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Ocean Color Response to an Episode of Heavy Rainfall In the Lagoon of New Caledonia Cécile Dupouy * a,e , Robert Frouin b , Rüdiger Röttgers c , Jacques Neveux d , , Francis Gallois a , Jean-Yves Panché a , Philippe Gerard a , Clément Fontana e , Christel Pinazo e , Sylvain Ouillon f , Audrey Minghelli-Roman g a Centre IRD de Nouméa, BP A5, 98848, Nouméa, New Caledonia b Scripps Institution of Oceanography, La Jolla, California, USA c GKSS, Geesthacht, Hambourg, Germany d UPMC (Paris 6), CNRS, LOBB, 66651 Banyuls sur Mer, France e LOBP, Université de la Méditerranée, 13007, Marseille cedex 09, France f IRD, Université de Toulouse, 31400, France g LSEET UMR CNRS 6017, 83162, La Seyne sur Mer, France ABSTRACT Inherent optical properties (IOPs) and remote sensing reflectance were measured in the southern part of the lagoon of New Caledonia during the VALHYBIO cruise (March-April 2008). The goal was to validate satellite chlorophyll data from MODIS and MERIS and to validate simulations of surface chlorophyll by a biogeochemical model. Physical parameters were collected from a Seabird CTD. Particulate and detritus absorption were measured with the filter pad technique. Backscattering was measured with a Hydroscat-6. Mapping of IOPs and Rrs were done for the whole southern lagoon area and compared with pigment maps. The cruise provided a description of the IOPs in different water types including bays, open ocean waters, mid-shelf lagoon, and above reefs. With respect to climatology, the heavy rainfall episode of March-April 2008 resulted in a large increase in chlorophyll-a concentration (by a factor of 3) attributed to increased nutrient availability from land drainage. Low backscattering ratios characterized the chlorophyll-rich plumes associated with the nutrient increase. The data are useful for the development of a specific algorithm for chlorophyll concentration retrieval by satellite in all oligotrophic lagoons during dry and wet seasons. Keywords: chlorophyll, algorithm, coral reef, lagoon, ocean color, New Caledonia, tropical Pacific ocean, La Nina, sea surface reflectance 1. INTRODUCTION Coral reef lagoon systems are very sensitive to anthropogenic (nutrients, mining) perturbations [1] as well as to interannual changes linked to the balance between dry El Nino and wet La Nina episodes, which are amplified in lagoons [2]. Sea surface chlorophyll is a proxy of phytoplankton biomass and is a direct integrator for the nutrient status of water masses and chlorophyll monitoring by satellite will greatly expand our knowledge of the functioning of coral reef lagoons [3]. Lately, it would allow the validation of simulations of chlorophyll by recently developed coupled biogeochemical models [4]. Tropical coastal environments are characterized by a range of extremely oligo- to eutrophic waters [5-8]. Lagoon waters belong to the class of optically complex waters (classified as Case 2 waters) where mineral particles and colored dissolved organic matter mix with phytoplankton [9]. Indeed, current algorithms such as OC4v4 for SeaWiFS and OC3 for MODIS [10, 11] are suitable for oceanic waters where chlorophyll drives variability of bio-optical properties (absorption and backscattering) of the waters. Attempts have been made to retrieve chlorophyll from remote sensing data in turbid case 2 waters [11-15]. Other algorithms tend to minimize the effect of bottom reflectance which increase surface reflectance values and therefore cause chlorophyll concentrations overestimation using algorithms developed for optically deep data [16]. The New Caledonian lagoon (22 177 km 2 , 25 m as a mean depth) lies in the South Western Tropical Pacific from 20°S to 22°S, and 166° to 167°E, with a heterogeneous bathymetry due to a complex geomorphology and a variety of different bottom colors. It is largely connected to the open ocean in the south part of the lagoon, but only by narrow passes in the south west part of the lagoon. Exchanges with the sea can modify the phytoplanktonic assemblage in the central lagoon characterized by oligotrophic to mesotrophic waters (yearly average chlorophyll-a concentration of 0.25 ± 0.01 mg m -3 ) [17, 18]. With relatively low river inputs and a low turbidity range compared with other tropical lagoons (0.20-16 g m -3 , [8], its trophic state is linked to spatial variations in flushing times [19, 20]. Ocean Remote Sensing: Methods and Applications, edited by Robert J. Frouin, Proc. of SPIE Vol. 7459, 74590G · © 2009 SPIE · CCC code: 0277-786X/09/$18 · doi: 10.1117/12.829251 Proc. of SPIE Vol. 7459 74590G-1 DownloadedFrom:http://proceedings.spiedigitallibrary.org/on09/13/2013TermsofUse:http://spiedl.org/terms

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Page 1: Ocean Color Response to an Episode of Heavy Rainfall In ...genius.ucsd.edu/Public/Reviews/B77_74590G.pdfOcean Color Response to an Episode of Heavy Rainfall In the Lagoon of New Caledonia

Ocean Color Response to an Episode of Heavy Rainfall

In the Lagoon of New Caledonia

Cécile Dupouy *a,e

, Robert Frouinb, Rüdiger Röttgers

c, Jacques Neveux

d, , Francis Gallois

a,

Jean-Yves Panchéa, Philippe Gerard

a, Clément Fontana

e, Christel Pinazo

e ,

Sylvain Ouillonf, Audrey Minghelli-Roman

g

a Centre IRD de Nouméa, BP A5, 98848, Nouméa, New Caledonia

b Scripps Institution of Oceanography, La Jolla, California, USA

c GKSS, Geesthacht, Hambourg, Germany

d UPMC (Paris 6), CNRS, LOBB, 66651 Banyuls sur Mer, France

e LOBP, Université de la Méditerranée, 13007, Marseille cedex 09, France

f IRD, Université de Toulouse, 31400, France

g LSEET UMR CNRS 6017, 83162, La Seyne sur Mer, France

ABSTRACT

Inherent optical properties (IOPs) and remote sensing reflectance were measured in the southern part of the lagoon of New

Caledonia during the VALHYBIO cruise (March-April 2008). The goal was to validate satellite chlorophyll data from

MODIS and MERIS and to validate simulations of surface chlorophyll by a biogeochemical model. Physical parameters were

collected from a Seabird CTD. Particulate and detritus absorption were measured with the filter pad technique.

Backscattering was measured with a Hydroscat-6. Mapping of IOPs and Rrs were done for the whole southern lagoon area

and compared with pigment maps. The cruise provided a description of the IOPs in different water types including bays, open

ocean waters, mid-shelf lagoon, and above reefs. With respect to climatology, the heavy rainfall episode of March-April 2008

resulted in a large increase in chlorophyll-a concentration (by a factor of 3) attributed to increased nutrient availability from

land drainage. Low backscattering ratios characterized the chlorophyll-rich plumes associated with the nutrient increase. The

data are useful for the development of a specific algorithm for chlorophyll concentration retrieval by satellite in all

oligotrophic lagoons during dry and wet seasons.

Keywords: chlorophyll, algorithm, coral reef, lagoon, ocean color, New Caledonia, tropical Pacific ocean, La Nina, sea

surface reflectance

1. INTRODUCTION

Coral reef lagoon systems are very sensitive to anthropogenic (nutrients, mining) perturbations [1] as well as to interannual

changes linked to the balance between dry El Nino and wet La Nina episodes, which are amplified in lagoons [2]. Sea surface

chlorophyll is a proxy of phytoplankton biomass and is a direct integrator for the nutrient status of water masses and

chlorophyll monitoring by satellite will greatly expand our knowledge of the functioning of coral reef lagoons [3]. Lately, it

would allow the validation of simulations of chlorophyll by recently developed coupled biogeochemical models [4].

Tropical coastal environments are characterized by a range of extremely oligo- to eutrophic waters [5-8]. Lagoon waters

belong to the class of optically complex waters (classified as Case 2 waters) where mineral particles and colored dissolved

organic matter mix with phytoplankton [9]. Indeed, current algorithms such as OC4v4 for SeaWiFS and OC3 for MODIS

[10, 11] are suitable for oceanic waters where chlorophyll drives variability of bio-optical properties (absorption and

backscattering) of the waters. Attempts have been made to retrieve chlorophyll from remote sensing data in turbid case 2

waters [11-15]. Other algorithms tend to minimize the effect of bottom reflectance which increase surface reflectance values and therefore cause chlorophyll concentrations overestimation using algorithms developed for optically deep data [16].

The New Caledonian lagoon (22 177 km2, 25 m as a mean depth) lies in the South Western Tropical Pacific from 20°S to

22°S, and 166° to 167°E, with a heterogeneous bathymetry due to a complex geomorphology and a variety of different

bottom colors. It is largely connected to the open ocean in the south part of the lagoon, but only by narrow passes in the south

west part of the lagoon. Exchanges with the sea can modify the phytoplanktonic assemblage in the central lagoon

characterized by oligotrophic to mesotrophic waters (yearly average chlorophyll-a concentration of 0.25 ± 0.01 mg m-3

) [17,

18]. With relatively low river inputs and a low turbidity range compared with other tropical lagoons (0.20-16 g m-3

, [8], its

trophic state is linked to spatial variations in flushing times [19, 20].

Ocean Remote Sensing: Methods and Applications, edited by Robert J. Frouin, Proc. of SPIE Vol. 7459, 74590G · © 2009 SPIE · CCC code: 0277-786X/09/$18 · doi: 10.1117/12.829251

Proc. of SPIE Vol. 7459 74590G-1

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The ValHyBio project objectives were:

• to analyse the magnitude and spectral dependency of in situ optical data in the New Caledonia lagoon and open

ocean waters

• to examine the performance of the SeaWiFS OC4v4 algorithm to retrieve Chla from remote sensing reflectance, Rrs

• to elaborate an algorithm for Chla retrieval from MODIS and MERIS including the elimination of the bottom

reflectance effect

In this paper we analyse data from a particular cruise in March-April 2008 (ValHyBio) in comparison with a climatological

data set [21]. This cruise was representative of a heavy rainfall episode of the El Nino/La Nina changes in the Tropical

Pacific Ocean.

2. METHODOLOGY

2-1. In situ Sampling

The Valhybio cruise was designed to characterize the in situ inherent bio-optical properties of waters (absorption and

backscattering, IOPs) driving the remote sensing reflectance in the New Caledonia lagoon.

Figure 1: Map showing location of stations sampled during the Valhybio cruise. Dots indicate the station locations. In red, the

southernmost stations which were sampled only during the first network (22-28 March 2008). In blue, stations sampled twice during the

first (22-28 March 2008) and second network (7-9 April 2008) of the Valhybio cruise. Triangles indicate stations sampled every hour

during a 24h cycle. REF = reference oceanic station.

The main goal of these measurements was 1) to define the bio-optical properties of waters on different bottom types, 2) to

determine if chlorophyll and other optically significant constituents co-vary in lagoon waters. Sampling included a first

network of 53 stations spaced ~10 km apart in the lagoons and in the ocean off the barrier reefs (22 March-1 April: Map

Figure 1, blue and red circles). A total of 170 profiles were realized. The Valhybio cruise was during the peak of the 2008 La

Nina episode in the Tropical Pacific, and after 4 months of heavy rainfall.

2-2. Optical parameters and discrete water samples

Water sample was collected with Niskin bottles at different depths of the upper-water column and filtered onto 47 mm GF/F

filters. The different biogeochemical and optical parameters are the pigment concentrations as measured by

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spectrofluorometry, the suspended particulate matter (SPM; g m-3), and the particulate organic carbon (POC; mg m-3).

Filtrations were performed immediately onboard at low vacuum pressure. For pigments, the 25 mm GF/F filters were dipped

in 5.4 ml 100 % acetone (final concentration � 90% acetone taking into account water retention by the filter, i.e., 0.621 ±

0.034 ml) and ground with the freshly broken end of a glass rod for chlorophylls and phaeopigment extraction. This method

allows to get chlorophyll a, divinyl chlorophyll a (“Chla”, the sum of chlorophyll a and divinyl chlorophyll a, in mg m-3) and

accessory chlorophylls (b, c) and divinyl chlorophylls (b, c) [22]. For POC, the pre-combusted 25 mm GF/F filters were used.

The concentration of particulate organic carbon (POC) was determined using a Carbo-Elba elemental analyzer. For the

determination of the absorption coefficient, 1L sample was filtered. All filters were kept in liquid nitrogen until the analysis

at the laboratory. Before filtration for SPM dry weight, 47 mm polycarbonate filters used as recommended by the JGOFS

protocol were pre-combusted at 60°C during one hour in order to remove any trace of organic matter and preweighted. After

filtration, filters were rinsed with a solution of 1.08 M of formiate acid to eliminate salt, and kept in a dessicator in Petri

dishes and re-dried in an owen at 60°C during one hour before the determination of the SPM dry weight using a Perkin Elmer

microbalance. In situ optical parameters are measured from an optical package (WET Labs, Inc.) including a chlorophyll

fluorometer, a 10 cm pathlength beam transmissometer (at the wavelength 660 nm), and a Hydroscat-6 (HOBILabs, Inc.)

which measures the optical backscattering at 6 wavelengths (442, 488, 510, 555, 620 and 670 nm). The method used to get

the beam cp(660) from the beam transmissometer is the one used in [23]. The scattering coefficient bp(660) was obtained as

the difference between the beam cp(660) and the measured ap(660) from the filter pad method. The method used to get bbp

from the Hydroscat-6 is described in [24]. The backscattering ratio was calculated as the ratio of the average of bbp values at

620 and 670nm from the Hydroscat-6 to the beam cp(660).

3. RESULTS

3-1. Spatial distribution of biogeochemical and bio-optical parameters

The spatial distribution of physical parameters (temperature, salinity at 5 meters) obtained during the first network of the

Valhybio cruise (22 March to 31 March: Julian Days 82 to 92= 10 days) is shown at Figure 2AB. Fresher (salinity < 34.2)

and colder waters (SST<26) are observed in the South Western lagoon (Stations b08 to a20) and in the South Eastern lagoon

(Station t01 and t17, t04 and t05). They are issued from the two main rivers, the Dumbea river on the west off Noumea city

and the Coulée river at the south of the Noumea city and from rivers at the southern tip of the of the main land (connected to

the large Yate lake). Fresher and colder waters than normal were advected in the lagoon and both low salinity (Fig. 2A) and

colder waters (Fig 2B) are tracers of the rain impact on the lagoon.

Figure 2: Spatial distribution of (A) Temperature, (B) Salinity for the first network (22 March to 31 March: Julian Days 82 to 92= 10 days)

of the Valhybio cruise.

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Figure 3: Spatial distribution of (A) Chla, (B) POC, (C) SPM, (D) POC:Chla for the first network (22 March to 31 March: Julian Days 82

to 92= 10 days) of the Valhybio cruise.

Chla and POC concentrations increase (Fig. 3AB) from coastal stations to offshore. They are maximum in the South Western

lagoon (between n43 and oc1 to p04-p12) and over a large area of the South Eastern lagoon (t01 to t16-t17). Concentrations

higher than 0.4 mg m-3 for Chl and higher than 100 mg m-3 for POC are tightly associated with fresher and colder waters

(Fig 2AB). Chla and POC distributions are similar in the fresh plume identified from stations t1 to t24-t25. Higher

concentrations of Chla and POC concentrations result from a large input of silica with the maximal values in the fresher

waters (not shown).

High SPM (> 0.9 mg/L) are observed in coastal areas, and are well related to low salinity waters. Isolated maxima at the

barrier reef (st 11 and 12) and in the region of a21-a22 are also observed. Low POC:Chla (< 200) characterize fresh and cold

waters influenced by land drainage. The POC:Chla increases from the coast to offshore waters with maxima (> 300) in the

southern part of the lagoon in oceanic oligotrophic waters between the two barrier reef tips (st 21 to st 27). The low POC:

Chla ratio is associated with rich in phytoplankton nearby the coast and in the fresh plumes. High POC:Chla ratios are

characteristic of oligotrophic waters where detrital carbon dominates.

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Regarding the optical parameters (Fig 4 ABCD), spatial distributions of ap(442) and bbp(510) strongly differ. The plume of

high ap(442) is found at stations t04-t05 (> 0.12 m-1

) and correspond to fresh waters as defined by a salinity lower than 34.4.

The distribution of bbp(442) shows a different pattern than ap(442). There is no increase of bbp(442) in the freshwater plume

(t04-t05). Moreover, freshwater plume values around t04-t05 do not differ from the values observed in the rest of the south-

eastern lagoon (0.005 m-1

). Contrary to ap(442), the maximal values of bbp(510) are found in the south-western part of the

lagoon (around the city of Noumea, between d27 to p12). This difference between the ap(442) and bbp(510) distributions is

typical of phytoplankton rich waters which exhibit comparatively high absorption and a low backscattering efficiencies

compared to mineral particles [7, 9].

Figure 4: Spatial distribution of (A) ap(442), (B) bbp510, (C) log(abs(cp(660)), (D) bbp/bp(650) for the first network (22 March to 31 March:

Julian Days 82 to 92= 10 days) of the March-April 2008 Valhybio cruise.

The distribution of cp(660) mimics the one of the Chla (note the inverse scale of Figure 4) with the highest values in the

chlorophyll-rich plumes. The backscattering ratio bbp/bp(660) shows the inverse pattern than cp(660) and reaches its lowest

values (< 0.02) in the fresh chlorophyll-rich water plume. This is in favour of a low backscattering ratio for the particles

advected in the large freshwater plume at the southeast and composed mainly of phytoplanktonic cells.

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3-2. Relationships between IOPs and Chla for the Valhybio cruise

Chla frequency distributions in the New Caledonia lagoon are linked to the lagoon bathymetry, with a median value of 0.16

mg m-3 in oligotrophic waters offshore (depths larger than 200 meters), and with a median value of 0.25 mg m-3

in deep

stations (> 20 meters). The higher Chla values are usually observed in shallower waters and in enclosed bays (< 20 meters

depth) as described previously (Figure 6, from [21]).

Figure 5: Histograms showing the frequency distribution of the chlorophyll concentrations for the A) open ocean waters, B) D deep lagoon

stations (bottom depth > 20 m), C) S shallow lagoon stations (bottom depth < 20m), D) Valhybio stations.

As the majority of the Valhybio stations are belonging to the D group (all deeper than 20 meters, except B08 and N25), Chla

must be compared to the frequency distribution of D stations of the 2001-2007 databasis (Figure 5C). The median value

during the Valhybio cruise is much higher (0.63 mg m-3

) than the median value of the D stations (Figure 6 ABCD) and

approaches the median value of the shallowest stations. This Chla increase in D stations is due to a large input of silica and

nitrate in the fresh water plume (data not shown).

The relationships between IOPs and Chla are shown in Figure 6A for ap(442) and Figure 6B for bbp(510) with the same

classification of stations as before. For the set of open ocean waters, relationships between ap(442) and bbp(510) are in good

agreement with the curves representing published models for case I waters [25, 26]. For the lagoon D stations or for the

Valhybio stations, the relationship between ap(442) and Chla is good before 1 mg m-3. There is much scatter in this

relationship for the S stations. A the opposite, there is a much larger variability for the relationship between bbp(510) and

Chla in the lagoon waters. This is related to the larger amount of mineral particles in lagoon waters than offshore [21].

A strong relationship is observed between bbp(510) and SPM for the shallow stations in the lagoon (Figure 7A). Note the

small range of SPM values of the Valhybio cruise stations (maximum value of SPM=2 g m-3

). Different relationships are

observed between bbp(510) and cp(660) according to the station classification used above [21]. For the Valhybio stations, the

bbp(510)/cp(660) relationship differs (in red, Figure 7B) from the one obtained for the D stations. This backscattering ratio at

Valhybio is lower for the same Chla than for the D stations during the period 2001-2007 and is also closer than the one of

0

10

20

30

40

50

600.1

0.1

6

0.2

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0.4

0.6

3 1

1.5

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3.9

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Chl a (mg m-3

)

Frequency (

%)

VALHYBIO

D

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)

Frequency (

%)

Lagoon D >20m

B

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%)

Lagoon S <20m

C

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%) open ocean

A

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open ocean waters. This shows that the proportion of mineral particles is less than usually in the deep stations of the lagoon

of New Caledonia during the Valhybio cruise.

Figure 6: Scatter plots as a function of measured Chla of A) phytoplankton absorption coefficient at 676 nm and B) particulate

backscattering coefficient at 510 nm. Open ocean waters from nine Diapalis cruises at 167°E, 21°S (light blue); D lagoon stations (bottom

depth > 20 m; dark blue), S lagoon waters (bottom depth < 20 m, orange losanges). Thick grey curves correspond to the model of [25] for

ap(676) and [26] for bbp(510)

Figure 7: A) Relationship between particulate backscattering coefficient at 510 nm and Turbidity as expressed in NTU (S stations, orange

losanges), B) relationship between bbp(510) and cp(660) for the different type of stations. Symbols are the same as in Figure 6AB.

3-3. Performance of OC4 algorithm for the Valhybio cruise

The performance of the OC4 algorithm [10, 11] was tested using reflectance calculated with our measured IOPs as in [21]

(Figure 8). The performance of the OC4v4 algorithm (the correlation between Chla and the maximum of reflectance ratios) is

poor when evaluated for the global data set, which corresponded to a mixture of case 1 and case 2 water types (Fig. 8 AB)

[21]. The OC4v4 algorithm was inadequate for the S stations (overestimation by a factor of about 3 on average and almost no

correlation with Chla), because the bio-optical properties driving remote sensing reflectance, viz. the absorption and

Hydroscat-6 backscattering coefficients are poorly related to Chla. On the other hand, at the open ocean stations where

essentially Model (Rrs) values were available (Chla in the range 0.07-0.4 mg m-3

), OC4v4 underestimated Chla with a mean

bbp(510)= 0.0116* Turb

R2 = 0.9871

0

0.02

0.04

0.06

0.08

0.1

0 2 4 6 8 10

Turbidity (NTU)

bb

p(510) (m

-1)

C

D stations

y = 0.0328x - 0.0001

R2 = 0.63

y = 0.0091x + 0.0002

R2 = 0.4089

VALHYBIO

y = 0.0177x

R2 = 0.4482

0

0.01

0.02

0.03

0 0.2 0.4 0.6

cp (660nm) (m-1

)

bb

p510 (

m-1

)

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)

aph

(676) (m

-1

)

VALHYBIO

Lagoon S < 20m

Lagoon D > 20m

case 1 model

open ocean

A

0.00

0.01

0.02

0 0.5 1 1.5 2

Chl a (mg.m-3

)

bbp

(510) (

m-1

)

VALHYBIO

Lagoon S < 20 m

Lagoon D > 20m

Open ocean

Case I model

B

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bias of -33% [21]. As expected from the analysis of the IOPs, the relationship between the OC4v4 retrieved Chla and

measured Chla does not differ from the one obtained on the D stations of the climatological data set.

Figure 8: A) Relationship between modeled Max[Model(Rrs)/(Rrs)]-OC4-Chl a and measured Chla for the global data set. The grey thick

curve is obtained by the NASA OC4v4 algorithm. B) Comparison between Chla derived from the OC4v4 algorithm for SeaWiFS and field

spectrofluorometric determinations on surface water samples. The line represents one-to-one perfect agreement. Open ocean waters, light

blue: D lagoon waters (> 20m), dark blue, S lagoon waters (< 20 m) orange losanges, and ValHyBio stations in red.

4. CONCLUSIONS

The influence of a heavy rainfall episode of the El Nino/La Nina changes in the Tropical Pacific Ocean was described on the

IOPS of the lagoon of New Caledonia compared with a database compiled over the drier period 2001-2007 including

Bissecote and Echolag cruises [21].

Major changes were observed in relation with an increase in pigment concentration by a factor of 3 related to an increase in

nutrients from strong land drainage after a 4 months rain episode. The increase in SPM was moderate compared to the

climatology. The SPM distribution was not fundamentally different than that of Chla.

Chlorophyll-rich plumes mainly related to the coast were later advected in the middle part of the lagoon. These rich plumes

had different characteristics according to the specific area concerned. The Southwestern plume (near the city of Noumea) was

both absorbing and backscattering. The Southeastern plume was strongly absorbing but only weakly backscattering. Both

plumes had a low backscattering ratio, indicating a majority of living particles.

The particulate backscattering ratio (bbp/bp) which describes the nature of the bulk particulate assemblage in the ocean color

variability [27, 28] was useful in differentiating the fresh and cold water influenced by land runoff. The range of this ratio

was 0.01-0.05 (at 660 nm) which is a low range compared to other tropical ecosystems such as the Australian Great Barrier

Reef [7, 29]. Low backscattering ratios and low POC:Chla characterized the chlorophyll rich plumes issued from the nutrient

increase and higher backscattering ratios (> 0.05) and high POC:Chla ratios were typical of the oligotrophic oceanic waters.

The performance of OC4 was not affected in the deep stations of the lagoon of New Caledonia by the rainfall runoff in

relation to the small modification in the particle composition (mainly an increase in the size and abundance of

phytoplanktonic cells rather than an increase in mineral particles).

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0.01

0.1

1

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Chl a (spectrofluorometric) mg.m-3

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