi
Fis 150
5 6 8 0 1 2 4 5 6 7 8 0 1 2 3 4 5
8 8 8 9 9 9 9 9 9 9 9 0 0 0 0 0 0
9 9 9 9 9 9 9 9 9 9 9 0 0 0 0 0 0
1 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2
Figure 3: Fishing effort expressed in terms of the number of
workdaysinthenorth-westernIonianSeafrom1985to2005.
Figure 4: Percentage of commercial catches in some demersal
species obtained with different gears in the north-western Ionian
Sea.
The effects at the first levels of the trophic chain, that is
on very short-lived species, are probably easier to identify Maynou [76] found a positive correlation between
([18, and references therein] [25, 68]) while those on fish the average annual NAO index and the annual catches
and other high-trophic level organisms are far from being of the shrimp Aristeus antennatus in six ports along the
well understood. In the Mediterranean, Lloret et al. [72] Catalonia coast. This author hypothesized that the NAO-
reported that the recruitment of continental shelf species, induced environmental variability could favour secondary
amongwhich,E.cirrhosa,M.merluccius,andM.poutassou, productionenhancingthefoodsupplyforA.antennatus.The
waspositivelycorrelatedwiththeriverrun-offand/orwind greaterfoodavailabilityforthisshrimpwouldstrengthenthe
mixing index. Orsi Relini et al. [73] found a positive reproductive potential of a particular year class resulting in
relationshipbetweenthesummerbiomassofE.cirrhosaand increasedcatches1to3yearslater.
winterNAOindex.AdirectinfluenceofNAOonthewater Asimilarhypothesiscouldalsobesuggestedforhakein
mass circulation and consequent effects on distribution of the Ionian Sea in agreement with the fact that the changes
M. merluccius was established by Abella et al. [74] from during1987-1988affectedthefeaturesoftheplanktoncom-
the Tyrrhenian to Ligurian Sea and on distribution of M. munities.Indeed,anenrichmentoftheepipelagicsystem,in
merlucciusandAristeusantennatusbyMassut´ıetal.[75]off terms of chlorophyll a concentrations, primary production
theBalearicIslands. rates, ciliate abundance and zooplankton biomass, was
JournalofMarineBiology 9
(a) (b)
Figure5:(a)Densityindiceswithstandarddeviationincrustaceanspeciescaughtfrom1985to2005inthenorth-westernIonianSea.(b)
Densityindiceswithstandarddeviationinfishspeciescaughtfrom1985to2005inthenorth-westernIonianSea.
detectedduringtheyearsofcycloniccirculationofthewater effortandNAOindexseemtoplayacomparableroleonits
masses along the north-western Ionian coasts [18, 21–24]. abundancechanges.ThesamewasdetectedforP.blennoides.
Hakes feed actively in neritic water during their yearly life However,theverylowR2valuesestimatedinthemodelsfor
stagesandthehigherabundanceoftheirpotentialpreycan these two fishes would indicate that the set of explanatory
favour the recruitment success according to the “match- variables selected does not completely explain the variance
mismatch hypothesis” [1]. The marked increase in density in the abundance data of these species. In fact, mostly for
observed in this fish during 1990 can be considered a M.barbatus,theabundanceintheIonianSeacouldalsobe
consequenceofsuccessfulrecruitment.Nurseryareasofthis affected by small vessels mostly working with gillnets the
specieshavebeenidentifiedalongthenorth-westernIonian effortofwhichhasnotbeenconsideredinthepresentwork.
Sea [41, 42]. Thus, the period of higher food availability, In conclusion, apart from hake abundance and recruit-
indirectly related to positive NAO index, would lead to ment data, for which a clear positive relationship was
enhancement of the recruitment in M. merluccius. The detected with the NAO index alone, the changes observed
remarkable presence of juveniles in the catches of hake in the other species appear to be an interaction between
plays more in favour of environmental factors than fishing bottom-upeffectslinkedtochangesinphysicalenvironment
pressureasmaincauseoftheirabundanceincrease. and top-down ones due to fishing. In agreement with
In the stock of M. barbatus of the Strait of Sicily it Gislason et al. [77], the effects of fishing seem to manifest
was shown that, for a given spawning stock level, higher themselves within a background of natural environmental
recruitment levels correspond to positive SST anomalies variationsthatarethemajoragentofchangeinthesystem.
[62].InthestockofM.barbatusintheIonianSeathefishing Analyzing the changes in different species, in contrasting
10 JournalofMarineBiology
partoftheprogramme“AssessmentofDemersalResources.”
StatisticalanalysesweresupportedbyagranttotheUniver-
sity of Bari (Local Research Unit CoNISMa) as part of the
VECTORproject(vector.conismamibi.it).
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