biologi al y lesintheCanaryIslandregion C.Troupin
a,
∗
,P.Sangràb
andJ.Arísteguib
a
UniversitédeLiège,GeoHydrodynami sandEnvironmentResear h(B5a),B4000Liège,Belgium
b
Fa ultaddeCien iasdelMar,UniversidaddeLasPalmasdeGranCanaria,35017LasPalmasdeGranCanaria,Spain
Abstra t
TheCanaryIslandregionisri hinmesos ale phenomenathatae t y lesofphysi aland biologi alpro esses.A 1Dversion
of theRegional O eani Modeling System(ROMS)is usedsouthof theGranCanaria Islandto simulateseasonal limatologies
of these y les.The modelis for ed with monthlyair-sea uxes averaged from1993 to 2002 and initialized with mean insitu
prolesoftemperature,salinity,oxygenandnitrate on entrations.TheK-ProleParameterization(KPP)mixed-layersubmodel
is ompared withother submodels usingidealized numeri al experiments.When for ed with realisti air-sea uxes, the model
orre tlyreprodu estheannual y leoftemperature(mixedlayerdepth),withminimumsurfa evaluesof18
◦
C(maximaldepth
>
105m)inFebruaryduring onve tivemixingresultingfromanegativeheatux.Maximumtemperaturesabove23◦
C(minimal
depth
<
20m)aresimulatedfromSeptembertoO toberafterstrongsummerheatingandade reaseinTradeWindsintensity. Asimplee osystemmodelis oupledtothephysi almodel,whi hprovidessimulatedbiologi al y lesthatareinagreementwithregional observations.Aphytoplanktonbloomdevelopsinlatewinter,drivenbytheinje tionofnewnutrientsintotheeuphoti
layer.Chlorophyllhasadeepmaximumu tuatingaround100mwith on entrationsaround1mg
Chl
a
m
−3
,whilesurfa evalues
arelow(around0.1mg
Chl
a
m
−3
)duringmostoftheyear.Thephysi alandbiologi almodelresultsarevalidatedby omparisons
withdatafromregional ampaigns, limatologi aleldsandtime-seriesfromtheESTOCstation.
Keywords: CanaryIslands,Mixedlayerdepth,ROMS1Dbio-physi almodel,biologi al y les,KPPmodel,NPZDmodel,Latitude
27.5
◦
N,longitude15.5◦
W PACS:92.60.Aa,92.10.Lq,92.60.C ,92.05.Fg 1. Introdu tionTheCanaryIslandAr hipelagois omposedofseven
is-landslo atedinthepassageofthe ool,southwestward
Ca-naryCurrent(CC), onsideredastheeasternboundaryof
theNorthAtlanti subtropi algyre.Thestudyofphysi al
and biologi al y les in this regionis an importanttopi
onsideringthenumerousphysi alpro essesthatinuen e
these y les.
Sin ethe1990's,manystudieswere arriedoutinthe
Ca-naryIslandareainordertoexamineintera tionsbetween
physi alandbiologi alpro esses(e.g.Arísteguietal.,1994,
1997; Arístegui and Montero, 2005; Barton et al., 2004;
Pelegrí et al., 2005a,b). Alongwith these in situ studies, time-series data were a quired throughmooring stations,
su hastheEuropeanStationforTime-SeriesintheO ean,
∗
Correspondingauthor.
Emailaddress: troupinulg.a .be (C.Troupin).
CanaryIslands(ESTOC,Fig.1),lo ated100kmnorthof
GranCanaria(Neueretal.,2007).Satelliteimageryfrom theregionwasusedto onrmtheseobservations(Barton etal.,2001;Davenportetal.,2002).
Braun(1976) arried outa survey5miles o Tenerife from1971to1972betweentheo eansurfa eand1200
me-ters.Lowlevelsofnutrientsalongwitha learseasonal
vari-ationofphytoplanktonwerereported, onrmingthe
olig-otrophi onditionsofCanaryIslandwaters(e.g.DeLeón andBraun,1973;Arísteguietal.,1997).
Summeris hara terizedbyasharpseasonalthermo line
that preventsthe inje tionof nutrientsinto the euphoti
zone,resultinginlowphytoplankton on entrations.
Maxi-malphytoplankton on entrationsappearinlatewinter
un-dertheee tof onve tivemixingandthermo lineerosion,
whi h makenutrientsavailable for phytoplankton. These
high on entrationswerealsoobservedbyHernández-León et al.(1984)insouthern Canarywaters,Hernández-León etal.(1984)onthesoutheasternshelfoftheGranCanaria
Island,andArísteguietal.(2001)in oastalwaterseastof GranCanaria.
Despite interest in understanding biologi al y les in
this area, only a few oupled physi al-biologi al models
have been developed. Zielinski et al. (2002) used a one-dimensional model in order to test dierent
parameteri-zations of the photosyntheti ally a tive radiation (PAR)
atESTOCstation.Althoughtheirmodelprodu ed
realis-ti resultsfortemperatureand hlorophyll on entration,
there wasno validation of the mixed layer depth model,
whi h isof ru ial importan e forthemodulationof
bio- hemi al y les.
Bahamón and Cruzado (2003) ompared the NW Mediterranean and NE Atlanti using a one-dimensional
model, but theirresults forthe physi al variables donot
representa uratelytheseasonal y lesinthese ond
sys-tem.Theyshowedstrati ationstartinginApril,whi his
tooearly,andtwomaximaforthephytoplankton.Therst
maximumwasinwinterandthese ondinfall,asituation
thatismoretypi alforhigherlatitudes.
Therst obje tiveof thepresent work wastosimulate
theseasonal y leoftheuppero eani layersimilartowhat
wasobserved in theCanaryIslandregion. Tothisend, a
one-dimensional physi al-biologi al oupled model for ed
withre entlydeveloped limatologiesforair-seauxes(e.g.
OAFluxesforheat,Tab.2)andinitializedwith
hara ter-isti onditionsofthisparti ularregionwasused.
ApointsouthofGranCanaria(Fig.1)wassele tedfor
modelimplementationbe ausethebiologi alpropertiesin
this area were more frequently sampled. The model
im-plementation site is lo ated in the transitional zone
be-tweentheupwellingri hwatersandtheoligotrophi
inte-rioro ean.Itmaybeae tedbymesos alevariability
in-trodu edbytheAr hipelago,buttheseee tswereltered
outinthe limatologi alobservations.Be auseofthe
re og-nition that dynami sof theo ean upperlayerare
funda-mentalforthemodulationofbio hemi aluxes,spe ial
at-tentionwaspaidtothemixedlayerseasonal y le,making
omparisons withother physi al model outputsand with
availablemeasurements.
A se ondobje tivewastheestablishmentofaseasonal
limatology of biogeo hemi al properties of the o eani
upper layer, whi h will help in implementing a
three-dimensional physi al-biogeo hemi almodel urrently
un-der development(Mason et al., 2008b,a), asis a oupled larvaemodel fo usedonthe sameregion (Bro hieret al.,
2008).
To a hieve these goals, the work is divided into ve
parts: the rst is the des ription of the dierent
ompo-nents(physi s,biology,mixedlayer)ofthemodel,aswell
astheassumptionsbehindthem(Se tion2). Nextdetails
ofthemodelsetuparedes ribedbyoutliningfor ingand
initialization onditions(Se tion3),asthesearekeyto
gen-eratingrealisti solutions.Themodelbehavioristestedin
idealizedexperiments(Se tion4)andtheresultingannual
eldsarepresented(Se tion5)andvalidated(Se tion6).
18
o
W
17
o
W
16
o
W
15
o
W
14
o
W
13
o
W
26
o
N
27
o
N
28
o
N
29
o
N
30
o
N
T
P
H
G
GC
F
L
Fig.1.TheCanaryAr hipelago:P=LaPalma,H=ElHierro,G=
LaGomera,T=Tenerife,GC=GranCanaria,F=Fuerteventura,
L = Lanzarote. The region of interest is lo ated south to Gran
Canaria(square).Cir le denotes ESTOCstation and trianglesare
thelo ationsoftheprolesusedintheinitialization.
2. Modeldes ription
Simulations were arried outwith the one-dimensional
version of ROMS (Regional O ean Modeling System,
Sh hepetkin and M Williams, 2005); the mixing losure sub-model,treatmentsof verti aladve tionand diusion
are ompatible between the 3D and the 1D model. A
stret hedverti al oordinatesystemprovidesanin reased
resolutionnearthesurfa eboundarylayer(Penven,2006).
2.1. Hydrodynami smodel
The hydrodynami s model omputes velo ity
ompo-nents(
u
andv
),temperature(T
)andsalinity(S
).Density isderivedfromthestateequationofJa kettandM Dougall(1995).Hydrostati and
f
-planeapproximationsaremade. Twoimportant onsequen es arise from theuse of a1-Dmodel:
(i) theverti alvelo ityisnulleverywhere:
w = 0
, (ii) adve tionisnotmodeled(u · ∇)X = 0
,withu
, thevelo ityve tor,and
X
,anys alarvariable.Temperatureandsalinityarebothrelaxedtowardtheir
ini-tialvaluewitharelaxationtime
τ
dependingofthedepth:τ = 2+23 exp(z/100)
.Thisformulagivesavalueofτ
about 5yearsbelow200mandupto25yearsatthesurfa e.2.2. Mixedlayermodel
Small-s ale pro esses are parametrized with the
K-Prole Parametrization (KPP, Large et al., 1994). With this s heme, the mixed-layerdepth (MLD hereinafter) is
denedasthedepth
d
atwhi hthebulkRi hardson num-ber, whi h measures the ratio between strati ation andRi
b
(d) =
[B
r
− B(d)]d
|V
r
− V(d)|
2
+ V
2
t
(d)
= Ri
cr
= 0.3,
(1) whereB
r
andV
r
are estimatesof the averagebuoyan y
andvelo ity,respe tively,and
V
t
/h
istheturbulentvelo ity shear.When buoyan y for ing is stable, the boundary layer
depthistakenastheminimumofthevalue omputed
a - ordingtoEq.(1),theMonin-Obukovlengthandthe
Ek-mandepth.
The KPP model enables to take into a ount mixing
throughsaltngering(warmsaltywaterover oolfresh
wa-ter),atypi alpro essinregionswithlowpre ipitationand
intensesolarradiation.
2.3. E osystemmodel
Theboxmodel(Fig. 2) from Fashamet al.(1990)was designed to simulate annual y lesof plankton dynami s
and nitrogen y lingin theuppero eani layer.Nitrogen
isgenerally onsidered asthelimitingnutrientofprimary
produ tion,hen eits hoi eas entralelementofthe
bio-logi almodel.TheFashametal.(1990)modelhasbeen ap-pliedtonumerousareasoftheo ean(e.g.Sarmientoetal.,
1993;Drange,1996).TheversionimplementedinROMSis anevolutionofthismodel(Gruberetal.,2006),whi hhas alsobeenappliedtoanothertransitionzone(California).
It was hosenforits simpli ity, a ordingtothe
obje -tiveofgettingmeanseasonal y les.Moreover,themodelis
ompartmental,sothatitpermitsthedistin tionbetween
newprodu tion(drivenbynewnutrients,primarilynitrate,
thatoriginatesfromoutsidetheeuphoti zone)and
regen-eratedprodu tion(drivenbynitrogenre y ledwithinthe
euphoti zone,mainlyammonium)(DugdaleandGoering,
1967). Ba teriawere eliminated asan expli itlymodeled statevariable,andrepla edwithimpli itparametrizations
ofremineralizationpro esses(Gruberetal.,2006). Biologi al variables are oupled to physi s by an
adve tion-diusion-rea tion equation and through
tem-perature. The latter inuen es various produ tion terms
of the e osystemmodel. This again emphasizesthe need
ofa orre trepresentationoftemperatureseasonal y le.
Themodelisalsofor edbythephotosyntheti ally
avail-ableradiation(PAR),derivedfromtheshortwavesolar
ra-diation(Fig.3)a ordingtothefollowingsteps(Frentzel,
2006):
(i) A linearly interpolated value is al ulated for ea h
timestep.
(ii) Adiurnal y leforinsulationissuperimposedonthe
solar shortwave radiation. If the al ulated surfa e
radiationisnegative,zeroisusedinstead.
(iii) Startingfromthetop,the
P AR
is al ulatedforea h gridboxa ordingtoP AR
=
P AR
+1
exp {(−0.5 (k
w
+ k
Chla
[Chla]
) ∆z
}
(2)with
k
w
andk
Chla
, the attenuation oe ients for waterand hlorophyll(Tab.1),[Chla]
the hlorophyll on entration,∆z
,theheightoftheverti algridlayer and
the ellindex,from1atthebottomtoNatthe top.Aslightabundan eisanessentialfa torforprimary
pro-du tioninpelagi systems,ana urateparametrizationof
itsvariationwithdepth isrequired.Zielinskietal.(2002) testedseveral
P AR
parametrizationsandshowedthatthe lightmodelhassigni antee tsonthesimulateddistri-butionof hlorophyll,bothinspa eandtime.They
re om-mendusingamodelwithanin reasedvalueofthe
attenua-tion oe ientnearthesurfa e,duetoabsorptionatlarger
wavelengths,andwithaspe i hlorophyllattenuation.
Modelparametersfortheuxesbetween ompartments
(Tab.1)areeitherbasedonmeasuredvaluesintheregion,
orarestandardvalues.Whennoinformationwasavailable
fortheregionofinterest,modelpres ribedvalues,originally
adaptedtoCaliforniaregion,wereused.
Fig.2. Compartments and nitrogenuxes ofthe biologi al model
(reprodu edfromFrentzel(2006)).
3. Simulationssettings
3.1. For ing
TheROMS1Disfor edbyheatuxes,windstressand
freshwaterux.Theseelds wereextra tedfromdierent
databases (Tab.2) andmonthly averagedvaluesfor the
1993-2002 period were omputed at a lo ation south of
GranCanaria(
27.5
◦
N,
15.5
◦
W).The hoi eofthe10-year
averagedfor ingeldswasmotivatedbythedesireto
ob-tainameanseasonal y le.Shortertimes alepro essesare
Parameters ofthe biologi almodel:(1)Fasham etal.(1990), (2)Gruber et al.(2006), (3) Redeld(1934), (4) Zielinskiet al.(2002), (5)
Sarmientoetal.(1993),(6)Dadouetal.(2001);(
⋆
)modeldefaultvalues.Parameter Symbol Values Unit Sour e
Lightattenuationduetoseawater
k
w
0.04m
−1
(1,2)
Lightattenuationby hlorophylla
k
Chla
0.025m
2
(mg Chla)
−1
(2)
Initialslopeofthe
P − I
urveα
5.00mg C(mg Chla W m
−2
d)
−1
(
⋆
)C : N
ratioforphytoplanktonr
C:N ;phyto
6.625mM ol C(mM olN )
−1
(3)
Maximum ellular hlorophyllto
C
Ratioθ
m
0.053mg Chla : mg C
(2) Inversehalf-saturationforphytoplanktonN O
3
uptakeK
N O
3
1/0.51/(mM olN m
3
)
(4)
Inversehalf-saturationforphytoplankton
N H
4
uptakeK
N H
4
1/0.11/(mM olN m
3
)
(
⋆
)Phytoplanktonmortalitytosmalldetritusrate
t
P mort
0.07d
−1
(5)
Zooplankton-spe i maximumgrazingrate
t
Zgraze
0.75d
−1
(
⋆
)Zooplanktonassimilatione ien y
AE
0.75−−
(2,4,6)Zooplanktongrossgrowthe ien y
GGE
0.65−−
(⋆
)Zooplanktonhalf-saturation onstantforingestion
Z
P
1.00mM ol N m
3
(2)
Zooplanktonspe i ex retionrate
t
Zbmet
0.10d
−1
(
⋆
)Zooplanktonquadrati mortalitytodetritus
t
Zmort
0.10d
−1
(mM ol N m
3
)
−1
(2)
Smalldetritalbreakdownto
N H
4
ratet
SDremin
0.1d
−1
(
⋆
)Spe i (Per unit
P hyto
+SDet
)aggregationratet
coag
0.005(mM ol N m
3
)
−1
d
−1
(2)
Spe i rateoflargedetritusre y lingto
N H
4
t
LDremin
0.1d
−1
(
⋆
)Sinkingvelo ityforsmalldetritus
w
SD
0.1m d
−1
(
⋆
)Sinkingvelo ityforlargedetritus
w
LD
10.0m d
−1
(2,5)
Sinkingvelo ityforphytoplankton
w
P hyto
0.1m d
−1
(
⋆
)Sinkingvelo ityfor hlorophylla
w
Chla
0.1m d
−1
(
⋆
)Oxidationof
N H
4
toN O
3
(nitri ation)t
nitri
0.1d
−1
(
⋆
)Figure3showsthenetheatux obtainedfrom OAux
data base. Maximal values (around 150
W/m
2
) o ur
in July-August and minimal negative values o ur from
November to February, leadingto winter onve tion, the
mainfa torprodu ingmixedlayerdeepening.
Figure4illustratestheannual y leofwindstressfrom
NCEP reanalysis. Maximum intensities about 0.1
N/m
2
wererea hedduringsummermonthsduetoanin reasein
thestrengthoftheTradeWinds.AsexplainedinSe .5the
stabilizing ee t of amaximum positive netheat ux in
summerandthedestabilizingee t ofwind stressduring
thisperiodleadstothemaximalstrati ationinautumn.
Freshwater ux annual y le (ECMWF reanalysis) is
showninFig.5.Valuesrangefrom0.065
cm/day
in sum-merto 0.1cm/day
in winterandare weakin omparison withtemperateregions.Seasurfa etemperature(SST,Fig.5)isrequiredto
or-re ttheheatuxesbe auseofthesensitivityofthesurfa e
net heat ux to the
SST
(Haney, 1971; Killworth et al.,2000). The maximum is rea hed in September, after the TradeWindsweaken.
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
−150
−100
−50
0
50
100
150
200
250
300
Months
Heat fluxes (W/m
2
)
Net heat flux
Solar shortwave radiation
Fig.3.Mean y les(boldline)andindividualyearsfrom1993to2002
(thinlines)ofsurfa enetheatux(plainlines)andsolarshortwave
radiation(dashedlines)fromOAux.
3.2. Initialization
The model was initialized in Mid-August with in situ
on-Jan Feb Mar Apr May Jun
Jul
Aug Sep Oct Nov Dec
0
0.02
0.04
0.06
0.08
0.1
0.12
0.14
Months
Wind stress (N/m
2
)
Fig.4.SameasFig.3,butforwindstressfromNCEPReanalysis.
Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
0.04
0.06
0.08
0.1
0.12
0.14
Months
Precipitation minus evaporation (cm/day)
P−E
Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
18
19
20
21
22
23
24
25
Months
Sea Surface Temperature (
°
C)
SST
Fig.5.SeaSurfa eTemperaturefromNOAAOISSTV2and
fresh-wateruxfromECMWF40YearsRe-Analysis.
Table2
Sour esofreanalyzeddatausedformodelfor ing.
Data sour e
SST NOAA optimum interpolation SST V2 (Reynolds etal.,2002)
NHFLux Obje tively Analyzed Air-Sea Fluxes (OAFlux) for
theGlobal O eans(Yuet al.,2004;Yu andWeller,
2007;Zhangetal.,2004)
FWFlux ECMWF40YearsRe-Analysis(Uppalaetal.,2005)
WStress CDC Derived NCEP Reanalysis Produ ts Surfa e
Flux(Kanamitsuetal.,2002)
entrationsfromtheFAX ruise(Bartonetal.,2004). Ver-ti al proles were averaged within the region of interest
andttedwithfourth-orderpolynomials,in ordertohave
analyti alexpressionsforinputtothemodel.Higherorder
polynomialsweretested but resulted in largeos illations
thatwerenotphysi allyrealisti .Withthisaveraging,the
spatialvariabilityinherenttothezonesurveyedduringthe
FAX ampaignisremoved.
Salinity proles (Fig. 6) show the de rease of salinity
with respe t to depth below the mixed layer.This
nega-tivesalinitygradientis hara teristi ofsubtropi alregions,
whereevaporationandsolarradiationarehighthroughout
theyear(Figs.3and5).Thissituationisfavorableto
dou-blediusionthroughsalt-ngeringandthereforeenhan es
mixing(Pérezetal.,2001).
8
10
12
14
16
18
20
22
24
−500
−450
−400
−350
−300
−250
−200
−150
−100
−50
0
Temperature
Depth (m)
T (
°
C)
35.5
36
36.5
37
37.5
38
38.5
39
−500
−450
−400
−350
−300
−250
−200
−150
−100
−50
0
Salinity
Depth (m)
S (PSU)
Fig.6.Proles oftemperature and salinityusedfor model
initial-ization.Boldlinesrepresent meanvaluesobtainedfromindividual
FAXproles(thinlines).
Oxygen surfa e on entration is slightly lower than 5
µmol/m
3
andrea hesitsmaximumaround75m.Nitrate
hasverylow on entrations(
≤ 1 µmol/m
3
)intherst100
m,thenqui klyin reasestorea hmorethan
15 µmol/m
3
at500m.
Theotherphysi alandbiologi alvariablesareassigned
with onstantvaluesoverthewater olumn:
Theinitialvelo itiesweresettozero.
The initial value for the verti al mixing oe ient for
momentum(
K
M
)was0.01 m
2
/s
.
The verti al mixing oe ients for salinity (
K
S
) and temperature(K
T
)weretakenequalto0.001 m
2
/s
.
Theremainingbiologi alvariableswereinitializedwith
onstantvaluesrepresentativeoftheregion(Tab.3). In
the aseofzooplankton,measuredvalueswere onverted
into units ompatible with the model (
mM ol N
2
m
−3
)
using onversions fa torsof zooplankton biomass from
Harris et al. (2000). For ammonium, the mean value obtained from numeri al simulations (Bahamón and Cruzado,2003)isassignedastheinitial ondition.
Initialvaluesforthebiologi alvariables.(
⋆
)denotesdefaultvalues.Phytoplanktonwasinitializedusingthemaximal hlorophyll/phytoplanktonratio.
Variable Unit Pres ribedvalue Sour e
Ammonium
mM ol N
2
m
−3
0.1
BahamónandCruzado(2003)
Chlorophylla
mg Chl
a
m
−3
0.5
Arísteguietal.(1997) PhytoplanktonmM ol N
2
m
−3
0.1187
Redeld(1934) ZooplanktonmM ol N
2
m
−3
0.06
Hernández-Leónetal.(2007) SmalldetritusmM ol N
2
m
−3
0.04
(⋆
) LargedetritusmM ol N
2
m
−3
0.02
(⋆
)2
2.5
3
3.5
4
4.5
5
−500
−450
−400
−350
−300
−250
−200
−150
−100
−50
0
Oxygen concentration
Depth (m)
[O
2
]
10
12
14
16
18
20
22
24
26
28
−500
−450
−400
−350
−300
−250
−200
−150
−100
−50
0
Nitrate concentration
Depth (m)
[NO
3
]
Fig.7.SameasFig.6butforoxygenandnitrates.
3.3. Modelsetup
Thesimulationparametervalueswere hosentoprodu e
anannual y leandtosatisfythenumeri al onstraintsof
stability.Numerousrunsweremadetodeterminatesuitable
values.
Themodelwasrunfor6yearstosteadystate onditions,
withatimestep
∆t = 60s
.Resultswerestoredon eaday, whi hremovesshorttimes alevariability.The water olumn was limited to the rst 500 m and
dis retizedonastaggeredgrid.Thedepthofthe
i
th
layer isgivenbyd(i) = −h
sinh θ
N −i
N
sinh(θ)
,
i = {0, 1, . . . , N },
with
h
the totaldepth,N
thenumberof layersandθ
, a parameterfor the adjustment of the grid spa ing.Thenumberoflayers
N
was hosentobe40.Alargernumberoflayersin reasedthe omputationaltimewithoutgiving
signi antdieren esintheresults.
θ
wastakenequalto4 tohaveaneresolution(≈
1.5m)nearsurfa e.4. Modeltesting
Before performing simulations with the for ing and
initial onditionsdes ribedinse tions3.1and3.2,
respe -tively,theKPPmodelis omparedwithothermixedlayer
models using idealized experiments. These experiments
werenotdesignedtoberepresentativeofthe onditionsof
theregion of interest, but to provide ommon
ongura-tionsforevaluatingthemixingsubmodels.
4.1. Des riptionoftheexperiments
Inea h experiment,latitudewastakentobe
29.91
◦
N
,
whi hgivesaninertialperiod loseto24hours,andallows
omparisonswith similar resultsfrom other models. The
verti alresolution isuniformoverthewater olumnwith
∆z = 2 m
.Thevelo ityisinitializedwithzerovalueinea h dire tion.Theexperimentsaredes ribedhereinafter.(i) thedeepeningexperiment(Exp.I)wasdesignedto
ex-aminetheee tsofwindstressonthewater olumn
stability.It onsistsofawindwith onstantdire tion
andintensity(
τ = 0.4 N/m
2
)for5 days.Theinitial
temperaturewassetto
24
◦
C
atsurfa e,witha
ther-malstrati ationof
0.05
◦
C/m
.Thesurfa enetheat
uxwassettozero.
(ii) the oolingexperiment(Exp.II)showedmixedlayer
deepening due to onve tive mixing produ ed by a
negativeheat ux. Initial onditionswerethe same
asthepreviousexperiment;windstresswasassigned
a onstantvalue
τ = 0.1 N/m
2
andsurfa enetheat
ux wastaken to be
−96.8 W/m
2
. The experiment
wasrunfor120days.
(iii) the heating experiment (Exp.III) illustrated mixed
layershallowingwhen the o eanexperien es a
pos-itive net heat ux. The initial onditions were the
sameasthetwoprevious ases,ex eptthat
19
◦
C
.The wind stress wasequal to
τ = 0.1 N/m
2
andthenetheatuxwas
290.4 W/m
2
.
4.2. Des riptionofthemodels
TheKPPmodelis omparedwithveothermodels:the
Mellor-Yamada level2(MYL 2) andlevel 2.5(MYL 2.5)
losures(MellorandYamada,1974),theNiiler(Niilerand Kraus, 1977, Nii.), Garwood (Garwood, 1977,Gar.) and Pri e-Weller-Pinkel(Pri eetal.,1986,PWP)models.The rsttwomodelsbelongtothe ategoryofdierential
mod-els,i.e.governingequationsintheirprimitiveformareused,
whilethethreeothermodelsbelongtothe ategoryof
inte-gratedmodels,i.e.governingequationsareintegratedover
themixedlayerdepth.Resultsoftheexperimentsare
sum-marizedinTab.4.Forea hexperimenttheMLD omputed
withROMS1Dis ompatiblewiththevaluesobtainedwith
theothermodels.
Table4
Mixedlayerdepth(inmeters)indierentexperimentswith6models.
Models MYL2 MYL2.5Nii. Gar. PWP ROMS1D
Exp.I 39 39 54 51 54 42
Exp.II 101 103 127 117 101 110
Exp.III 14 14 4 14 8 7
5. Numeri alresults
5.1. Physi s
Annualeldsgeneratedwiththefor ingdes ribedin
se -tion 3.1 and with the parametersof se tion 3.3 are
pre-sented.
5.1.1. Temperature
Temperature(Fig.8)rea hedamaximumfromlate
Au-gustto mid-O toberwithvaluesaround
23
◦
C
. The
tem-peraturemaximumdoesnoto urwhentheheatuxwas
maximum (July-August), but rather in September, after
the period of peak insulation. This arises be ause of the
destabilizingee tofTradeWinds,whi hrea htheir
maxi-malintensityinJuly-August(Fig.4).Alongwiththis
max-imaltemperature,astrongstrati ationtakespla eunder
theinuen eofintensesolarheating.
FromFebruaryto May, strong onve tivemixing took
pla eandgeneratedminimumtemperaturesaround
18
◦
C
.
Thevariabilityindu edbyair-seauxesae tedonlythe
upper 125 m of the water olumn, sin e temperature is
fairly onstantbelowthisdepth.
5.1.2. Mixedlayerdepth
TheMLD istheupperpartofthewater olumn where
propertiesaredistributedquasihomogeneously.This
phys-i al property does not only a ountsfor the inuen e of
Months
Depth (m)
(
°
C)
13
14
15
16
17
18
19
19
20
22
23
21
Jan Feb Mar Apr MayJun Jul AugSep Oct NovDec
−450
−400
−350
−300
−250
−200
−150
−100
−50
14
16
18
20
22
Fig.8.AnnualtemperatureeldobtainedwithROMS1D.
ea hfor ingparameter(wind,surfa etemperature,
pre ip-itation,heatuxes),butalsostronglydetermines
bio hem-i al y les.
The annual evolution of mixed layer omputed with
ROMS 1D model (Fig. 9, plain line) follows the
temper-ature behavior: the mixed layer is deepest in February
(
> 100
m) after a period of onve tive mixing starting in November.Then shallowing startsasnet heat uxin- reasesandrea hesavaluearound25mbetweenJulyand
September,despiteintensesummerTradeWinds(Fig.4).
Deepening startsagain after September,under the ee t
ofde reasingsurfa enetheatux.
Jan
Fev
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
−180
−160
−140
−120
−100
−80
−60
−40
−20
0
Months
Mixed Layer Depth (m)
ROMS 1D −− KPP submodel
∆
T = 0.5
°
C from 10 m
Kara criterion with ROMS1D solution
Kara criterion with WOA05
Fig.9. Mixed layer depth: ROMS 1D KPP model (plain line),
sametemperature riterionasNeueretal.(2007)(dashedlinewith
),density riterionfromKaraet al.(2000)appliedto ROMS1D solution(dashedlinewith∇)
and toWOD05 limatology(dashed5.2. Biology
5.2.1. Phytoplankton
AsillustratedinFig.10,maximalphytoplankton
on en-trationsappearinFebruaryintherst100m.This
maxi-mumisexplainedbyanutrientinje tionintotheeuphoti
zoneprovokedbythethermo lineerosionindu edby
win-ter onve tivemixing.From mid-June,adeepmaximum,
alsovisibleinthe hlorophylleld(Fig.11),develops.
Beyond 150 m,phytoplankton on entrations are very
small (
O(10
−4
)
mMol
N
2
m
−3
). From asimple point of
view,phytoplanktonneeds two ingredientsto grow:light
and nutrients. Sin e light de reases exponentially with
depth, it be omes insu ient to sustain phytoplankton
growth, even though a su ient amount of nutrients is
available.
Months
Depth (m)
(mMol N
2
m
−3
)
0.05
0.1
0.15
0.2
0.25
0.25
0.25
0.25
0.3
0.3
0.35
Jan Feb Mar Apr MayJun Jul AugSep Oct NovDec
−150
−100
−50
0
0.05
0.1
0.15
0.2
0.25
0.3
Fig.10.Phytoplankton on entrationobtainedwithROMS1D.
5.2.2. Chlorophyll
Among thenumerous variablesof themodel, itis
par-ti ularlyinterestingtopresent hlorophyll on entrations,
sin ethisvariablewasinitializedwitha onstantvalue,
al-lowingthemodelto showits apabilityto reprodu ereal
prolesbyitself.
Figure 11 shows a homogeneous mixed layer of low
hlorophyll above a deep hlorophyll maximum (DCM)
developedinspringandsummer,withmaximal
on entra-tionsaround1mg
Chl
a
m
−3
at100m.Surfa e
on entra-tionsrangefrom 0.4mg
Chl
a
m
−3
,whenthemixed layer
isdeepest,to1mg
Chl
a
m
−3
insummer.
5.2.3. Summary
Inorderto summarizetheannualbiologi al y le,
on- entrationsbetweensurfa eandareferen edepth(the
eu-photi depth)areintegrated.Thelight-depthrelationship
usedinthemodelgivesanestimatedeuphoti depthof72.8
m.
Time [months]
Depth [m]
(mg Chl
a
m
−3
)
0.1
0.2
0.2
0.30.3
0.3
0.3
0.4
0.4
0.4
0.5
0.5
0.6
0.6
0.7
0.7
0.7
0.8
0.8 0.9
0.9
0.9
1
Jan Feb Mar Apr MayJun Jul AugSep Oct NovDec
−150
−100
−50
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
Fig.11. Chlorophyll on entration between surfa e and 150 m as
obtainedfromROMS1D.
Zielinski et al. (2002) ompared four water light-eld modelsandfoundarangeforthe1%depthbetween81and
139minwinter,andbetween83and133minspring.The
hoi e of therangefor theintegrationmodies
quantita-tivelytheresults,butthestudiedme hanismsarestillthe
same.
Thelate-winterbloomismadeupofthreestages:
(i) ApeakinnutrientsinearlyFebruary,dueto
onve -tivemixing.
(ii) Highphytoplankton on entrationsinmid-February
around25mMol
N
2
m
−3
.
(iii) AzooplanktonbloombeginningbytheendofMar h.
A ordingtothebiologi almodel,zooplankton
on- entrationis enhan edbygrazingofphytoplankton
and isdiminished bymetabolism, death and
ex re-tion.Itisthenassumedthattheoriginofthetimelag
andthelongerdurationofzooplanktonbloom omes
from the quantityof phytoplankton made available
duringtheshortbloomperiod.
Afterthesethreestages, on entrationsde reaseas
nu-trientsbe omeslessavailablewhilethethermo linestarts
toreform.Theevolutionofdetritus on entrationis
simi-lartothatforthezooplankton,withamaximumbetween
Mar handMayandaquasisteadyde reaseuntiltheend
ofDe ember.Thissimilarityarisesbe ausethemain
pro- essesthatdrivedetritusevolutionaremortalitiesof
phy-toplankton and zooplankton along with the part of
zoo-planktonthatisnotassimilated.
6. Validation
Resultsare omparedwiththefollowingsour es:
limatologi aleldsextra tedfromtheWorldO ean
At-las2005(Lo arninietal.,2006,WOA05herinafter)and averagedoverthe areaof interest.Using limatologi al
valuesis ompatiblewiththegeneralgoalofsimulating
Jan Fev Mar Apr May Jun Jul Aug Sep Oct Nov Dec
0
5
10
15
20
25
30
Months
Concentrations (mMol N
2
m
−2
)
NO
3
NH
4
Phyt
Zoo
SDet
Fig.12. Con entrations integrated fromeuphoti depthto surfa e
asobtainedfromROMS1D.
fortemperature,salinityfrom 0to1500mon24levels,
andforoxygenandnitratesfrom0to500mon14levels;
in situdata extra tedfrom theWorldO eanDatabase
andfrom ampaigns arriedaroundGranCanaria.
measurements from ESTOC station, keeping in mind
thatthe onditionsarenotexa tlythesame.
6.1. Temperatureeld
Figures8and13showthesamenear-surfa estru ture:
intense mixing and low temperatures during winter,
for-mationof thermo linein spring,strongstrati ation and
maximaltemperaturesinsummer.
Dieren es betweenthemodelresultsand WOA05
li-matology are attributed to the use of a one-dimensional
model,whi hpreventstheexisten eofverti alvelo ityand
adve tion.TheCanaryIslandsareinuen edby the ool
CanaryCurrent,whi h isnotin ludedin themodel
for -ing, whi h results in themodeled temperaturemaximum
beinghigherthanvaluesfromtheWOA05.
Thenumeri al resultsarealso in agreementwith
mea-surementsatESTOCstation:Neueretal.(2007): asurfa etemperatureof
18
◦
C
alongwithadeepmixing
fromJanuarytoMar h,
astrati ationstartinginJuneandamixedlayerdepth
around40-50m,
amaximaltemperatureduringSeptemberaround
24
◦
C
.
6.2. Mixedlayerdepth
There are several ways to ompute the MLD, as
illus-trated onFig.9.A ompletereview isfound in deBoyer Montgutet al.(2004).TheMLD omputedbyROMS1D isbasedonKPPmodel.Inthisformulation(Largeet al.,
1994,Eq.(21)),theMLD isthesmallestdepthvaluethat makes the bulk Ri hardson number equal to its riti al
Months
Depth (m)
12
13
14
15
16
17
18
19
19
20
21
22
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
−500
−400
−300
−200
−100
0
12
14
16
18
20
22
Fig.13.AnnualtemperatureeldfromWOA05MonthlyLongTerm
Meantemperature.
value
Ri
cr
. Hen eusingthisdenition,bothstrati ation andshearingare onsidered.Themain hara teristi softheMLDannual y le
al u-latedfromthemodel(plain urvewith ir les)are orre tly
reprodu ed:awinterdeepening ausedby onve tive
mix-ingandasummershallowingindu edbystrongheatux.
InNeueret al.(2007),awinterMLD varyingfrom 100 min1997to200min1994isshown.For omparisonswith
ESTOCstation, themodelMLDwas omputedfollowing
thesame riterion (temperature dieren e of
0.5
◦
C
with
respe ttothevalueat10m).Doingso,themixedlayeris
deepest(around120m)fromFebruarytoMar h(Fig.9).
A more sophisti ated method from Kara et al. (2000) isbasedonadensitydieren e riterionfrom areferen e
depth,hen eitpermitsthe onsiderationofvariable
salin-ity.ThismethodwasappliedonbothROMS1Dresultsand
WOA05 limatology oftemperatureand salinity,yielding
aunique riterionfor omparison.
Resultsare very similar from June to De ember, with
amaximal dieren e being lose to 20 m. In winter and
spring,dieren eslargerthan50m areen ountered.
Pos-sibleexplanationsarethela kofresolutionofthemodelat
thesedepthsandthemathemati aldenitionofMLD.
Nevertheless,thefour urvesrepresentthesamephysi al
pro esses,regardlessofthesele tedMLDformulation.
6.3. Nitrates
InFig.14 limatologi al(WOA05,Gar iaetal.,2006b) andinsitu(Braun,1976)dataare omparedwiththe an-nual y les of nitrates obtainedwith the model. In ea h
plot,ordersofmagnitudeare lose,suggestingthatthe
ou-plede osystemmodelprodu ed oherentresults.
Howevertwomaindieren esappear:1.theshapeofthe
iso-
N O
3
: in thenumeri al results,theyare lose to hori-zontal,whilein the limatology andin situ data,verti aldispla ements o ur. On e again, the absen e of verti al
verti algradientof
N O
3
islargerinthesimulationsthanin the limatology.Thismaybea onsequen eofthestret hedgrid(lowerresolutionasthedepth in reases)orofthe
in-trinsi e ien yofthemodel.
Near surfa e on entrations areverylow(maximumof
0.45mMol
N
2
m
−3
in therst75m), in agreementwith
Neueretal.(2007),whoreportedundete tablysmall mea-surablevalues(<1mMol
N
2
m
−3
)inthemixedlayer.
6.4. Phytoplanktonandzooplankton
Validation of plankton variables is not asdire t as for
nutrientsoroxygen,sin eno limatologiesexist forthese.
Moreover,allthe phytoplanktonandzooplanktonspe ies
are representedbytwoaggregatedstatevariables(
[phyto]
and[zoo]
)in themodel, whereasseveral groupsare usu-ally onsidered(e.g.pi o-,nano-phytoplankton,mesozoo-plankton, ...). Forthese reasons,their evolution will be
dis ussedqualitatively.
6.4.1. Phytoplankton
Arístegui et al. (2001) measured the biomass between Mar h1988andJune1989,withthe losestsampling(one
week)duringwinterbloom:maximaofintegrated
on en-trationo urinlatewinterandare omposedoftwopeaks
(see theirFig.3a), onein mid-Februaryandthe otherin
mid-Mar h.Theseobservationsare oherentwith the
nu-meri alresults(Figs.10and15),wheretwodistin tperiod
of high phytoplankton on entrationin therst 20-25m
arepresent.
Jan Fev Mar Apr May Jun Jul Aug Sep Oct Nov Dec
8
9
10
11
12
13
14
15
16
Months
Phytoplankton concentration (mMol N
2
m
−2
)
Fig.15.Integrated phytoplankton on entrationobtainedwiththe
modelinthe40rstm.
6.4.2. Zooplankton
InFig.12thezooplanktonbloomtakespla ewithatime
lagwithrespe ttothephytoplanktonbloom,inagreement
with observationsfrom Hernández-Leónet al.(1984)and
Arísteguiet al.(2001). However,with this ex eption,the
biologi almodelwasnotabletoreprodu ethetypi al
fea-turesofthezooplankton y le.
Hernández-Leónetal.(2004)measuredtheaverageand zooplanktonbiomasses(theirFigs.3and6)andshoweda
learmaximumby theend of February, while ROMS1D
resultsshowamaximumextendingoverseveraldaysinlate
Mar h.TheyalsoreportedamaximuminJuly,mentioned
inBraun(1976),andattributedittointenseTradeWinds duringthese months.Thisfeaturewasnotreprodu edby
themodel.
Asstatedpreviously,dieren esbetweenmodeland
re-alityoriginatefromtheuseofasinglevariabletorepresent
zooplankton. TheCanaryIslandwatersare hara terized
byaprodu tionmainlydrivenbymesozooplankton.
Adve -tionof biologi alproperties, throughupwelling laments
forexample,mayalso ontributetosu hdis repan ies.
6.5. Chlorophylla
Figure11showsresultsthatareinagreementwith
mea-surementsofNeuer et al. (2007),whoshowed on entra-tionshigherthan0.5mg
Chl
a
m
−3
neartheeuphoti depth
andBartonet al.(2000)whoreportedvalueshigherthan 0.6mg
Chl
a
m
−3
intheleeregionofGranCanaria.
A ord-ingtoTett etal.(2002),thisDCMresultsfroma steady-statebalan ebetweenthelimitationofphytoplankton
pro-du tionduetolightandlossesduetoexportedprodu tion
andlo almetabolismofmi roplankton.
Simulatedsurfa e on entrationsagree with the
obser-vationsofNeueretal.(2007):highestvalues(upto0.4mg
Chl
a
m
−3
)whenthemixedlayerisdeepestandlowervalues around0.05mgChl
a
m
−3
(1mg
Chl
a
m
−3
insimulations)
insummer.Arísteguietal.(2001)reportedasmall hloro-phyllbloom,withvalueslargerthantheannualmean,but
lowerthanthoseen ounteredintemperateregions.
ComparisonwithinsitudatafromtheFAX ruise( Bar-tonetal.,1998)ismadeinFig.16.Variationsfromoneday toanother annotbe apturedbyone-dimensionalmodel,
astheymay omefromthe omplex ir ulationpatternof
the region. However, the model shows distributions that
agreeinstru tureandmagnitude.
ComparisonswithZielinskietal.(2002)showgood agree-ment.Thesimulatedand observedDCMappearsat
sim-ilardepths. Moreover,ROMS1Dresults ompare
quanti-tativelybettertotheESTOCobservedvaluesthandothe
simulatedresults presented in Zielinski et al. (2002)(see theirFig.10).
6.6. Oxygen
Thesimulatedoxygenannual y le(Fig.17(a))is
om-paredwith limatologi alelds(Gar iaetal.(2006a),Fig. 17(b)).Bothshowsimilarordersofmagnitudeandahigh
near-surfa e on entrationsabove5mMol
O
2
m
−3
at the
endofwinter.Anothersurfa estru turewithoxygen
on- entrationsaround5mMol
O
2
m
−3
Months
Depth (meters)
(mMol N
2
m
−3
)
1
1
2
2
3
4
5
6
7
8
9
10
11
12
12
13
13
14
14
15
15
16
17
16
18
18
18
19
19
20
20
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
−500
−400
−300
−200
−100
0
2
4
6
8
10
12
14
16
18
(a)WOA2005 (b)Insitu
Months
Depth (m)
(mMol N
2
m
−3
)
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Jan Feb Mar Apr MayJun Jul AugSep Oct NovDec
−450
−400
−350
−300
−250
−200
−150
−100
−50
2
4
6
8
10
12
( )ROMS1DFig.14. Annual eldof nitrate on entration: (a)WorldO ean Atlas 2005,(b)measurements 5mileseasto Tenerife(reprodu ed from
Braun(1976)),( )modelresults.
0
0.5
1
1.5
2
−150
−100
−50
0
(mg Chl
a
m
−3
)
Depth (m)
FAX data
Simulation
Fig.16.Modelresults(boldlines)andinsitumeasurements(Barton etal.,1998)forthe hlorophyll on entrationbetween7thand9th ofAugust.
Thedieren es seenbetweenthesimulated andobserved
nitrate valuesmodel arealso visible in theoxygen
distri-butions.
7. Dis ussion
More than 50 years ago, Sverdrup (1953) developed a modelto explain thespring bloomof phytoplankton and
the onditionsunderlyingitsformation.Histheorystates
that the phytoplankton growth season starts in early
spring, when the mixed layer depth be omes shallower
thanthe riti aldepth. Criti aldepthdepth isdenedas
the depth above whi h the depth-integrated daily gross
primaryprodu tionequalsrespiration,i.e.thedepthabove
whi hintegratednetdailyprimaryprodu tionequalszero
(Nybakken,2001).
Theassumptionsunderlyingthistheoryare:
(i) phytoplankton ellsareuniformlydistributed inthe
mixedlayer,
(ii) photosynthesisisassumedtode reaseexponentially
withdepth,asthelightdoes,
Months
Depth (m)
(mMol O
2
m
−3
)
3.8
4
4.2
4.4
4.6
4.8
5
5
5.2
Jan Feb Mar Apr MayJun Jul AugSep Oct NovDec
−500
−400
−300
−200
−100
0
3.8
4
4.2
4.4
4.6
4.8
5
(a)ROMS1DMonths
Depth (m)
[mMol O
2
m
−3
]
3.2
3.4
3.6
3.6
3.8
3.8
4
4.2
4.4
4.6
4.8
4.8
5
5
5.2
5.2
5.2
5.2
5.4
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
−500
−400
−300
−200
−100
0
3.5
4
4.5
5
(b)WOA2005Fig.17.Annualeldofoxygen on entration:(a)Resultsfrom
mixedlayer.
The riti aldepthwas al ulatedusingtheaveragevalue
oflight
I
¯
D
penetratingintotheeuphoti zone. This aver-ageamountoflightisobtainedbyintegratingtherelationbetweendepthandlightintensity
I
λ
(z) = I
λ,0
exp(−k
λ
z)
(3) fromthesurfa e(z = 0
)tothedepthD
,whi hleadsto¯
I
D
=
¯
I
0
kD
(1 − e
−kD
).
(4)The previousequation is used to getan expression for
the riti aldepth
D
cr
.Assumingthatphytoplankton ells moveupanddowninthemixedlayer,theyre eiveonav-erage alight intensity equal to
I
¯
D
. The riti al depth is thenthedepthatwhi hI
¯
D
equalstoI
C
,the ompensation lightintensity, dened asthe amountof lightthat makesprodu tionandlossequal.The ompensationdepthisthe
depthatwhi hthatequalityissatised.
CombiningthepreviousdenitionwithEq.(3),the
om-pensationdepthis al ulatedby
D
c
=
ln(I
0
) − ln(I
C
)
k
,
(5)Finally,thesubstitutionof
I
¯
D
bythe ompensationlight intensityI
C
in(4)yieldsD
cr.
=
¯
I
0
k I
C
[1 − exp(−k D
cr.
)]
(6) The ompensationdepthdependsonthe larityofwaterandthusvariesovertheworldo eans.However,a ording
toNybakken(2001),theeuphoti depthisagoodestimate. Solving Eq. (6)with this approximation, a riti aldepth
equal to 113 m wasfound. As the MLD depends on the
wayitisdened(seeFig.9),interpretationofSverdrup's
riti aldepththeoryisnotstraightforward.
TheMLD omputedfromtheROMS1Dsimulation
re-sults results in a riti al depth that is deeper than the
MLDallyearlong,whi histypi alforsubtropi alregions
(Arísteguietal.,2001).However,ifothermethodsareused to omputetheMLD,thereisaninterse tionbetween
rit-i al and mixed layer depths, taking pla e during April,
whi hdisagreeswiththeobservationofalate-winter
phy-toplanktonbloom.
Hen eitis on ludedthatthemixedlayerdepthisa
on-trolonthetimingofthebiologi al y les,evenifSverdrup's
theory ouldnotbeappliedintheregionofinterest.
8. Con lusions
8.1. Physi s
Physi alpro essessouthofGranCanariawere
su ess-fullysimulatedusingtheROMD1Dmodel.Maximal
sur-fa e temperatures and strongest strati ation were
pro-du ed in late summerearly fall, when the Trade Winds
100mo ursinFebruary,undertheee tofnegativeheat
ux,inagreementwithobservationsandwithexisting
li-matologies.The orre trepresentationofthisphysi al
fea-tureisarststepindevelopmentofamodeltounderstand
thebiogeo hemi alpro esses.
Thedieren esbetweennumeri alresultsand data(in
situor limatologi al)arebelievedtobeduetothe
follow-ing:
Thenatureofa1-Dmodel:neitheradve tionnorverti al
velo ityexists.
For ing is arried out with monthly valueslinearly
in-terpolatedatea htimestep,thuspro esseswithshorter
times alearelteredout.
However,evenwiththese dieren esmeanseasonal
y- lesweresimulated.
8.2. Biology
Astrong ouplingbetweenair-seauxesand biologi al
y leswasobserved:
the phytoplankton bloom o urs in late winter, when
surfa etemperature is minimumbe ause of onve tive
mixingprodu edbynegativeheatux.
Fromearly springto latefall, biologi al on entrations
tendtode reaseundertheee toftheshallowseasonal
thermo lineformedbyintenseheating.
This y le ontrasts with what is usually observed in
temperateregions,whereaphytoplanktonbloomo ursin
early spring,strongest mixing in winter,due to maximal
windvelo ities duringthisseason,andmaximal
tempera-turesinsummer.
Primary produ tion in the Canary Island region is
nutrient-limited, as there is a su ient amount of light
re eivedbythe water olumn allyearlong,while in
tem-perateregions,primaryprodu tionislightlimited,asthe
mixing is strong enough to feed the euphoti layer with
newnutrientsduringmostoftheyear.
Numerousbiologi almodelshavebeendes ribedin the
literatureandsomewereprimarilydesignedtoworkwithin
a pre-determinate domain of the sea. As a model is
al-waysan idealizationofreality,ithasto bea eptedthat
some pro esses may not be a urately modeled or well
parametrized.Nevertheless,thee osystemmodel oupled
withthephysi almodelyieldedresultsthat orrespondto
observedpatternsformostofthesimulatedvariables.
8.3. Perspe tives
Furtherworkshould on entrateonthe omputationof
the riti aldepthwithamoredetailed model, totest the
appli abilityofSverdup'stheorytotheCanaryIsland
wa-ters. Some of the parameters in the biologi al model
re-quiremodi ationsthatshouldbebasedonmeasurements
and/or experiments arried out in the Canary Island
sim- omponentintoseveral ompartments.
9. A knowledgments
Theauthorsareverygratefultotwoanonymous
review-ersfor their onstru tive ommentswhi h ontributesto
deeplyimprovetheoriginalmanus ript,toP.Penven(IRD)
formakingavailabletheROMS1Dmodelandforhelping
foritsuse, E.Mason(ULPGC)forhis orre tionsonthe
manus ript and S. Hernández-León for his omments on
thezooplankton y le.
NOAA-ESRLPhysi al S ien es Division,theECMWF
and the WHOI are a knowledged for providing the data
setsusedinthefor ing,andtheNODCformakingavailable
theWorldO eanAtlas.
FundingbyFondsde la Re her he S ientique- FNRS
throughaFRIA Grantandby theMECthroughRODA
Proje t(CMT2004-06842-CO3/MAR)are greatly
appre- iated. This resear h was initialized during a stay of C.
TroupinattheULPGCfa ilitatedbyanERASMUSgrant
fromtheEuropeanCommission.
ThisisMAREpubli ation172.
Referen es
Arístegui,J.,Hernández-León,S.,Montero,M.F.,Gómez,
M.,2001.Theseasonalplanktoni y lein oastalwaters
oftheCanaryIslands.S i.Mar.65,5158.
Arístegui,J.,Montero,M.F.,2005.Temporalandspatial
hangesin planktonrespirationandbiomassinthe
Ca-naryIslandsregion:theee tofmesos alevariability.J.
Mar.Syst.54,6582.
Arístegui, J., Sangrà, P., Hernández-León, S., Cantón,
M.,Hernández-Guerra,A.,Kerling,J.L.,1994.
Island-indu ededdiesintheCanaryIslands.Deep-SeaRes.41,
15091525.
Arístegui,J.,Tett,P.,Hernández-Guerra,A.,Basterretxea,
G., Montero, M. F., Wild, K., Sangrà, P.,
Hernández-León, S., Cantón, M., Gar ía-Braun, J., Pa he o, M.,
Barton, E. D., 1997. The inuen e of island-generated
eddieson hlorophylldistribution:astudy ofmesos ale
variationaroundGranCanaria.Deep-SeaRes.44,7196.
Bahamón,N.,Cruzado,A.,2003.Modellingnitrogenuxes
in oligotrophi environments: NW Mediterranean and
NEAtlanti .E ologi alModelling163,223244.
Barton,E.D.,Arístegui,J.,Tett,P.,Cantón,M.,
Gar ía-Braun, J., Hernández-León, S., Nykjaer, L., Almeida,
C.,Almunia,J.,Ballesteros,S.,etal.,1998.The
transi-tionzoneoftheCanaryCurrentupwellingregion.Prog.
O eanogr.41,455504.
Barton,E. D., Arístegui,J., Tett, P., Pérez, E. N., 2004.
Variabilityin the CanaryIslandsareaoflament-eddy
ex hanges.Prog.O eanogr.62,7194.
Ja ob,E.G.,Jones,B.,Arístegui,J.,Felix,H.,2000.Lee
region of Gran Canaria.J. Geophys. Res. 105,17173
17193.
Barton,E. D.,Flament,P., Dodds,H., Mit helson-Ja ob,
E. G., 2001. Mesos alestru tures viewed by SAR and
AVHRRneartheCanaryIslands.S i.Mar.65,167175.
Braun,J.G.,1976.Produ tionStudies inCanaryIslands
waters.I.Hydrography,nutrientsand primary
produ -tion.
Bro hier,T.,Mason,E.,Sangrà,P.,Lett,C.,2008.
I hthy-oplanktontransportfromtheafri an oasttothe anary
islands: a ase study using a high-resolution
hydrody-nami .In:S ien eandtheChallengeofManagingSmall
Pelagi FisheriesonSharedSto ksinNorthwestAfri a.
Casablan a,Morro o.
Dadou,I.,Lamy,F.,Rabouille,C.,Ruiz-Pino,D.,
Ander-sen,V.,Bian hi,M.,Garon,V., 2001.Anintegrated
biologi al pump model from the euphoti zone to the
sediment:a1-Dappli ationintheNortheasttropi al
At-lanti .Deep-SeaRes.48,23452381.
Davenport, R., Neuer, S.,Helmke, P., Pérez-Marrero,J.,
Llinás,O., 2002.Primary produ tivity in the northern
CanaryIslandsregionasinferredfromSeaWiFSimagery.
Deep-SeaRes.49,34813496.
de BoyerMontgut, C., Made , G., Fis her, A. S.,Lazar,
A.,Iudi one,D.,2004.Mixedlayerdepthovertheglobal
o ean:Anexaminationofproledataandaprol-based
limatology.J.Geophys.Res.109,C12003.
DeLeón,A.R.,Braun,J.G.,1973.Ci loanualdela
pro-du iónprimariaysurela ión onlosnutrientesenaguas
anarias.Bol.Inst.Esp.O ean.167,124.
Drange, H., 1996. A 3-dimensional isopy ni oordinate
modeloftheseasonal y lingof arbonandnitrogenin
theAtlanti O ean.Phys.Chem.Earth21,503509.
Dugdale,R.C.,Goering,J.J.,1967.Uptakeofnewand
re-generatedformsofnitrogeninprimaryprodu tion.
Lim-nol.O eanogr.12,196206.
Fasham,M.J.R.,Du klow,H.W.,M Kelvie,S.M.,1990.
A nitrogen-based model of plankton dynami s in the
o eani mixedlayer.J.Geophys.Res.48,591639.
Frentzel, H., 2006. Thebiogeo hemi almodel of the
Re-gional O eanModelingSystem (ROMS). Available
on-linehttp://www.brest.ird.fr/Roms_tools.
Gar ia, H. E., Lo arnini, R. A., Boyer, T. P., Antonov,
J. I., 2006a. World O ean Atlas 2005, Volume 3:
Dis-solvedoxygen,apparentoxygenutilization,andoxygen
saturation.Te h.rep.,NOAA,WashingtonD.C.,342pp.
Gar ia,H.E.,Lo arnini,R.A.,Boyer,T.P.,Antonov,J.I.,
2006b. World O ean Atlas 2005, Volume 4: Nutrients
(phosphate,nitrate,sili ate).396pp.
Garwood,R.W.,1977.Ano eani mixedlayermodel
a-pableof simulating y li states. J.Phys.O eanogr. 7,
455471.
Gruber, N., Frenzel, H., Doney, S. C., Mar hesiello, P.,
M Williams,J.C.,Moisan,J.R.,Oram,J.J.,Plattner,
simu- urrentsystem.Deep-SeaResear h53,14831516.
Haney,R. L.,1971.Surfa ethermal boundary onditions
foro ean ir ulationmodels.J.Phys.O eanogr.1,241
248.
Harris, R., Wiebe, P., Lenz, J., Skjoldal,H.-R., Huntley,
M. E., 2000. Zooplankton Methodology Manual.
A a-demi Press,London,UK.
Hernández-León,S.,Almeida,C.,Bé ognée,P.,Yebra,L.,
Arístegui,J.,2004.Zooplanktonbiomassandindi esof
grazingand metabolism during alate winter bloomin
subtropi alwaters.Mar.Biol.145,11911200.
Hernández-León,S.,Gómez,M.,Arístegui,J.,2007.
Meso-zooplanktonintheCanaryCurrentSystem:The
oastal-o eantransitionzone.Prog.O eanogr.74,397421.
Hernández-León,S.,Llinás,O., Braun,J.G.,1984. Ci lo
anualdelabiomasadelmesozooplan tonsobreunarea
de plataforma en aguas del Ar hipielago Canario. Inv.
Pes .52,316.
Hernández-León,S.,Llinás,O.,Braun,J. G.,1984. Nota
sobrelavaria ióndelabiomasadelmesozooplan tonen
aguasdeCanarias.Inv.Pes .48,495508.
Ja kett, D. R., M Dougall, T. J., 1995. Minimal
adjust-mentofhydrostati prolestoa hievestati stability.J.
Atmos.O ean.Te h.12,381389.
Kanamitsu, M.,Ebisuzaki, W., Woollen, J.,Yang,S.-K.,
Hnilo,J.J.,Fiorino,M.,Potter,G.L.,2002.NCEP-DOE
AMIP-IIReanalysis(R-2).Bul.oftheAtmos.Met.So .
83,16311643.
Kara,A.B.,Ro hford,P.A.,Hurlburt,H.E.,2000.An
op-timaldenitionforo eanmixedlayerdepth.J.Geophys.
Res.105,1680316821.
Killworth,P.D.,Smeed,D.A.,Nurser,A.J.G.,2000.The
ee tsono eanmodelsofrelaxationtowardobservations
atthesurfa e.J.Phys.O eanogr.30,160174.
Large, W. G., M Williams, J. C., Doney, S. C., 1994.
O eani verti almixing:areviewandamodelwitha
non-lo al boundary layer parameterization. Rev. Geophys.
32,363403.
Lo arnini, R.A., Mishonov,A.V., Antonov,J.I.,Boyer,
T. P., Gar ia, H. E., 2006. World O ean Atlas 2005,
Volume1:Temperature.Te h.rep.,NOAA,Washington
D.C.,182pp.
Mason,E.,Colas,F.,Molemaker,J.,Sh hepetkin,A.,
San-grà, P., M Williams, J. C., 2008a. Seasonal variability
ofthe anary urrentsystem.In:EasternBoundary
Up-wellingE osystems.LasPalmasdeGranCanaria,Spain.
Mason, E., Sangrà,P., Colas,F., Molemaker,J.,
Sh hep-etkin, A., Hughes, M., Dong, C., M Williams, J. C.,
2008b.Ahigh resolutionnumeri almodel study atthe
Canary islands. In: O ean S ien e Meeting. Orlando,
USA.
Mellor,G.L.,Yamada,T.,1974.Ahierar hyofturbulen e
losuremodelsforplanetaryboundarylayers.J.Atmos.
S i.31,17911806.
Neuer,S.,Cian a,A.,Helmke,P.,Freudenthal,T.,
Daven-port,R.,Meggers,H.,Knoll,M.,Santana-Casiano,J.M.,
hemistry and hydrography in the eastern subtropi al
NorthAtlanti gyre. Results from the European
time-seriesstationESTOC.Prog.O eanogr.72,129.
Niiler,P.P.,Kraus,E.B.,1977.ModellingandPredi tion
of the Upper Layers of the O ean. Oxford: Pergamon
Press,NewYork.
Nybakken,J.W.,2001.MarineBiology,AnE ologi al
Ap-proa h.BenjaminCummings,SanFran is o.
Pelegrí,J.L.,Arístegui,J.,Cana,L.,González-Dávila,M.,
Hernández-Guerra, A., Hernández-León, S.,
Marrero-Díaz,A.,Montero,M.,Sangrà,P.,Santana-Casiano,M.,
2005a.Couplingbetweentheopeno eanandthe oastal
upwellingregiononorthwestAfri a:waterre ir ulation
and oshorepumping of organi matter. J. Mar. Syst.
54,337.
Pelegrí, J. L.,Marrero-Díaz, A., Ratsimandresy, A.,
An-toranz,A.,Cisneros-Aguirre,J.,Gordo,C.,Grisola,D.,
Hernández-Guerra,A., Laíz, I., Martínez, A., Parrilla,
G.,Pérez-Rodríguez,P.,Rodríguez-Santana,A.,Sangrà,
P.,2005b.Hydrographi ruisesonorthwestAfri a:the
Canary urrentandtheCapeGhirregion.J.Mar.Syst.
54,3963.
Penven, P., 2006. 1D ROMS model
do umentation. Available online
http://www.brest.ird.fr/Roms_tools/roms1d. Pérez,F.F.,Mintrop,L.,Llinás,O.,González-Dávila,M.,
Castro, C. G., Alvarez, M., Körtzinger, A.,
Santana-Casiano, M., Rueda, M. J., Ríos, A. F., 2001. Mixing
analysisofnutrients,oxygenandinorgani arboninthe
CanaryIslandsregion.J.Mar.Syst.28,183201.
Pri e,J.F.,Weller,R.F.,Pinkel,R.,1986.Diurnal y ling:
Observationsandmodelsoftheuppero eanresponseto
diurnalheating, oolingand windmixing. J.Geophys.
Res.91,84118427.
Redeld,A.C.,1934.Ontheproportionsoforgani
deriva-tionsin seawaterandtheirrelationtothe omposition
ofplankton.In:Daniel,R.(Ed.),JamesJohnson
Memo-rialVolume.UniversityPressofLiverpool,pp.177192.
Reynolds, R. W., Rayner, N. A., Smith, T. M., Stokes,
D.C.,Wang,W.,2002.Animprovedinsituandsatellite
SSTanalysisfor limate.J.Clim.15,16091625.
Sarmiento,J.L.,Slater,R.D.,Fasham,M.J.R.,Du klow,
H.W.,Toggweiler,J.R.,Evans,G.T.,1993.Aseasonal
three-dimensionale osystemmodel of nitrogen y ling
intheNorthAtlanti euphoti zone.GlobalBiogeo hem.
Cy les7,417450.
Sh hepetkin,A.F.,M Williams,J.C.,2005.Theregional
o eani modelingsystem(ROMS):asplit-expli it,
free-surfa e,topography-following- oordinateo eani model.
O eanModell.9,347404.
Sverdrup,H.U.,1953.On onditionsforthevernal
bloom-ingofphytoplankton.J.Cons.Perm.Int.Explor.Mer.
18,287295.
Tett,P.,Arístegui,J.,Barton,D.,Basterretxea,G.,Armas,
J.D. D., Es ánez,J. E., Hernández-León,S.,Lorenzo,
Uppala, S.M., Kallberg,P. W.,Simmons, A. J.,Andrae,
U., daCosta Be htold, V., Fiorino, M., Gibson, J. K.,
Haseler, J., Hernandez, A., Kelly, G. A., et al., 2005.
TheERA-40re-analysis.Quart.J.R.Meteorol.So .131,
29613012.
Yu, L.,Weller, R. A., 2007. Obje tivelyanalyzed air-sea
heat uxes for the global i e-free o eans (1981-2005).
Bull.Am.Meteorol.So .88,527539.
Yu,L.,Weller,R.A.,Sun,B.,2004.Improvinglatentand
sensibleheatuxestimatesfortheAtlanti O ean
(1988-1999)byasynthesisapproa h.J.Clim.17,373393.
Zhang, Y.-C., Rossow, W. B., La is, A. A., Oinas, V.,
Mish henko,M.I.,2004.Cal ulationofradiativeuxes
from thesurfa etotopofatmospherebasedonISCCP
andotherglobaldatasets:Renementsoftheradiative
transfermodelandtheinputdata.J.Geophys.Res.109,
D19105.
Zielinski,O.,Llinás,O.,Os hlies,A.,Reuter,R.,2002.
Un-derwater lighteld and itsee t on aone-dimensional
e osystemmodelatstationESTOC,northoftheCanary