Biodeposit production by Bractechlamys vexillum (bivalve: pectinidae) in a tropical lagoon

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Journal of Sea Research xxx (2010) xxx–xxx

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Biodeposit production by Bractechlamys vexillum (bivalve: pectinidae) in atropical lagoon

Jacques Clavier ⁎, Laurent ChauvaudLEMAR, Laboratoire des Sciences de l'Environnement Marin, UMR CNRS 6539, Institut Universitaire Européen de la Mer, Technopôle Brest-Iroise, Place Nicolas Copernic,29280 Plouzané, France

⁎ Corresponding author. LEMAR (UMR CNRS 6539), Ide la Mer, Technopôle Brest-Iroise, Place Nicolas CoperTel.: +33 2 98 49 87 39; fax: +33 2 98 49 86 45.

E-mail address: [email protected] (J. Clav

1385-1101/$ – see front matter © 2010 Elsevier B.V. Aldoi:10.1016/j.seares.2010.03.001

Please cite this article as: Clavier, J., ChauvaJ. Sea Res. (2010), doi:10.1016/j.seares.201

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Article history:Received 5 January 2010Received in revised form 22 February 2010Accepted 1 March 2010Available online xxxx

Keywords:BivalvePectinidBiodepositionLagoonSestonFiltration

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OIn the SW lagoon of New Caledonia (South Pacific), the pectinid Bractechlamys vexillum (Reeve) is acharacteristic species of the 1000-km² grey sand bottoms benthic community, with an average density of1.65 ind m−2 and a mean dorso-ventral shell height of 47 mm. To assess the role of B. vexillum populations inthe transfer of matter at the water–sediment interface of the lagoon ecosystem, in situ measurements wereperformed of individual biodeposition of total particulate matter, particulate organic matter, particulateorganic carbon and nitrogen, chlorophyll a, and pheopigments. Fluxes of particulate matter in sedimenttraps, with and without individuals placed in cages, were compared at various seasons (66 experiments) andrelated to concentrations in seston. Particulate matter biodeposit production was significantly linked to B.vexillum biomass and to suspended chlorophyll a and pheopigment concentrations, but not to temperatureor to other particulate material concentrations in seston. Daily average biodeposit production of totalparticulate matter and particulate organic carbon by unit of biomass were 298 mg g AFDW−1 d−1 and45 mg g AFDW−1 d−1, respectively. At the study site, biodeposition by the B. vexillum population was570 mg m−2 d−1 of total particulate matter and 68 mg m−2 d−1 of organic carbon, which represent 4.8%and 6.4% of natural sedimentation, respectively. This relatively low figure at the local scale results, however,in a total contribution of B. vexillum to particulate sedimentation in the whole lagoon of 34·103 metrictons year−1 for total particulate matter and 4·103 metric tons year−1 for organic carbon.

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nstitut Universitaire Européennic, F-29280 Plouzané, France.

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l rights reserved.

ud, L., Biodeposit production by Bractechlamy0.03.001

© 2010 Elsevier B.V. All rights reserved.

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The two main sources of energy for coastal zoobenthic commu-nities are benthic primary production and sedimentation of organicmatter (Mann, 1982). Biological activity of autotrophic organismsdrives the former (Gattuso et al., 2006) whereas the latter is mostlydetermined by physical–chemical processes influencing transport andsedimentation of particulate material (Passow and De La Rocha,2006). Suspended particulate organic matter is composed of bothallochthonous material and autochthonous living organisms, mainlyplankton (Otero et al., 2000). In coastal areas, the rate of benthicprimary production increases where low depth allows high irradianceto reach the seafloor and where the vicinity of the shore increasesnutrient availability (Gattuso et al., 2006). However, particulatematter sedimentation remains an important factor in benthicecosystem functioning (Lundsgaard and Olesen, 1997).

Biological activity influences fluxes of particulatematter at thewater–sediment interface (Graf and Rosenberg, 1997). The sedimentation of

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organic and mineral particles from seston can be passively enhancedwhen hydrodynamics slows near the bottom as a result of biogenicstructures fromvegetation, suchas seagrass beds (Duarte et al., 2005), andfrom macrozoobenthos (Friedrichs et al., 2009), such as polychaete tubeassemblages (Peine et al., 2009). It can also depend on active filtration ofwater by suspension feeders with biodeposition of the non-digestedmaterial (Dame, 1996; Ragueneau et al., 2005). Filter-feeding bivalvespump water through the gills for respiration, and at the same time, theparticle retaininggillsfilter the through-flowingwater for suspended foodparticles, with an associated biodeposition of fecal pellets (feces) andmucus-bound pseudofeces (Urrutia et al., 2001) originating from theparticles retained by the gills. When abundant, bivalves can be importantcontributors to mineral and organic matter transfer towards the water–sediment interface, both in natural and cultivated populations (Cranfordet al., 2009;Newell, 2004). Suspension feeders are amajor feature of coralreef communities (Yahel et al., 2002) and mostly correspond to soft orcalcified corals (Fabricius and Dommisse, 2000; Houlbrèque and Ferrier-Pagès, 2009), but filter-feeder bivalves can be abundant in some lagoons.This abundance is particularly the case for natural and cultivatedpopulations of pearl oysters (Berthe and Prou, 2007), but other bivalvescan also live at high densities (Niquil et al., 2001).

The SW lagoon of New Caledonia covers about 2000 km², 95% ofwhich is occupied by sediments (Chardy et al., 1988). The average

s vexillum (bivalve: pectinidae) in a tropical lagoon,

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depth of the lagoon is 17.5 m (Douillet et al., 2001). Three main softbottom macrobenthic assemblages have been recognized in thislagoon: mud deposits near the coast and in the deeper parts, the greysand bottoms community in the middle, and the white sand bottomscommunity behind the reef. They make up 35%, 50%, and 15% of thelagoon surface, respectively (Chardy et al., 1988). The average softbottom macrofaunal biomass of the lagoon has been estimated to be13.0 g AFDWm−2 (Chardy and Clavier, 1988). Suspension feeders,representing two-thirds of the total animal biomass, are dominated bysolitary corals like Heteropsammia cochlea (Spengler) and mollusks.The bivalve Bractechlamys vexillum (Reeve) is a medium-sizedpectinid with a maximum dorso-ventral shell height of about65 mm, distributed throughout the Western Pacific region (Dijkstra,1985). In the SW New Caledonia lagoon, this bivalve is a dominantspecies in terms of biomass, as is characteristic of a grey sand bottomscommunity (Chardy and Clavier, 1988), where it lives unattached atthe surface of the sediment. The aim of this study was to evaluate insitu the individual daily biodeposit production by B. vexillum inrelation to environmental parameters and to estimate the contribu-tion of populations to the transfer of particulatematerial at thewater–sediment interface of the lagoon.

2. Material and methods

2.1. Study site

The “Rocher à la Voile” site is located in the SW lagoon of NewCaledonia, near the Nouméa peninsula (Fig. 1). The average depth is

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Fig. 1. Location of the study site in th

Please cite this article as: Clavier, J., Chauvaud, L., Biodeposit productionJ. Sea Res. (2010), doi:10.1016/j.seares.2010.03.001

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10 m. According to Chardy et al. (1988), this place corresponds to atypical “grey sand bottoms” benthic community with the highest B.vexillum density (10 ind m−2) recorded in the lagoon. Currents aremoderate and driven both by tide and SE trade winds with velocitieslower than 0.1 m s−1 (Douillet et al., 2001). The average salinity in theSW lagoon is 35.5 with low seasonal and inter-annual variation(±0.3) except during tropical cyclone events (Ouillon et al., 2005),which did not occur during the present study periods.

2.2. Biodeposit processing

The study was conducted using sediment traps made of PVC tubes,16 cm in diameter and 50 cm high, with an internal volume of ca. 10 L.Two traps were fastened by divers to a vertical mooring, with the top1.5 m above the bottom. Stainless steel cages (8×8 cm)with a 10-mmmesh were secured in the middle of the top of the traps, with thelower part 1 cm under the rim (Fig. 2). B. vexillum was collected bydiving from the sea floor under the traps. The shells were cleaned ofdeposits and of possible epibionts, which are seldom found on thespecies. One individual was closed in a cage while the other cage waskept empty to provide a reference for natural sedimentation in thetraps. B. vexillum individuals were selected to cover the range of sizesobserved in the natural population (Fig. 3). At the initiation of eachexperiment, two identical double-trap (mollusk+reference) systemswere simultaneously installed at the same depth, at a distance of ca.10 m from each other. To bypass the effects of any short-termvariation in sedimentation fluxes, the traps were deployed for 24 h.Then, the cages were removed and the bivalves kept for subsequent

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e SW lagoon of New Caledonia.

by Bractechlamys vexillum (bivalve: pectinidae) in a tropical lagoon,


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Fig. 2. Schematic representation of the experimental device. (A) Cylindrical sediment traps,(B) cagewith oneB. vexillum individual, (C) cagewithoutB. vexillum. Themooring apparatusincludes (D) a ballast weight, (E) a mooring line tensioned by (F) a subsurface buoy.

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analysis. The traps were closed by a cylindrical lid fit over the top andthen moved up to the surface and transferred to the laboratory in thevertical position. A total of 33 experiments (66 individuals) werecarried out at various seasons.

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198Fig. 3. Shell height distribution of Bractechlamys vexillum in the (A) experimental and(B) natural populations at the study site.


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2.3. Seston sampling

To compare the characteristics of seston at the trap level and nearthe sediment where scallops naturally live, divers collected two sets oftriplicate samples of water just after each trap deployment. Sampleswere taken at the top of the trap level and about 5 cm above thewater–sediment interface, with care to avoid sediment suspension.Divers collected the water in 500-mL bottles, which were filled withnatural seawater at the surface. At the required depth, the bottleswere placed head-down, flushed with air from a diving regulator, andcarefully filled anew with ambient seawater. Sea surface temperature(SST) was recorded during each experiment with 0.1 °C precision.

2.4. Filtrations

Filtration of the trap contents was achieved in the laboratory lessthan 2 h after the end of the experiments. The trap contents werestirred, poured into a 20-L jar, and kept homogenized under agitationduring subsampling. Four subsamples of about 150 mLwere collected.Samples were filtered on pre-ashed and pre-weighed GF/C filters(1.2 μm nominal pore size) to assess sedimented total particulatematter (TPM), particulate organic matter (POM), particulate organiccarbon (POC), particulate nitrogen (PN), chlorophyll a (Chl a), andpheopigment (Pheo) concentrations. Filters were rinsed twice usingan isotonic ammonium formate solution to expel salt. For comparisonpurposes, only total particulate and organic matter concentrationswere considered in seawater at the trap level. Water samples forsuspended (SPM) and organic (SPOM) particulate matter concentra-tion assessment at the trap level, and SPM, SPOM, suspendedparticulate organic carbon (SPOC), nitrogen (SPN), chlorophyll a(SChl a), and pheopigments (SPheo) at the bottom level, were alsofiltered on pre-ashed and pre-weighed GF/C filters and treated asdescribed above. Filters were kept frozen (−20 °C) until analysis (lessthan one month later).

2.4. Analytical procedures

Color was used to assess the sex of B. vexillum, with white or creamindicating a male gonad and orange indicating the female gonad.Individuals without distinctive gonads were classified as undifferen-tiated and corresponded to both immature and post-spawning.Dorso-ventral shell height was measured using a vernier caliper.Ash-free dry weight (AFDW) was used to determine biomass andcalculated as the difference between the flesh weight (DW) after 8 dof drying at 60 °C and the ashweight after calcination at 450 °C for 2 h.

TPM and SPM were calculated as the difference in weight of thefiltered material before and after drying at 60 °C for 72 h. POM andSPOM correspond to the difference in weight before and aftercalcination at 450 °C for 2 h. Suspended and sedimented POC andPN content of the freeze-dried material was measured using a CNHanalyzer (Hewlett-Packard 185B), after fuming the filters overconcentrated HCl in a closed container to remove inorganic carbon.Photosynthetic pigments were extracted using 90% acetone at 4 °C inthe dark for 8 h and measured on a fluorometer (TURNER 112).Suspended and sedimented Chl a and Pheo concentrations weredetermined according to Lorenzen (1967).

Naturally sedimented TPM, POM, POC, PN, Chl a, and Pheo werecalculated as the average value for the two reference traps with emptycages. Biodeposit production of TPM, POM, POC, PN, Chl a, and Pheocorresponded to the difference in content of two coupled traps(mollusk and reference). To assess the volume of water processed byB. vexillum, we calculated the weight standardized clearance rate (CR,L g AFDW−1 h−1) as the ratio between the biodeposition of inorganicmatter (in feces+pseudofeces) measured in the trap, divided by theindividual biomass (mg AFDW) (Iglesias et al., 1998), and the



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Table 1t1:1

Characteristics of the experimental population of Bractechlamys vexillum.t1:2t1:3 N Average dorso-ventral

shell height (mm)Average biomass (g)

t1:4 Male 30 47.03 0.173t1:5 Female 26 50.23 0.200t1:6 Undifferentiated 10 37.24 0.127t1:7 Global 66 46.81 0.177


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particulate inorganic matter concentration in seston (mg L−1) whichis the difference between SPM and SPOM.

2.4. Statistical analysis

Because the number of measured parameters was relatively low,we used bivariate rather than multivariate analysis to establish therelationships between variables. These relationships were investigat-ed by plotting and inspecting the data. We then tested differentmodels to fit the data using least-squares curve fitting with theLevenberg–Marquardt algorithm, and selected the model thatexplained the maximum of variance (highest r²). When modelswere almost the same in their explanatory value, we reported resultsfor the simplest one (usually the linear regression model). Model IIlinear regression analysis was used when measurement erroroccurred for both X and Y variables (Laws and Archie, 1981), andparameters were compared using Z tests (Scherrer, 1984). One-wayANOVA and Student t-tests were used to compare means. Data werefirst tested to check the homogeneity of variances (Levene's test). In

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Fig. 4. Monthly evolution of (A) measured sea surface temperature, concentrations of (B) s(C) suspended organic carbon (SPOC) and suspended particulate nitrogen (SPN), and (D) s


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the absence of homoscedasticity, samples were compared using theKruskal–Wallis and Mann–Whitney U tests.

3. Results

3.1. B. vexillum characteristics

B. vexillum dorso-ventral shell height (Table 1) varied from 19.5 to64.9 mm and individual biomass from 0.01 to 0.381 g. Average shellheight and biomass (AFDW) differed significantly according to sex(ANOVA, pb0.01 for both), but a post-hoc analysis (least significantdifferences) indicated that only mean shell height and biomass ofundifferentiated individuals were significantly lower. The generalshell height distribution of the experimental population was morespread out than that of natural population (Fig. 3), but their averagesizes (46.47 mm and 46.62 mm, respectively) did not differ signifi-cantly (t test, pN0.05). AFDW to dry weight (DW) ratio calculated forthe experimental population was 0.83 (N=66; r²=0.91).

3.2. Environmental parameters

SST (Fig. 4A) varied from 23.0 °C (September) to 28.3 °C(February) during the experiments, with an average value of 25.9 °C(SD 1.9). SPM and SPOM concentrations in seston at the trap and atthe bottom level were highly correlated (Fig. 5), and the slopes of themodel II regressions (SPM: 1.02; SPOM: 1.05) did not differsignificantly from 1 (Z test, pN0.05). Monthly SPM, SPOM, SPOC,and SPN concentrations measured at the bottom level (Table 2) variedseasonally with a maximum in June (cool season) and a minimum in

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uspended particulate matter (SPM) and suspended particulate organic matter (SPOM),uspended chlorophyll a (SChl a) and suspended pheopigments (SPheo).



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Fig. 5. Relationships at the bottom and the trap level between (A) suspendedparticulate matter (SPM) and (B) particulate organic matter in seston (SPOM), withadjusted linear models. r² is the coefficient of determination.

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January (warm season) (Fig. 4B–C). These parameters were signifi-cantly positively correlated with each other (pb0.01 for all correla-tions). SChl a and SPheo varied also seasonally but with a minimum inJuly and a maximum in February (Fig. 4D), they were significantlynegatively correlated to other variables in seston (pb0.01). Eachvariable changed seasonally according to SST (pb0.05) except SPON(p=0.07). SPM, SPOM, and SPOC were negatively correlated, andphotosynthetic pigments were positively, linearly related to SST. POMrepresented 32% of SPM, and SPOC made up 19% and SPON 1% ofSPOM. SPheo concentrations were relatively high and correspondedto 69% of SChl a concentration.

3.3. Clearance rates

The annual average CR was 6.9 (SD 6.6) L g AFDW−1 h−1 (5.7 L gDW−1 h−1). With a mean individual biomass of 0.177 g AFDW, B.vexillumwouldprocess a volumeofwater of 29 L d−1. CR increasedwith

UNTable 2Characteristics of suspended particulate material (seston) and sedimented material in trapcorresponding ratio. Standard deviations are in parentheses.

Seston (μg L−1) Trap

Sedimentatio

Total particulate matter 4495.033 (4231.319) 240.453 (108Particulate organic matter 1455.789 (1085.050) 49.433 (28.Particulate organic carbon 282.334 (149.926) 21.452 (8.5Particulate nitrogen 23.324 (9.337) 1.052 (0.4Chlorophyll a 0.468 (0.226) 0.012 (0.0Pheopigments 0.323 (0.143) 0.018 (0.0


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SST (Table 3); the average value was 3.03 (SD 1.69) L g AFDW−1 h−1

belowand 9.63 (SD6.17) L g AFDW−1 h−1 above 27 °C. CR dramaticallydecreased with SPM concentration (Fig. 6A), with an average value ofonly 2.74 (SD 1.27) L g AFDW−1 h−1 above 2.6 mg SPM L−1. Similarly,CR decreased with increasing SPOM, SPOC, and SPN concentrations(Table 3). In contrast, CR had a significant positive linear relationshipwith Chl a (Fig. 6B) and Pheo (Table 3). CR was not significantly relatedto individual biomass (Fig. 6C, r²=0.01, p=0.51) and did not varysignificantly with sex (Kruskal–Wallis test, p=0.49).

3.4. Daily biodeposition rates

Daily individual biodeposit production of POM, POC, PN, and Pheoin the traps represented, on average, about one-third of naturalsedimentation (Table 2), whereas TPM contribution was slightlylower. In contrast, the average Chl a biodeposition was clearly inferiorto natural sedimentation in the traps (Table 2). The slope of the modelII regression between biodeposited TPM and biodeposited POM (3.92,SD 0.18) differed significantly from the slope of the regressionbetween SPM and SPOM (2.58, SD 0.20), indicating an enrichment oforganic material in biodeposits. All parameters measured on biode-posits were positively correlated with each other (pb0.01), and theywere significantly related positively to biomass of B. vexillum andnegatively to both SChl a and SPheo concentrations (Tables 3 and 4).Biomass accounted for more than 50% of the variance for anybiodeposition parameter, and it was by far the main explicative factorof daily individual biodeposition. Relationships were not significantfor the other seston variables. Biodeposit production was not relatedto SST and to natural sedimentation, except for TPM and PN, whichwere significantly related to sedimented POM and PN, respectively(Tables 5 and 6). C/N did not differ significantly in biodeposits (24.8,SD 14.4) from natural sedimentation (20.9, SD 5.0) but wassignificantly lower (12.0, SD 3.9) in seston (U-test, pb0.01).

The individual hourly biodeposit production per unit of biomassdid not vary significantly according to sex for TPM, POM, POC, PN, orChl a (Kruskal–Wallis test, pN0.05) but differed significantly for Pheo(Kruskal–Wallis test, p=0.01). Mann–Whitney U-test results indi-cated that only undifferentiated individuals differed from males andfemales for Pheo (pb0.01). The relationships between individualbiomass and biodeposit production of TPM, POM, POC, PN, and Pheoper unit of biomass revealed maximal fluxes for the smallerindividuals and then a decrease towards an asymptote (see Fig. 7Afor an example of POC). Only Chl a showed an opposite pattern, withnegative fluxes for juveniles increasing toward a positive asymptotefor larger individuals (Fig. 7B). Individual hourly biodeposit produc-tion per unit of biomass was not related to CR (Fig. 6D) for anyparameter.

3.5. Contribution to water and particulate fluxes

B. vexillum density in the 1000-km2 grey sand bottoms communityis 1.65 ind m−2 (Chardy et al., 1988). The average volume of waterprocessed by the species is 48 L m−2 d−1 and 48·106 m−3 d−1 for

s both naturally (sedimentation) and by biodeposition by Bractechlamys vexillum, and

n (mg trap−1 d−1) Biodeposition (mg ind−1 d−1) Ratio

.671) 57.007 (29.201) 0.24640) 18.455 (11.317) 0.3734) 6.809 (4.455) 0.3211) 0.358 (0.282) 0.3409) 0.002 (0.002) 0.1708) 0.006 (0.005) 0.33



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Table 3t3:1

Relationships between the clearance rate (L h−1 g AFDW−1), sea surface temperature(SST, °C), and suspended particulate variables. Parameters for SST and suspendedparticulatematter (SPM,mg L−1), suspended particulate organicmatter (SPOM,mg L−1),suspended particulate organic carbon (SPOC, μg L−1), and suspended particulate nitrogen(SPN, μg L−1) correspond to the equation y=a /x+b. Parameters for suspendedchlorophyll a (SChl a, μg L−1) and suspended pheopigments (SPheo, μg L−1) correspondto the equation y=ax+b. r² is the coefficient of determination, and p is the relatedprobability. The relationships are illustrated by the fitted curves for SPM and Chl a in Fig. 5.

t3:2t3:3 Variable a b r²

t3:4 SST −1046.40 46.88 0.30t3:5 pb0.001t3:6 SPM 21.08 0.29 0.69t3:7 pb0.001t3:8 SPOM 6.02 0.38 0.32t3:9 pb0.001t3:10 SPOC 1859.87 −1.79 0.40t3:11 pb0.001t3:12 SPN 226.16 −4.51 0.38t3:13 pb0.001t3:14 SChl a 13.58 0.03 0.19t3:15 p=0.001t3:16 SPheo 20.05 −0.12 0.14t3:17 p=0.004


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the whole lagoon. The water volume of the 2000 km² lagoon (averagedepth 17.5 m) can be estimated to 35·109 m3, and corresponds to2 years of total through-flowing water filtered by the B. vexillumpopulation gills.

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Fig. 6. Relationships among clearance rate (CR) of Bractechlamys vexillum and (A) suspended p(C) individual biomass and (D) hourly biodeposit production per unit of biomass (TPM). The par0.01, respectively.


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With an average density of 10 ind m−2 (Chardy et al., 1988), thenatural population of B. vexillum at the Rocher à la Voile site accounts for570 mgm−2 d−1 of TPM and for 68 mgm−2 d−1 of POC, whichrepresent 4.8% and6.4% ofmeasurednatural sedimentation, respectively.B. vexillum TPM and POC biodeposit production in the 1000-km2 greysand bottoms community are 94 mg m−2 d−1 and 11 mgm−2 d−1,respectively. The total contribution of the species to fluxes of particulatematter at the water–sediment interface of the grey sand bottomscommunity of the SW lagoon of New Caledonia can be estimated at34·103 metric tons year−1 for TPM and 4·103 metric tons year−1 forPOC.

4. Discussion

4.1. Methodological considerations

Filtration and biodeposition in bivalves can be studied in threemainways. Laboratory experiments generally rely on cultured phytoplanktoncells to assess, under fully controlled conditions, the clearance rate andthe size distribution of particles retained by gills. Other studies usenatural seston pumped from thenearby coastalwater into experimentalenclosures and provide more representative estimates of filtration innature (MacDonald and Ward, 2009; Strohmeier et al., 2009). Thesemethods are of particular interest to study the ecophysiologicalpotential of filter feeders and related processes. However, controlledexperimental conditions are often far from true representations of thecomplex web of interactions that govern the functioning of the natural

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articulate matter concentration (SPM), (B) chlorophyll a concentration in seston (SChl a),ameters for thefitted curves inA andB are given inTable 3. r² values in B andDare 0.04 and



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Table 4t4:1

Relationships between individual hourly biodeposition per unit of biomass andindividual biomass (g AFDW). POM, biodeposited particulate organic matter (mg gAFDW−1 h−1); POC, biodeposited organic carbon (mg g AFDW−1 h−1); Chl a,biodeposited chlorophyll a (μg g AFDW−1 h−1); Pheo, biodeposited pheopigments(μg g AFDW−1 h−1). Parameters correspond to the equation, y=a /x+b. r² is thecoefficient of determination, and p is the related probability. The fitted curves for POCand Chl a are drawn in Fig. 7.

t4:2t4:3 Variable a b r²

t4:4 POM 0.11 3.87 0.42t4:5 pb0.001t4:6 POC 0.06 1.24 0.58t4:7 pb0.001t4:8 Chl a −0.04 0.76 0.75t4:9 pb0.001t4:10 Pheo 0.04 1.24 0.43t4:11 pb0.001

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environment. An evaluation of the ecological influence of filter feederson natural communities and their environment requires in situexperiments in which tested individuals are as close as possible totheir natural condition (Hewitt and Norkko, 2007).

In the SW lagoon of New Caledonia, high particulate matterconcentration is often observed near the water–sediment interfacewhere B. vexillum is living (Ouillon et al., 2004). However, we foundsimilar concentrations of SPM and SPOM at the bottom and at the traplevel, 1.5 m above ground. We extrapolated this result to the othervariablesmeasured in seston and inferred that our experimental protocolcan provide a realistic view of natural biodeposition of B. vexillum.

4.2. Filtration of B. vexillum

CR was calculated by the “biodeposition method” based on theconservative nature of inorganic matter in pooled feces and pseudo-feces. Differentmethods are used formeasurement offiltration rates inbivalves (see Riisgård, 2001, for a review). Petersen et al. (2004) foundthat this method provided an underestimation of the actual clearance

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Table 5Relationships among total particulate matter (TPM, mg trap−1 d−1), particulate organic matter (nitrogen (PN,mg trap−1 d−1), chlorophyll a (Chl a, μg trap−1 d−1), andpheopigments (Pheo, μg t(SPM, mg L−1), suspended particulate organic matter (SPOM, mg L−1), suspended particulatechlorophyll a (SChl a, μg L−1), and pheopigments (SPheo, μg L−1). N is the number of observationcorrespond to significant correlations (pb0.05).


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rate, whereas Bayne (2004) stated that the procedure was suitable forbivalves under natural conditions. The major drawback of thetechnique is that pseudofeces can be dispersed by hydrodynamicsfrom the container where mollusks are kept, but such a problem islikely to be minor in 50-cm-high sediment traps. Furthermore, 24-hmeasurements allow for short-term changes in the feeding behaviorrelated to bivalve behavior, fluctuations in current speed, or sestonconcentration or composition (Strohmeier et al., 2009). The weight-standardized CR of B. vexillum (5.7 L g DW−1 h−1) was smaller thanthat of Pecten maximus (L.) (8.0–11.6 L g DW−1 h−1) in laboratorystudies in which low natural plankton concentrations at temperaturesbetween 4.6 and 19.6 °Cwere tested (Strohmeier et al., 2009), and thatof Aequipecten opercularis (10.67 L g DW−1 h−1) measured for 2individual scallops (Riisgård, 2001). In addition, B. vexillum CR wasmuch lower than that of Pinctada margaritifera L. (22 L g DW−1 h−1)in a Pacific atoll (Pouvreau et al., 2000), the highest reported CR in anatural environment.

The CR of bivalves may be related to gill area, which is generallyproportional to the body weight (Riisgård, 1988). However, a highrelative gill area has been observed for the pearl oyster (P.margaritifera) in coral lagoons and interpreted as an adaptation tooligotrophic conditions (Pouvreau et al., 2000). The CR of B. vexillumwas not significantly related to individual biomass or to sexualdevelopment. Our results do not allow drawing inferences about theeffects of these endogenous factors.

CRwas related to SSTwith a clear increase for temperatures higherthan 27 °C. In studies on cold or temperate areas, high temperatureshave been associated with a decline in CR or filtration rate (FR). Forinstance, FR decreases at temperatures over 20 °C for Mytilus edulis(Widdows and Bayne, 1971) and over 19 °C for Crassostrea gigas(Thunberg) (Bougrier et al., 1995). Our results show a differentresponse from tropical bivalves with the highest CR for the maximalSST (28.3 °C) encountered in nature during our experiments. SST was,however, significantly related to most seston variables, which can actas confounding factors. The steep decrease in CR with SPMconcentration suggests a major influence of this variable and the
T
POM, mg trap−1 d−1), particulate organic carbon (POC, mg trap−1 d−1), particulate organicrap−1 d−1) in biodeposits, and biomass of B. vexillum (gAFDW), suspended particulatematterorganic carbon (SPOC, μg L−1), suspended particulate nitrogen (SPN, μg L−1), suspendeds, r is Pearson product–moment correlation, and p is the related probability. The framed cells



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Table 6t6:1

Parameters of the functional linear regressions corresponding to significant correlations (framed cells) in Table 5. Biodeposit-dependent variables: total particulate matter (TPM,mg trap−1 d−1), particulate organic matter (POM, mg trap−1 d−1), particulate organic carbon and nitrogen (POC, PN, mg trap−1 d−1), chlorophyll a (Chl a, μg trap−1 d−1),pheopigments (Pheo, μg trap−1 d−1). Explicative variables: ash-free dry weight biomass of B. vexillum (g AFDW), suspended chlorophyll a (SChl a, μg L−1), and pheopigments(SPheo, μg L−1) in seston. Standard deviations are in parentheses.

t6:2t6:3 AFDW SChl a SPheo

t6:4 Slope Intercept Slope Intercept Slope Intercept

t6:5 TPM 298.56 (17.34) 5.15 129.25 (16.22) −3.49 204.35 (25.01) −9.08t6:6 POM 115.71 (8.30) −1.64 50.09 (6.13) −4.99 79.20 (10.00) −7.16⁎

t6:7 POC 45.55 (3.43) −1.10 19.72 (2.39) −2.42⁎ 31.17 (3.80) −3.27⁎

t6:8 PN 2.88 (0.25) −0.14⁎ 1.25 (0.15) −0.23⁎ 1.97 (0.24) −0.28⁎

t6:9 Chl a 19.96 (1.58) −1.36⁎ 8.64 (1.09) −1.93⁎ 13.66 (1.71) −2.30⁎

t6:10 Pheo 47.07 (3.82) −1.97⁎ 20.38 (2.66) −3.33⁎ 32.22 (4.16) −4.21⁎

⁎ Indicates intercepts significantly different from 0 (pb0.05).t6:11


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related SPOM, SPOC, and SPN. In contrast, CR increased withphotosynthetic pigment concentration. Thus, food quality affects thefeeding behavior of B. vexillum. The highest Chl a used as a microalgalbiomass index stimulates the animal's FR whereas the other sestonvariables have an opposite effect. A similar decrease in CR withincreasing SPM concentration has been observed for Placopectenmagellanicus (Gmelin) (Bacon et al., 1998). This scallop can selectparticles containing Chl a to improve the nutritional value of the

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Fig. 7. RelationshipsamongBractechlamysvexillum individual biomass and (A)biodepositedparticulate organic carbon by unit of biomass (POC) and (B) biodeposited chlorophyll a byunit of biomass (Chl a). The parameters for the fitted curves are given in Table 4.


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often respond to an increase in seston concentration by increasingmucus-coated pseudofeces production (MacDonald et al., 1998), theresponse of B. vexillum can be interpreted as an energy-savingbehavior, even if a mechanism to get rid of excessive particle loads isalso possible. Further information is needed, however, on the ingestionand absorption rates of the species to draw reliable conclusions.

In regionswith high seston level, bivalve growth is very limitedwhenChl a concentration is low, and bivalves may stop suspension feedingwhen the concentration decreases to 0.5–0.9 μg L−1 (Strohmeier et al.,2009). In tropical lagoons, Chl a concentration is often lowbecause of lownutrient concentration (Fichez et al., 2005). The average Chl aconcentration measured during the present study (0.47±0.2 μg L−1)was similar to that previously identified in the SW lagoon (Torréton et al.,2007) and to values obtained in various other coral reef lagoons (seeJacquet et al., 2006, for a review). A very high clearance activity of P.margaritifera in an oligotrophic environment has been associated withthe large relative size of gills and maximal use of the pumping capacityper unit gill area (Pouvreau et al., 1999). In the SW lagoon of NewCaledonia, the b2 μm algal size fraction predominated and mainlyconsisted of Synechococcus (Jacquet et al., 2006). The fraction N2 μmrepresented about 30% of the Chl a in seston. The minimum size ofparticles collected by the pectinid gill is about 5 μm (Beninger and LePennec, 1991); retention efficiency decreases for smaller size seston,being only 15% for 2-μm particles in Argopecten irradians L. (Riisgård,1988) even though particles as small as 1 μm could be retained althoughwith a very lowefficiency (Pouvreau et al., 1999). The clearance rate for P.maximuswasmaximal for average Chl a concentrations (0.3–0.8 μg L−1)and decreased for higher concentrations (Strohmeier et al., 2009). Asimilar pattern was observed for Chlamys farreri (Johnes and Preston)(Hawkins et al., 2001). In the sameway,we found a negative relationshipbetween biodeposition by B. vexillum and Chl a concentration in seston.

4.3. Biodeposition by B. vexillum

Seston was collected once at the onset of each experiment. Sestoncharacteristics are likely to change during the course of 24 h inrelation to hydrodynamic conditions (Iglesias et al., 1998), and short-term scallop biodeposit production may vary accordingly. Sestonconcentration measured at the sampling station was low and close tothe general characteristics of the SW lagoon of New Caledonia (Clavieret al., 1995), which could lead to a dominance of feces in biodeposits(Cranford et al., 2005). However, the threshold for pseudofecesproduction (see Pouvreau et al., 2000, for a review) is generally lowerthan the average SPM concentration measured during our study, andpseudofeces production by B. vexillum is very likely to occur in thelagoon environment. Furthermore, the noticeable concentration of Chla in some sedimented material suggests that biodeposits did notoriginate only from feces. The amount of deposited Chl a in traps waslower than in the reference trap only for young specimens. Thenegative significant relationship between biodeposit production and



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photosynthetic pigments in seston can derive from the oligotrophicstatus of the lagoon. To obtain enough food, organisms must increasethe FR when algae cell concentration is low. This food seeking mayalso account for the relative depletion of Chl a concentration inbiodeposits compared to natural sedimentation.

The biodeposit production of B. vexillum is related to body size. Asimilar relationship has been established for Crassostrea virginica(Gmelin) (Haven and Morales-Alamo, 1966) and Laternula elliptica(King and Broderip) (Ahn, 1993). B. vexillum biodeposit productionper unit biomass is, however, markedly higher for small individuals, ashas already been observed for mussels (Callier et al., 2006; Tsuchiya,1980). Biodeposition of Chl a differed from that of other variables; thelowest values were identified for small individuals, suggesting theirpreferential ingestion rate for algae. This finding can be related to ahigh particle selection by juveniles. The average total B. vexillumbiodeposit production was 10.3 mg g DW−1 h−1, a value in the rangeof feces production by P. margaritifera (6–12 mg g DW−1 h−1) in atolllagoons, to which pseudofeces production (0–6 mg g DW−1 h−1) isadded (Pouvreau et al., 2000). In situ fecal production for the scallop P.magellanicus during a tidal cycle has been identified as distinctlylower, with a maximum rate of 1.2 mg g DW−1 h−1 (Cranford andHargrave, 1994).

As shown by C/N values, organic matter mineralization rate wassimilar between biodeposits and naturally deposited material.However, nutrient content in egested feces may be influenced byscallop digestion and absorption, and pseudofeces bound in mucusproteins may influence nutrient composition. We found no relation-ship between biodeposits and seston concentrations, with theexception of photosynthetic pigments, for which the relationshipwas negative. This relative independence from seston suggests thatparticle retention by B. vexillum does not depend directly on thequantity and quality of suspended particulate matter. Independencefrom seston characteristics may also be a consequence of a selectivefiltration related to suspended particle concentration. The amount ofbiodeposits mainly depends on the individual size, i.e., the volume ofwater filtered through the gills. Different results were obtained for thegastropod Crepidula fornicata (Barillé et al., 2006). A good relationshipbetween the biodeposit production rate and the concentration ofparticulate matter in seston (r2=0.86) was found, but the tested SPMconcentration reached 196 mg L−1, whereas maximal value in thelagoon was 21 mg L−1 and the average value was five times lower.However, the proportion of organic matter was higher in B. vexillumbiodeposits than in naturally sedimentedmaterial. This difference canbe attributable tometabolic fecal loss or to themucus production fromgills, coating the rejected material. Such a phenomenon wouldsupport the hypothesis of pseudofeces production by B. vexillum. Inthe New Caledonia lagoon, available particulate matter did not limit B.vexillum, and biodeposit production mainly depends on the biomassof mollusks without significant seasonal variation.

4.4. Contribution to particulate matter and energy fluxes in the lagoon

The large difference in SPM concentration previously establishedbetween near-bottom water and subsurface (Clavier et al., 1995) andthe high variation in measured seston concentration at the bottomlevel (1.086 to 21.801 mg L−1) indicate that resuspension is a majorfeature in the SW lagoon of New Caledonia. Resuspension mainlyrelies on the influence of wind-waves on sediment (Ouillon et al.,2004) and results in a mixing of benthic material and plankton,making it available to local or remote suspension feeders. Resuspen-sion also derives from biological activity of mobile fauna like foragingcrustaceans or, on the study site, the solitary cnidarianHeteropsammiacochlea (250 ind m−2), which can be moved about by the commensalsipunculid worm Aspidosiphon jukesi Baird (Chardy et al., 1987).Deposition of resuspended fine grain size benthic particles accountedfor more than 80% of the material collected by sediment traps in the


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SW lagoon (Clavier et al., 1995). Resuspended sediment was howevermainly inorganic, with only 10% of organic matter. In nature,individuals of B. vexillum are capable of rapid short-distanceswimming burst to escape predators, but they are standing most ofthe time at the water–sediment interface where they excavateshallow pits in the substrate in order to lie with the left (upper)valve even with the sediment surface. Therefore, biological resuspen-sion of sediment by this bivalve is likely low or moderate. Physicalresuspension of feces and pseudofeces is related to current speed.Widdows and Navarro (2007) found that critical erosion thresholdsfor Cerastoderma edule (L.) biodeposits were recorded at currentvelocities of 15 and 25 cm s−1, for pseudofeces and feces respectively.Currents are generally lower than 10 cm s−1 in the SW lagoon(Douillet et al., 2001) and physical resuspension of biodeposits canalso be regarded as low or moderate. Most of organic carbonbiodeposition remains at the water–sediment interface where it canbe used by benthic micro or macroorganisms.

Feces and pseudofeces result from active collection and transfer ofsuspended particles and therefore increase sedimentation. Biodepo-sits have a high nutritional value (Giles and Pilditch, 2004; Kautskyand Evans, 1987) and show high bacterial activity (Grenz et al., 1990;Kaspar et al., 1985). Biodeposit production increases the flux oforganic carbon to the sediment, and the mineralization of biodepositsincreases sediment respiration and supplies regenerated nutrients tothe overlying water and to benthic primary producers (Dame, 1996).Biodeposit contribution to benthic metabolism can be high in areaswhere filter feeders dominate invertebrate communities (Newell,2004), such as mussel or oyster farming areas (Cranford et al., 2009;Haven and Morales-Alamo, 1966). Less-detailed information isavailable about the contribution of natural populations of bivalvesto water–sediment transfer of particulate material, particularly incoral reef areas (Pouvreau et al., 2000).

Anadara scapha (L.) and B. vexillum are the major filter-feederbivalves in the 1000-km² grey sand bottoms community of the SWlagoon of New Caledonia, with a pooled biomass of 3 g AFDWm−2

(Chardy and Clavier, 1988). A. scapha biodeposition values are stillunknown, but members of the genus can produce large amounts ofbiodeposits (Miranda-Baeza et al., 2006). With an average carbonproduction of 4·103 metric tons year−1 for POC in the grey sandbottoms community, the general contribution of B. vexillum to energyfluxes is modest compared to the POC average natural sedimentationof 223·103 metric tons year−1 (Clavier et al., 1995), or compared tothe carbon benthic primary production of the grey sand bottomscommunity, estimated at 197·103 metric tons year−1 (Clavier andGarrigue, 1999). Therefore, the relative contribution of the species andof bivalves more generally to translocation of carbon from water tosediment is low in grey sand bottoms community. However, if weconsider the station studied, biodeposition of 68 mg C m−2 d−1 by B.vexillum only accounts for 12% of the carbon required for benthiccommunity respiration (Clavier and Garrigue, 1999). Carbon biode-position in places where bivalve density is high can therefore besignificant in the local biogeochemical functioning of the benthos.

Acknowledgements

We thank Pierre-Henri Lecques for assisting in the experimentalwork. We also thank Dr. Tore Strohmeier for his valuable commentson the manuscript. This research was supported by funding from theFrench Institute for Research and Development (IRD) and the FrenchNational Research Agency (ANR) project “CHIVAS”.

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