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Annual microplankton cycles in Villefranche Bay, Ligurian Sea, NW Mediterranean
FERNANDO GÓMEZ* AND GABRIEL GORSKY1
DEPARTMENT OF AQUATIC BIOSCIENCES, THE UNIVERSITY OF TOKYO,
1CNRS/UPMC, LABORATOIRE D’OCÉANOGRAPHIE DE VILLEFRANCHE/MER, UMR

 VILLEFRANCHE-SUR-MER CEDEX, FRANCE *CORRESPONDING AUTHOR: fernando.gomez@fitoplancton.com

-- YAYOI, BUNKYO, TOKYO -, JAPAN AND , OBSERVATOIRE OCÉANOLOGIQUE, BP ,

Abundance and composition of microplankton were studied over a period of 2 years at two depths in Villefranche Bay (Ligurian Sea, NW Mediterranean Sea). Diatoms dominated the microplankton in late spring and autumn, whereas dinoflagellates composed the major part of the microplankton in summer. The silicoflagellate Dictyocha fibula and the diatom Thalassionema frauenfeldii dominated in winter. Ciliates showed low variability throughout the year with the lowest abundance in February and an increase which coincided with the diatom maxima during autumn in both years. In 1998, the spring bloom (in May) was mainly composed of dinoflagellates near the surface and of diatoms in deeper layers. Subsurface diatom maxima were observed in August–September and November. In 1999, diatoms peaked in May both at the surface and at the depth of 50 m. They showed a strong maximum in October. Dinoflagellates and tintinnids showed maxima in early November. Comparisons with previous studies reveal that (i) changes in species composition have not been significant, (ii) the silicoflagellate’s abundance is lower during the present study, (iii) the sequential spring bloom is composed of a pico-nanoplankton bloom in March and microphytoplankton in May, whereas in other western Mediterranean areas the spring microphytoplankton bloom is reported in February and March, (iv) high water transport through the Corsica channel coinciding with low or negative winter values of North Atlantic Oscillation (NAO) index are associated with the anomalous strong development of the spring diatom blooms in the Bay of Villefranche, whereas the usual trend is the lack of or weak development of the spring diatom bloom. This feature may determine the nature and the fate of primary production and the interannual variability in the relative importance of the microbial food web versus the microbial loop.

I N T RO D U C T I O N
The Mediterranean Sea is an oligotrophic semi-enclosed basin with an eastward decrease in productivity (Sournia, 1973). However, the concentrations of nitrate and phosphate are increasing (Béthoux et al., 1998a) and the cultural eutrophication due to the rising demographic impact is becoming an important ecological issue for the Mediterranean coastal zones (UNESCO, 1988; Turley, 1999), along with the effects of the climatic change on the Mediterranean ecosystem (Francour et al., 1994; Bianchi and Morri, 2000). The trophic character of Villefranche Bay varies from oligotrophic [based on chlorophyll or primary production values (Bustillos-Guzmán et al., 1995; Lacroix and Nival, 1998)] to mesotrophic [based on the nitrogen dynamics

and plankton composition (Selmer et al., 1993)]. The food web is predominantly based on the microbial loop (Rassoulzadegan and Sheldon, 1986; Dolan et al., 1995; Thingstad et al., 1998). The waters are dominated by picoand nanoplankton, mainly consisting of cyanobacteria (Synechococcus sp.), nanoflagellates (responsible for the major part of the primary production) and heterotrophic bacteria (Lins da Silva, 1991; Zweifel et al., 1993; Jacquet et al., 1998). Even during the spring and autumn maxima, the abundance of microphytoplankton is low (Hagström et al., 1988; Lins da Silva, 1991; Ferrier-Pagès and Rassoulzadegan, 1994) in comparison to the Gulf of Lions ( Jacques, 1969) and to the Gulf of Marseille (Travers, 1971) where river input and wind forcing are more important. The studied area is influenced by the Ligurian current

Journal of Plankton Research 25(4), © Oxford University Press; all rights reserved

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[or Northern current; sensu (Millot 1999)], characterized by a low chlorophyll concentration (Goffart et al., 1995). This current is part of a basin-wide cyclonic circulation involving Modified Atlantic Water at the surface and the Levantine Intermediate Water at depth (Béthoux et al., 1988; Millot, 1999). It is formed from the merging of two independent currents flowing northwards on both sides of Corsica Island, the Western Corsica Current (WCC) and the Eastern Corsica Current (ECC or Tyrrhenian Current). In Villefranche Bay, the wind stress can modify the circulation pattern (Nival and Corre, 1976). During calm periods, the surface current circulates from the south (open sea) towards the north of the Bay and subsurface currents tend to circulate southwards near the bottom. During the events of eastward wind, surface waters circulate towards the open sea and subsurface waters are upwelled, allowing deeper waters to ascend the slope. Deeper waters reach the surface when the strong eastward wind regime is persistent (Nival et al., 1975). As oligotrophic areas are very sensitive to environmental variations, their monitoring is essential for the evaluation of the long-term changes in the community structure. The Bay of Villefranche, regularly sampled since 1957 (Etienne et al., 1991), is especially interesting due to the high number of studies performed in this area which facilitates comparison. The present work focuses on (i) the comparison of composition, seasonal cycle and dynamics of microplankton succession in 1998 and 1999 with previous studies, and (ii) the evaluation of the relationship between

hydrographic conditions and the causes of the interannual variability.

METHOD
Study site and sampling
The Bay of Villefranche is located in the northern part of the Ligurian Sea (NW Mediterranean Sea, Figure 1). Sampling was performed at a permanent coastal station called Point B. The station is located at the entrance of the Bay of Villefranche (43°41Ј10ЉN, 7°19Ј00ЉE, water column depth ~80 m) which is open towards the sea and is not protected much from the wind (Nival and Corre, 1976). Conductivity–Temperature–Depth (CTD) and water sampling were performed weekly between 10 and 11 a.m. Water column profiles of temperature and salinity were recorded with a Seabird SBE25 CTD. A stratification index calculated as the average density difference m–1 of the 0–80 m depth layer (␴t units m–1) was used to characterize the degree of water column stability. Phytoplankton analysis was carried out weekly during late spring and autumn when changes were more apparent, and fortnightly in winter and summer. Samples for microplankton analysis were collected using Niskin bottles at the surface (1 m) and at 50 m depth and fixed with acid Lugol’s solution. In the laboratory, 100 ml subsamples were allowed to settle (48 h) in composite chambers. The entire chamber bottom was always examined using the Utermöhl technique. Organisms that

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could not be identified to the species level were assigned only to genera, for example diatoms as Navicula or dinoflagellates as Gymnodinium, Gyrodinium or Amphidinium and suprageneric groups such as naked ciliates. Dominance and diversity indices were computed using taxa identified at species level only. Although the counting error of the abundance estimations was generally lower than 20% (Lund et al., 1958), it increased in the cases of low microplankton concentrations. Colonies (usually diatoms) were considered as one unit for abundance estimations and small cells mainly belonging to the flagellate group were not considered. It should be noted that these small cells deteriorate in fixed samples and that their abundance may be underestimated by the methodology used (Hewes et al., 1984). This can also affect the non-loricated ciliate’s quantification (Stoecker et al., 1994).

temperature was 13.37°C (minimum 12.84°C). In late summer 1999, temperature values in the surface layer were exceptionally high (24ºC isotherm reached 27 m depth). In summer 1998, temperatures were lower and the 24ºC isotherm extended to 18 m only (Figure 2). According to Romano et al., temperatures in 1999 were the highest in the NW Mediterranean since 1995 (Romano et al., 2000). The stratification of the water column started earlier in 1998 than in 1999 (Figure 2), probably due to the lower winter temperatures in 1999. Mixing events, induced by wind during the stratification period, resulted in lower values of the stratification index.

Seasonal cycles of the main microplankton groups
Total microplankton abundance and the main groups considered (dinoflagellates, diatoms, ciliates; nonloricated ciliates and tintinnids; and silicoflagellates) at the surface and 50 m depth are shown in Figure 3. The average microphytoplankton abundance during the period studied was ~1 ind ml–1. In both years, the microphytoplankton abundance remained low from December to March with a rapid increase in May, in late summer (August–September) and in October and November, when diatoms were the taxa responsible for abundance fluctuations. In 1998 the spring maximum was not observed at the surface as usual, but only at 50 m depth. Dinoflagellates were concentrated mostly in the surface layer. The highest abundance was recorded in May 1998 (up to 4 ind ml–1). It coincided with the deep (50 m) diatom peak. The comparison of the observed abundance between the surface and 50 m depth showed that dinoflagellates are mainly surface-dwelling organisms. However, the decrease in the surface abundance values was followed by an increase in the abundance at 50 m depth (November–December, 1998; Figure 3). Generally, this group displayed low abundance during winter. The annual cycle of silicoflagellates fluctuated inversely when compared with the annual cycle of the bulk microplankton, with maximal surface values in winter. From spring to autumn they were detected only under the thermocline (50 m depth). In winter 1999 their abundance was higher than in 1998, coinciding with lower temperatures observed during this year. The abundance of naked ciliates ranged between 0.2 and 1 ind ml–1 throughout the year. The lowest values were observed in winter, especially in February. The surface abundance of ciliates increased during the spring and during autumnal diatom maxima (more pronounced in 1999). The distribution of tintinnids showed large fluctuations with some of the maxima occurring shortly after a diatom peak (August 1998, 50 m depth; January 1999, 1 m). In autumn 1999, an increase in the diatoms 50 m

Data analysis
The phytoplankton species diversity was estimated according to Shannon’s formula (HЈ = –Σpi log2 pi, where p = n1/N, n1 = number of individuals of one species and N the total number of individuals) (Margalef, 1967). Microphytoplankton species dominance was calculated according to the formula ␦ = 100 (n1 + n2)/N, where ␦ is the dominance index equal to the percentage of the total standing stock contributed by the two most important species (n1 + n2) and N the total cell concentration (Hulburt, 1963). For the statistical analyses, the data for microplankton abundance were converted to log10(x + 1) and the whole dataset was reduced to 27 (surface) and 23 (50 m depth) microphytoplankton taxa by merging poorly represented species or congeneric species, which are supposed to have similar ecophysiological characteristics. The inclusion of supraspecific and heterogeneous taxa has probably resulted in overestimated similarity values. A hierarchical flexible cluster analysis with b = –0.3 on a Euclidean matrix of distance (Legendre and Legendre, 1998) was used in order to define groups of co-occurrent microplankton species. The statistical program PASSTEC developed at the Observatoire Océanologique of Villefranche was used for calculations (Ibañez and Etienne, 1998).

R E S U LT S
Hydrographic conditions
The average water column temperature from January to April 1998 was 14.13°C (minimum 13.47°C) during 1998, whereas in 1999 for the corresponding period the average

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Fig. 2. Seasonal variations of the temperature and density at the mouth of the Villefranche Bay (NW Mediterranean Sea) from January 1998 to January 2000. The stratification index was calculated as the average density difference m–1 of the 0–80 m depth layer each 5 m (␴t units m–1). Note the different scale of each group.

Stability index -1 ␴ t units m

deep (early September) was followed 2 weeks later by an increased abundance of tintinnids at the same depth.

Composition, dominance, diversity and succession
More than one 150 taxa (Table I), mainly composed of 25 genera of dinoflagellates (80 species) and of 37 diatom genera (65 species), were identified. The dominant bloom-forming diatoms are Dactyliosolen fragilissimus,

Leptocylindrus danicus, L. minimus, Guinardia striata and Pseudo-nitzschia spp. Regarding species number, the genus Chaetoceros, with nine species, is the most important, followed by Leptocylindrus (four species) and Guinardia (three species) (Table I). Within the dinoflagellates, the Ceratium group with 18 species is the most important, followed by the genus Protoperidinium (nine species), Dinophysis (seven species) and Prorocentrum (six species) (Table I). In addition to these taxonomic groups, the silicoflagellate

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Table I: Taxonomic composition of microplankton in the Villefranche Bay
Dinoflagellates Dinoflagellates Diatoms

Table I: Continued
Diatoms

Podolampas spinifera Okamura Amphidinium spp. Amphisolenia bidentata Schröder Asterodinium libanum Abboud-Abi Saab Ceratium arietinum Cleve Ceratium candelabrum (Ehr) Stein Ceratium carriense Gourret Ceratium contortum (Gourret) Cleve Ceratium declinatum (Karsten) Jörg Ceratium extensum (Gourret) Cleve Ceratium furca (Ehr) Clap et Lach Ceratium fusus (Ehr) Dujardin Ceratium hexacanthum Gourret Ceratium horridum (Cleve) Gran C. horridum var.buceros (Zach) Sournia Ceratium limulus Gourret Ceratium longirostrum Gourret Ceratium macroceros (Ehr) Vanh Ceratium massiliense (Gourret) Karsten Ceratium pentagonum Gourret Ceratium pulchellum Schröder Ceratium setaceum Jörg Ceratium symmetricum Pavillard Ceratium teres Kofoid Ceratium tripos (Müller) Nitzsch Ceratocorys armata (Schütt) Kof Ceratocorys gourretii Paulsen Ceratocorys horrida Stein Cladopyxis brachiolata Stein Cochlodinium spp. Corythodinium constrictum (Stein) Taylor Corythodinium elegans (Pav) Taylor Corythodinium frenguellii (Rampi) Balech C. tessellatum (Stein) Loeb and Loeb III Dinophysis acuminata Clap and Lach Dinophysis caudata Saville-Kent Dinophysis cf. punctata Jörg Dinophysis sacculus Stein Dinophysis schroederi Pav Dinophysis tripos Gourret Dinophysis spp. Erythropsidinium sp. Gonyaulax spp. Gonyaulax verior Sournia Gymnodinium spp. Gyrodinium spp. Heterodinium sp. Histioneis spp. Micracanthodinium sp. Ornithocercus heteroporus Kofoid Ornithocercus magnificus Stein Oxytoxum longiceps Schiller Oxytoxum milneri Murray and Whitting Oxytoxum scolopax Stein Phalacroma doryphorum Stein P. rotundatum (Clap and Lach) Kof and Mich Podolampas elegans Schütt Podolampas palmipes Stein Amphora spp. Asterionellopsis glacialis (Cast) Round Asteromphalus sp. Asteromphalus flabellatus (Bréb) Grev Bacteriastrum delicatulum Cleve Bacteriastrum elegans Pav Bacteriastrum hyalinum Lauder Bleakeleya notata (Grun) Round Cerataulina pelagica (Cleve) Hendey Chaetoceros affinis Lauder C. compressus Lauder C. curvisetus Cleve C. dadayi Pavillard C. danicus Cleve C. decipiens Cleve C. densus Cleve C. didymus Ehr C. lorenzianus Grun C. messanensis Castracane C. peruvianus Brightwell C. rostratus Lauder C. tortissimus Gran Coscinodiscus spp. Cylindrotheca closterium (Ehr) Reim and Lew Dactyliosolen fragilissimus (Bergon) Hasle Detonula pumila (Castracane) Gran Diploneis spp. Ditylum brightwellii (West) Grun Grammatophora spp. Guinardia delicatula (Cleve) Hasle Guinardia flaccida (Cast) Per Guinardia striata (Stolt) Hasle Hemiaulus hauckii Grun Hemiaulus sinensis Grev Hemidiscus cuneiformis Wallich Leptocylindrus danicus Cleve L. mediterraneus (Per) Hasle Leptocylindrus minimus Gran Licmophora spp. Melosira sp. Navicula spp. Nitzschia longissima (Brév ex Kutz) Ralfs in Prich Odontella mobiliensis (Bailey) Grun Paralia sulcata (Ehr) Cleve Pleurosigma spp. Proboscia alata (Brightwell) Sundström Pseudo-nitzschia delicatissima ‘complex’ Pseudo-nitzschia sp. ‘Nitzschia seriata complex‘ Pseudosolenia calcar-avis (Schultze) Sundström Rhabdonema adriaticum Kütz Rhizosolenia bergonii Per Rhizosolenia hebetata Bailey f. hebetata Rhizosolenia hebetata f. semispina (Hensen) Gran Rhizosolenia imbricata Brightwell Rhizosolenia styliformis Brightwell Skeletonema costatum (Grev) Cleve Striatella unipunctata (Lyngbye) Agardh Other phytoplankton groups Dictyocha fibula Ehr Dictyocha speculum Ehr Halosphaera viridis Schmitz Phaeocystis sp. Pterosperma sp. Rhizosomonas setigera (Pav) Patterson et al. Prorocentrum compressum (Bailey) Abé Prorocentrum dentatum Stein Prorocentrum micans Ehr Prorocentrum rotundatum Schiller Prorocentrum triestinum Schiller Protoperidinium claudicans (Paulsen) Balech P. crassipes (Kof) Balech P. depressum (Bailey) Balech P. diabolus (Cleve) Balech P. divergens (Ehr) Balech P. minusculum Pav (=Minuscula bipes Lebour) P. oceanicum (vanHöffen) Balech P. pellucidum (Schütt) Balech P. steinii (Jörg) Balech Protoperidinium spp. Pseliodinium vaubanii Sournia Pyrocystis elegans Pav Pyrocystis fusiformis Wyville-Thomp. Pyrocystis lunula Schütt Pyrocystis obtusa Pav Pyrocystis robusta Kofoid Scaphodinium mirabile Margalef Scrippsiella cf. trochoidea (Stein) Balech ex Loeb III Pronoctiluca spinifera (Lohm) Schiller Prorocentrum balticum (Lohm) Loeb III

Synedra spp. Thalassionema frauenfeldii (Grun) Hallegraeff Thalassionema nitzschioides Grun ex Mereschkowsky Thalassiosira spp. Thalassiothrix longissima Cl and Grun

Ciliates Ascamphelliella sp. Amphorides quadrilineata Clap and Lach Climacocylis sp. Codonella spp. Codonellopsis spp. Dadayiella ganymenes Entz Dictyocysta elegans Jörg Dictyocysta spp. Eutintinnus fraknoii Daday Eutintinnus lusus-undae Entz Epiplocylis spp. Favella sp. Mesodinium rubrum Lohman Rhabdonella spiralis Fol Salpingella acuminata Jörg Salpingella decurtata Jörg Salpingella spp. Steenstrupiella steenstrupii Clap and Lach Stenosemella nivalis (Meun) Kof and Campb Stenosemella ventricosa (Clap and Lach) Jörg Tintinniosis campanula (Ehr) Dad Tintinniosis radix (Imhof) Brandt Tintinniosis spp. Tintinnus inquilinus (Muller) Schrank Undella claparedei Entz Undella marsupialis (Brandt) Kof and Campb Xystonella treforti Daday Xystonellopsis cymatica Brandt

Dictyocha fibula was the most abundant. For ciliates, we were not able to make precise taxonomic identifications, although the observed species consisted mostly of Halteria sp., Laboea sp., Lohmanniella sp. and Strombidium-like cells (Rassoulzadegan, 1977; Rassoulzadegan and Sheldon, 1986). The dominant phytoplankton species and dominance index for both years are summarized in Figure 4. In winter 1998, the dominant species were Thalassionema frauenfeldii (which dominated until July at 50 m depth) and D. fibula. Both species generated the highest degree of dominance ( January at 50 m depth). Towards the summer, these species were progressively replaced in the surface layer by an assemblage composed of dinoflagellates. In May 1998, this assemblage was composed of

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T. frauenfeldii / D. fibula D. fibula / T. frauen. T. frauenf/ D. brightwelli T. frauenf/ D. fibula D. fibula / T. frauenfeldii D. fibula / T. frauenfeldii D. fibula / T. frauen. T. frauen./ Cyl. closterium T. frauenfeldii / Gymnod. T. frauenfeldii / Gymnod. T. frauenfeldii / Gymnod. D. fragil./P. seriata D. fragil./Chaeto. sp Gym./ T. frauenf. Gym./ T. frauenf. Gym./ Cyl. closterium Ch.peruvian./Gym. Ch.peruvian./Gym. D. fibula/T. frauenfeld. D. fibula/T. frauen. T. frauenf./ D. fibula D. fibula / Navicula D. fibula/T. frauenfeldii D. fibula / C. tripos D. fibula/Gymnodinium T. frauenfeld/ D. fragilis. T. frauenf./Gyrod. L. danicus / D. fragilisimus L. danicus / T. frauenfeldii L. minimus/Ch.curvisetus P. seriata/ Bacteriastrum P. seriata/ Ch. affinis L. danicus / D. fragil. G. striata/Bacteriastrum P. seriata/ L. danicus T. frauenfeldii / D. fibula T. frauenfeldii / D. fibula T. frauenf./ D. fibula

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J D. fibula / T. frauenfeldii D. fibula / Gyrodinium F D. fibula / Gymnodinium M D. fibula / Gymnodinium D. fibula / Gymnodinium A C. furca / P. balticum M C. furca / P. micans Gonyaulax / Gyr. J Gyrod. / Gonyaulax Gymnod. / Gyrod. J Gymno. / P. micans Gymno. / S. trochoidea A Gymno. / P. micans Gymno. / B. elegans S Gym. / T. frauenf. O Ch. peruvian./ Gymno. N Ch. peruv. / T. frauenf. D. fibula / T. frauenf. D Gymno./ T. frauenf. J D. fibula / Gymn. D. fibula / Gymnodinium F D.fib./ Pleurosigma M D. fibula / Gymnodinium A P. alata / D. fragilissimus M C. furca / P. micans sp/ Gyr. J Chaetoceros P. micans / C. furca

T. frauenfeldii / D. fibula

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P. micans / C. furca Gyr. / L. danicus Gym. / Gyr. C. furca / D. rotundatum L. minimus / D. fragilis. Chaetoceros sp/ G. striata T. frauenf. / Chaetoceros sp

T. frauenfeldii / D. fibula T. frauen./ C. furca T. frauenfeldii / P. seriata T. frauenf./ P. seriata

0 20 40 60 80 %

0 20 40 60 80 100 %

Fig. 4. Dominance (%) index and dominant species at the surface (1 m) and 50 m depth in 1998 and 1999 in the Bay of Villefranche.

Ceratium furca (2.4 ind ml–1) and Prorocentrum micans, followed by P. balticum, Ceratium fusus and also other typical red tide species (Dinophysis sacculus/acuminata, Dinophysis caudata, Mesodinium rubrum and Gonyaulax sp.). Two weeks after this Ceratium furca/Prorocentrum micans maximum, the diversity index reached its highest value (3.7 bits ind–1) for this year in an assemblage dominated by Gyrodinium and Gonyaulax. In late spring the assemblage of Ceratium and Prorocentrum was progressively replaced by small dinoflagellates belonging to the genera Gymnodinium and Gyrodinium. In early autumn the water column was dominated by Chaetoceros spp. Towards the winter, D. fibula and T. frauenfeldii dominated again. In winter 1999, D. fibula was the dominant species until April, followed by Gymnodinium sp. and by tychoplanktonic diatom species such as Navicula sp. and Pleurosigma sp. In early May, Proboscia alata and Dactyliosolen fragilissimus bloomed at the surface. In summer C. furca and P. micans dominated. At the end of August an assemblage composed of D. fragilissimus, Leptocylindrus danicus and L. minimus dominated at both depths. In September, Guinardia striata, Chaetoceros spp. and Bacteriastrum delicatulum were the dominant species. Finally in November 1999 (after the autumnal maximum), the abundance of tintinnids (mainly Amphorides quadrilineata and Salpingella spp.) became high. They were associated with the highest value of microphytoplankton diversity index (4.26 bits ind–1).

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1998

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In surface waters, spring and autumnal peaks coincided with the increase of the diversity index values (cf. May 1998). At 50 m depth, the abundance maxima generally occurred following an increase of the diversity index (Figure 5). Several groups of species can be separated in the dendrogram (Figure 6). At the surface one group is composed mainly of dinoflagellates. This group also includes diatoms such as Hemiaulus hauckii and Bacteriastrum elegans. Another group is composed from the most frequent species of bloom-forming diatoms. Two groups appear to be clearly separated. One composed of Ceratium furca and Prorocentrum micans is present all the year but only with episodic maxima from spring to autumn. The second, composed of T. frauenfeldii and D. fibula, is inversely related to the rest of the groups, showing the higher abundance in winter. At 50 m depth, the results are similar but the separation between dinoflagellates and diatoms is less clear. Ceratium furca and P. micans appear to be grouped with the rest of the dinoflagellates, whereas T. frauenfeldii and D. fibula are again clearly separated from the rest of the groups.

DISCUSSION
Microphytoplankton distribution
During winter, D. fibula and non-colonial individuals of T. frauenfeldii dominated the phytoplankton community. They are frequent components of the Mediterranean

deep flora (Kimor et al., 1987). Nival (Nival, 1965) observed a seasonal pattern of D. fibula in the Bay of Villefranche with the highest abundance in winter. This tendency is usual in many Mediterranean areas with a high development of silicoflagellates in winter, when they form an important fraction of the phytoplankton population (Tolomio et al., 1999). During the rest of the year the silicoflagellates are only observed in subsurface waters (Travers and Travers, 1968). Our results show that silicoflagellates reach their highest abundance when the winter temperature is low, as it was in 1999. In the Bay of Villefranche, Nival (Nival, 1965) reported a maximum of 5.5 silicoflagellate cells ml–1 and Rassoulzadegan (Rassoulzadegan, 1979) 1.8 cells ml–1. More recently, Lins da Silva (Lins da Silva, 1991) reported a maximum value of 0.4 cell ml–1 in March. This abundance is similar to the maximum value observed (also in March) during this study (0.45 cell ml–1). This long-term decrease in silicoflagellate abundance could be related to the progressive increase of water temperature in the Ligurian Sea since 1960 (Astraldi et al., 1995) and to the decrease of the wind force (Fromentin and Ibañez, 1994). At the permanent station Point B, Bougis and collaborators reported temperatures lower than 12ºC in February–March during the 1960s (Bougis et al., 1968). However, during this study the temperatures were always higher than 12.8ºC. Among the diatoms, T. frauenfeldii has been cited as the most frequent winter species in Monaco (Pavillard, 1913, 1937), Banyuls (Dangeard, 1932) and in other Mediterranean areas (Friligos and Gotsis-Skretas, 1987).

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Diversity index (bits cell-1)

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3 4 5 6

Euclidean distance

1

2

50 m

P. pellucidum

P. micans

C. pentagonum

Ch. peruvianus

G. flaccida

C. furca

P. alata

L. danicus

B. elegans

Pseudo-nitzschia sp

C. fusus

L. fragilissimus

D. rotundatum

Ch. curvisetus

Ch. tortissimus

T. frauenfeldii

P. spinifera

H. hauckii

G. striata

Ch. decipiens

L. minimus

0.8 1.6 2.4 3.2 4.0

C. candelabrum

surface

1m

Gonyaulax sp

B. elegans

Ch. peruvianus

P. pellucidum

P. depresum

D. caudata

G. flaccida

Chaetoceros sp

Ch. curvisetus

T. frauenfeldii

Pseudo-nitzschia sp

Chaetoceros sp

D. rotundatum

L. minimus

L. fragilissimus

Ch. decipiens

Ch. tortissimus

P. diabolus

H. hauckii

L. danicus

Fig. 6. Dendrogram of the Euclidean distance between the abundant species at the surface (1 m) and 50 m depth.

Thalassionema frauenfeldii is a perennial species developing successfully in winter when other diatoms are absent (Travers, 1971). The presence of quasi-perennial species decreases the specific diversity index in the surface waters between January and March of 1998 (Figure 5). During spring, diatom blooms were associated with a temporal transition between mixed and stratified conditions generated by the increase in temperature (usually in May). In summer, following the diatom peak, dinoflagellates became the dominant component of the microphytoplankton community (Figure 4). In the NW Mediterranean, according to the review by Velásquez and Cruzado (Velásquez and Cruzado, 1995), diatoms and dinoflagellate species represented, respectively, 51% and 36% of the total phytoplankton species. In Villefranche, our data showed that diatoms and dinoflagellate species represented, respectively, 43% and 52% of the total specific composition of microphytoplankton. For example in 1973, diatoms represented 8%, while dinoflagellates composed 28% of the average phytoplankton biovolume (Rassoulzadegan, 1979). In the Gulf of Genova, Bernhard and Rampi (Bernhard and Rampi, 1967) reported 31% of diatoms and 48% of dinoflagellates. The

C. carriense

relative abundance of dinoflagellates in the NW Mediterranean microphytoplankton community augments from west eastwards. The composition and abundance of dinoflagellates in this study did not show significant differences from previous observations in the area. The most frequent species are Prorocentrum micans, Ceratium furca and Ceratium fusus (Halim, 1960; Rassoulzadegan, 1979). This assemblage is also typical in other Mediterranean areas (Revelante and Gilmartin, 1976). In the surface waters of the Bay of Villefranche, C. fusus and P. micans were associated with diatom maxima and also with periods of dinoflagellate dominance. As a consequence, the position of both species in the dendrogram does not appear associated with the diatom or the dinoflagellate group (Figure 6). According to Halim (Halim, 1960), the dinoflagellates reach their maximal values in summer (up to 4 ind ml–1) with peaks in May and June–July. The usual presence of congeneric assemblages of Ceratium (C. furca, C. fusus and C. pentagonum), Prorocentrum (P. micans, P. balticum, P. triestinum and P. rotundatum) and Gonyaulax spp., organisms with apparently the same nutritional requirements, reveals that the interspecific competence is not clear. Even genera

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P. micans

G. striata

C. fusus

D. fibula

C. furca

P. alata

D. fibula

F. GÓMEZ AND G. GORSKY

ANNUAL MICROPLANKTON CYCLES IN NW MEDITERRANEAN SEA

such as Ceratium or Prorocentrum, considered as autotrophic because of the presence of photosynthetic pigment (Lessard and Swift, 1986), have often shown a phagotrophic behaviour (Kimor, 1981; Stoecker et al., 1997). Dominant species in the Bay of Villefranche such as C. fusus and C. furca are heterotrophic (Mikaelyan and Zavyalova, 1999; Smalley et al., 1999). Given that they succeed peaks of phototrophs, they may feed on the organic matter pool. Dinoflagellates are typically surface-dwelling organisms; however, the subsurface maximum observed after the decrease in the surface abundance could be related to the motility of this group. Vertical migration may allow the uptake of new nutrients in deeper waters during the period of surface nutrient depletion (Eppley et al., 1968; Blasco, 1978). During the summer and after the C. furca/P. micans peak, small dinoflagellates of the genera Gyrodinium and Gymnodinium became dominant. Ibañez and Rassoulzadegan (Ibañez and Rassoulzadegan, 1977) reported that Gymnodinium reaches its maximum abundance towards mid-June. The dominance of these species in nutrient-impoverished environments could be related to their trophic versatility. Many species of Gyrodinium and Gymnodinium present an unclear trophic behaviour (Novarino et al., 1997). During summer in the Bay of Villefranche, despite the presence of some dinoflagellate species capable of forming red tides, this phenomenon was not detected. It is probable that the poor nutritional conditions prevailing in the Bay with low nitrate concentration (<0.3 µM; Ferrier-Pagès and Rassoulzadegan, 1994) and the advection of water masses prevent the development of red tides. The abundance of dinoflagellates did not reach the values normally reported in red tide events ( Jacques and Sournia, 1978–79). However, episodes of massive algae proliferation, such as Chattonella, have been reported in the Bay of Villefranche during summer (Trégouboff, 1961). During the stratified period in Villefranche Bay, the Ceratium–Prorocentrum assemblages developed first, followed by the Gyrodinium–Gymnodinium group. Only a few species of diatoms such as Hemiaulus hauckii and Pseudosolenia calcar-avis are associated with a complete stratification of the water column (Margalef, 1967). The presence of genera such as Ornithocercus or Pyrocystis revealed the complete development of this stage until a new environmental fluctuation (usually wind) interrupted the succession. The presence of these large diatom and dinoflagellate species can be explained by the presence of symbiotic cyanobacteria (Taylor, 1982; Kimor et al., 1992). During summer stratified conditions, strong wind events were associated with diatom (fucoxanthin) maxima

(Bustillos-Guzmán et al., 1995) and with episodic development of Dactyliosolen, Leptocylindrus and Rhizosolenia species (i.e. August 1999; Figure 2). Phytoplankton may respond to environmental changes by rapid population increases in the superficial layer. Bustillos-Guzmán et al. (BustillosGuzmán et al., 1995) reported diatom maxima associated with wind events in August 1992. This environmental forcing can inject nutrients into the nutrient-limited surface layer and generate high biomass. Experimental studies showed that primary production in the Bay was limited by phosphorus in summer (Thingstad et al., 1998) and in early autumn (Berland et al., 1973; Zweifel et al., 1993). However, late summer events could overlap with the autumnal bloom (such as in 1998).

Microzooplankton distribution
The specific seasonal distribution of naked ciliates is not detailed in this study. Their population dynamics may be more related to the prey (bacteria or detritus) distribution than to the changes in the physical and nutritional conditions. The average abundance of naked ciliates in this study was ~1 ind ml–1, in the range of the abundance reported in previous studies (Rassoulzadegan and Gostan, 1976; Rassoulzadegan, 1977). There is no clear relationship between the tintinnid abundance and the seasonal cycle (Vaqué et al., 1997), chlorophyll concentrations (Dolan et al., 1999) or water column stability (Cariou et al., 1999). We reported tintinnid maxima in December–January and November (Figure 3) after the breakdown of the thermocline (Figure 2). Temperature-dependent distribution of different tintinnid species has been reported in Villefranche (Posta, 1963) and in other areas (Koray and Özel, 1983; Abboud-Abi Saab, 1989). Size and abundance of available prey may also influence the abundance and community structure of tintinnids. In this study, large tintinnids such as Amphorides quadrilineata (0.16 cell ml–1) and Salpingella sp. succeeded the microplankton maxima (after the autumnal bloom of 1999). This tendency is related to the nutritional interdependence with several groups of phytoplankton (Kimor and Golandsky-Baras, 1981) and to the availability of large sized prey (Bernard and Rassoulzadegan, 1990, 1993). An increase in large ciliate (>50 µm) populations ingesting microplankton cells (Smetacek, 1981) is observed following the nanoplankton maxima in Villefranche (Lins da Silva, 1991), peaking in May and November (Bernard and Rassoulzadegan, 1994). The isolated peaks in the abundance of tintinnids and the sudden appearance of predetermined species can be

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explained by the seasonal recruitment from cysts as a consequence of a particular environmental condition (Paranjape, 1987; Kamiyama and Tsujino, 1996).

Seasonal cycles of phytoplankton
The seasonal pattern of phytoplankton in temperate coastal areas usually shows two annual peaks, in late winter and autumn in open coastal ecosystems, and in spring and late summer in enclosed coastal ecosystems (Cebrián and Valiela, 1999). When compared with other Mediterranean coastal areas (Table II), it seems that the bimodal annual cycle is not the usual trend in the Ligurian Sea and Gulf of Lions (Table III). In the coastal Gulf of Lions (Banyuls), unimodal cycles were reported with a marked maximum in January, whereas in the Gulf of Genova the maximum was reported in September–October. In the Gulf of Lions, the phytoplankton abundance can be related to the strong winter northerly winds that generate intense mixing processes associated with high primary production (MEDOC Group, 1970; Lefevre et al., 1997). Near Banyuls, chlorophyll peaks reached 3 µg l–1 ( Jacques, 1970) and 6 µg l–1 (Ibarra, 1981). In the Gulf of Marseille, the phytoplankton is seasonally influenced by the Rhône river outflow plume (Leveau et al., 1990; Soto et al., 1993). Eastward, these phenomena can be less relevant and the trophic character is gradually more oligotrophic. In Villefranche Bay, the average chlorophyll a concentration

is 0.32 µg l–1 (ranging between 0.17 and 0.64 µg l–1) and is considered within the range of oligotrophic values (Bustillos-Guzmán et al., 1995). Villefranche Bay can represent an intermediate position between the adjacent zones (Gulf of Lions–Gulf of Genova). The higher biomass usually observed in autumn suggests that the seasonal cycle in Villefranche displays a transition from bimodal to unimodal annual cycles with the maximum around October–November. At the beginning of the last century, Pavillard reported the lack of the spring diatom bloom in some years during his study near Monaco, whereas the autumnal maxima were more prominent. This feature surprised him after his studies on the usual spring bloom in the Gulf of Lions (Pavillard, 1937). In Table IV, we have compiled studies on seasonal cycles performed in the Bay of Villefranche. The phytoplankton biomass has been evaluated using different estimators such as chlorophyll, fluorescence, particles (using a Coulter particle counter) and the abundance of several taxonomic groups. Generally the spring bloom of phytoplankton in Villefranche develops in two phases. In March–April, a first maximum is composed of pico-nanoplanktonic autotrophs followed by a second maximum in May composed of microphytoplankton. In some cases the peak of large diatoms was not observed. Bustillos-Guzmán et al. compared the hydrographic conditions between a typical diatom bloom year [1986, in (Claustre et al., 1989)]

Table II: Some examples of phytoplankton peak periods in coastal areas of the Mediterranean Sea
Location Months of peaks Year Reference

Venice Lagoon Saronicos Bay Saronicos Bay Saronicos Bay Israel coast Lebanon coast Gulf of Naples Gulf of Naples Castellón Coast Castellón Coast Castellón Coast Castellón Coast Catalonian Coast Catalonian Coast Catalonian Coast Catalonian Coast Catalonian Coast

Mar, Aug Mar, Oct Apr, Oct Apr, Oct Mar, Aug, Sep, Oct Mar, Oct Mar, May, Oct May, Oct Feb Feb, Nov Mar Feb, Dec Mar, Nov Mar, Nov Mar, Oct Feb, Aug Jan, Aug

1988–89 1967 1975–1976 1984–85 1983 1973–75 1955 1984–1988 1956–57 1957, 1961 1962 1970 1966 1965–66 1978–79 1992 1993

Tolomio et al., 1999 Ignatiades, 1969 Ignatiades, 1984 Siokou-Frangou, 1996 Azov, 1986 Lakkis and Novel-Lakkis, 1981 De Angelis, 1956 Zingone et al., 1995 Herrera and Margalef, 1957 San Feliu and Muñoz, 1975 San Feliu and Muñoz, 1975 San Feliu and Muñoz, 1975 Margalef and Castellví, 1967 Margalef and Ballester, 1967 Estrada,1980 Mura et al., 1996 Mura et al., 1996

The biomass descriptors are the chlorophyll a concentration or the phytoplankton abundance.

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F. GÓMEZ AND G. GORSKY

ANNUAL MICROPLANKTON CYCLES IN NW MEDITERRANEAN SEA

Table III: Phytoplankton peak periods in coastal areas of the French–Italian coast of the NW Mediterranean Seaa
Location Months of peaks Biomass estimator Year Reference

Gulf of Lions Gulf of Lions Gulf of Lions Gulf of Lions Gulf of Marseille Monaco coast Monaco coast Monaco coast Monaco coast Monaco coast Gulf of Genova Gulf of Genova Gulf of Genova Gulf of Genova
aVillefranche

Jan, Mar, Sep, Oct Feb/Sep, Oct Jan Jan Mar, May/Sep, Nov Oct–Nov Nov Feb, Aug, Oct Feb, Oct May, Oct Jun, Sep, Dec Jun, Aug, Dec Sep Oct

Nanoplankton volume Chlorophyll Chlorophyll Chlorophyll Phytoplankton Phytoplankton Phytoplankton Nanoplankton volume Chlorophyll Phytoplankton Microphytoplankton Microphytoplankton Microphytoplankton Organic matter

1937 1965–68 1973 1977 – 1910 1913 1937 1965 1965 1959 1962 1965 1980–81

Bernard, 1938 Jacques, 1970 Neveux et al., 1975 Ibarra, 1981 Travers, 1962, 1973 Pavillard, 1937 Pavillard, 1937 Bernard, 1938 Kane, 1967 Kane, 1967 Bernhard and Rampi, 1967 Bernhard and Rampi, 1967 Bernhard et al., 1969 Fabiano et al., 1984

Bay is not included.

and a non-typical diatom bloom year [1992, in (BustillosGuzmán et al., 1995)]. Air temperatures were colder in 1986 than in 1992 (Fromentin and Ibañez, 1994). Bustillos-Guzmán et al. reported that the diatom bloom in 1986 occurred when the water column began to stabilize while in 1992 this occurred prior to stabilization [late April, in (Claustre et al., 1988]. In March–April 1992 the irradiance increased when the water column was not stabilized. The authors concluded that the conditions of 1992 favoured the development of nanoflagellates versus diatoms. When the light and stabilization became optimal for diatoms, the nutrients had already been depleted by nanoflagellates. In an oligotrophic environment this phenomenon may generate a significant interannual variability. In more eutrophic areas, nutrient concentrations could support a regular spring diatom bloom despite the nanoflagellate population growth.

Interannual variability of microphytoplankton: comparison with previous studies
The Mediterranean climate is under the continental influence of Europe and Asia. The Sahara Desert and the Atlantic Ocean favour thermal anomalies and changes of circulation patterns (Maheras et al., 1999). The Mediterranean Sea is very sensitive to variations in heat or the water budget (Béthoux et al., 1999). Changes in the eastern and western basin are frequently out of phase as a consequence of different processes that affect the climate of the two basins (Reddaway and Bigg, 1996).

In southern Europe and the Mediterranean Sea, positive values of the North Atlantic Oscillation (NAO) are related to dry winter conditions, whereas negative NAO years are associated with wet winters (Hurrell, 1995). In the Ligurian Sea, precipitation is associated with the increase of wind stress (Andersen and Prieur, 2000) and with a rapid increase of nutrients in the surface waters (Martin et al., 1989). The variable intensity of the Corsica channel transport is correlated with the winter NAO index. Higher transports are associated with negative NAO and less water is transported in the Corsica channel during positive NAO years (Vignudelli et al., 1999). According to Astraldi et al. the long-term survey of the Corsica channel circulation provides the first evidence of the NAO influence on the interannual variability of the physical and biological processes in the Western Mediterranean (Astraldi et al., 1999). The Corsica channel transport that feeds the Ligurian current (Northern current) presents a strong seasonality. The prominent interannual variability of the Ligurian current (Le Floch, 1963; Béthoux et al., 1982; Astraldi and Gasparini, 1992; Sammari et al., 1995) may influence the nature of the spring phytoplankton bloom intensity on the northern coast of the Ligurian Sea. Years with high transports (and low or negative winter NAO indices) are associated with a remarkable development of the spring diatom bloom. When the intensity of the circulation is lower (positive NAO index) the level of the spring bloom is low (Table V). During productive years the spring bloom develops earlier (late March) than during

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Table IV: Interannual variability of biomass peaks (using different estimators) in the Bay of Villefranche
Biomass peaks Biomass descriptor Period Reference

Apr, Aug Apr, Nov May–Jun May, Aug, Sep Mar Apr/Mar Mar, Aug Mar, Sep Feb, Mar, Apr, Jun May May Apr, Aug, Oct Mar, Apr, May May (and Mar) Mar, May May Mar, May May May, Aug, Nov May, Sep, Nov

Cyanobacteria Picoflagellates Nanoflagellates Microphytoplankton Microphytoplankton Cyanobacteria Nanoplankton Cyanobacteria Nanophytoplankton Microphytoplankton Microphytoplankton Chlorophyll Fluorescence Particles (3–10 µm) Fluorescence/chlorophyll Particles (3–10 µm) Fluorescence Particles (2–15 µm) Microphytoplankton Microphytoplankton

1986 1986 1986 1986 1987 1988/1989 1989 1990 1990 1990 Nov’90–Jun’91 1992 Feb–Jun, 1995 Feb–Jun, 1995 Oct’95–May’96 Oct’95–May’96 Jan–Jun, 1999 Jan–Jun, 1999 1998 1999

Lins da Silva, 1991 Lins da Silva, 1991 Lins da Silva, 1991 Lins da Silva, 1991 Fernex et al., 1996 Bernard and Rassoulzadegan, 1994 Bernard and Rassoulzadegan, 1994 Ferrier-Pagès and Rassoulzadegan, 1994 Ferrier-Pagès and Rassoulzadegan, 1994 Ferrier-Pagès and Rassoulzadegan, 1994 Selmer et al., 1993 Bustillos-Guzmán et al., 1995 Therond, 1995 (unpublished data) Therond, 1995 (unpublished data) Senior, 1996 (unpublished data) Senior, 1996 (unpublished data) Mandleberg, 1999 (unpublished data) Mandleberg, 1999 (unpublished data) This study This study

the periods of lower production (May). In the English Channel, Irigoien et al. reported that the proportion of diatoms during the phytoplankton spring bloom was significantly correlated with the NAO index, whereas the total phytoplankton carbon and the timing of the spring bloom were not correlated (Irigoien et al., 2000). During the period 1985–1987 in the Corsica channel, transport was estimated to be the highest in the last 15 years, up to 2.0 Sverdrup (Astraldi et al., 1995; Vignudelli et al., 2000). During this period the winter air temperatures were the lowest since 1957 (Fromentin and Ibañez, 1994; Santoreli et al., 1995). During the 1980s, the highest levels of precipitation on the Mediterranean basin were recorded between 1985 and 1987 (Boukthir and Barnier, 2000). For example, in Sardinia the precipitation level recorded in 1985 was the highest since 1964 (Delitala et al., 2000). According to the model of Lacroix and Nival, based on meteorological data from 1985, the beginning of the stratification and the development of the spring diatom bloom took place at the end of March near the coast of Nice (Lacroix and Nival, 1998). Claustre et al. and Lins da Silva reported the development of an extraordinary diatom bloom in the Bay of Villefranche in 1986 (Claustre et al., 1988/1989; Lins da Silva, 1991). From a study between 1987 and 1989, Fernex et al. pointed out

high phytoplankton abundance in late March 1987, followed by high copepod concentration, whereas during 1988 and 1989 this latter sequence did not occur (Fernex et al., 1996). In the Gulf of Genova an exceptional increase in the zooplankton abundance was also reported in 1987 (Licandro and Ibañez, 2000). After the 1985–1987 period, winter transport in the Corsica channel decreased (higher values of the NAO index) and the intensity of the spring bloom declined. The analysis of the long-term temperature and salinity variations in the western Mediterranean shows an increasing trend in the deep water temperature (Béthoux et al., 1990; Francour et al., 1994; Roether et al., 1996; Krahmann and Schott, 1998). Thermal anomalies, as in late summer of 1999 (Figure 2), are associated with unprecedented events of mass mortality of sessile invertebrates in the western Mediterranean Sea (Perez et al., 2000; Romano et al., 2000). From the analysis of sapropels, the increase in the freshwater input appears to be correlated with the increase in the abundance of diatoms in the Mediterranean Sea (Kemp et al., 1999). The decrease in precipitation due to the NAO (Hurrell, 1995) and the damming of rivers (Milliman, 1997) could be associated with the progressive decrease in diatom abundance in the Mediterranean Sea, and conversely the increase in nanoflagellates and dinoflagellates over

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ANNUAL MICROPLANKTON CYCLES IN NW MEDITERRANEAN SEA

Table V: Spring phytoplankton bloom intensity (evaluated as high or low) Villefranche Bay compared with the winter transport (Sverdrups = 1 ϫ 106 m3 s–1) in the Corsica Channel reported by Astraldi et al. (Astraldi et al., 1995) and Vignudelli et al. (Vignudelli et al., 2000) and the winter NAO (North Atlantic Oscillation) index [(Hurrell, 1995) and further articles]
Year Flux NAO Bloom References

1962 1963 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1998 1999

– – 2.0 1.6 1.3 0.9 – – 0.6 0.4 0.2 0.8 1.0 –

–2.38 –3.60 –0.63 0.50 –0.75 0.72 5.08 3.96 1.03 3.28 2.67 3.03 0.72 1.70

High High High High High Low Low Low Low Low – – Low Low

Nival, 1965 Nival, 1965 Lacroix and Nival, 1998 Claustre et al., 1988; Lins da Silva, 1991 Fernex et al., 1996 Fernex et al., 1996 Fernex et al., 1996 Ferrier-Pagès and Rassoulzadegan, 1994 Selmer et al., 1993 Bustillos-Guzmán et al., 1995 – – This study This study

diatoms or silicoflagellates. There is a progressive increase in nitrate and phosphate concentrations in the Mediterranean basin, whereas the silicate concentrations appear to be stationary (Béthoux et al., 1998a). This could have consequences on the decrease of the relative importance of diatoms (Turley, 1999). The modification of the N:Si and Si:P molar ratios could induce a shift from the diatom communities to flagellate-dominated communities, which do not need silicate in their cell structures (Officer and Ryther, 1980, Escaravage et al., 1996). This shift may influence the community structure, favouring filter feeding zooplankton (Fernex et al., 1996; Gorsky and Fenaux, 1998; Gorsky et al., 1999) over selective feeders such as copepods (Berggreen et al., 1988). Thus, the consequence of anthropogenic and climatic changes such as the greenhouse effect and the decrease in the freshwater input (Rohling and Bryden, 1992; Béthoux and Gentili, 1996; Béthoux et al., 1998b; Boukthir and Barnier, 2000) may contribute to a progressive change in the structure of the pelagic food web in the Mediterranean Sea.

help during his stay at Villefranche. We thank P. Nival for helpful comments and suggestions on the manuscript. We acknowledge the support from the European Commission’s Marine Science and Technology Programme (MAST III-EURAPP) under contract MAS3-CT98-0161. This work was undertaken in the framework of the French PNEC ART 4 program. F.G. acknowledges financial support by the Spanish Ministry of Science and Technology and by the European Commission (IBC2-CT-2001-80002).

REFERENCES
Abboud-Abi Saab, M. (1989) Distribution and ecology of tintinnids in the plankton of Lebanese coastal waters. J. Plankton Res., 11, 203–222. Andersen, V. and Prieur, L. (2000) One month study in the open NW Mediterranean Sea (DYNAPROC experiment, May 1995): overview of the hydrobiological structures and effects of wind events. Deep Sea Res. I, 47, 397–422. Astraldi, M. and Gasparini, G. P. (1992) The seasonal characteristics of the circulation in the north Mediterranean basin and their relationship with the atmospheric–climatic conditions. J. Geophys. Res., 97, 9531–9540. Astraldi, M., Bianchi, C. N., Gasparini, G. P. and Morri, C. (1995) Climatic fluctuations, current variability and marine species distribution: a case study in the Ligurian Sea (north-west Mediterranean). Oceanol. Acta, 18, 139–149. Astraldi, M., Balopoulos, S., Candela, J., Font, J., Gacic, M., Gasparini, G. P., Manca, B., Theocharis, A. and Tintoré, J. (1999) The role of

AC K N O W L E D G E M E N T S
We thank S. Dallot for providing CTD data from the SOMLIT program, I. Palazzoli for technical assistance and the skippers D. Betti and J.-Y. Carval for collecting the samples at Point B. F.G. wishes to thank B. Gasser, P. Licandro, J. C. Molinero, M. Picheral and S. Souissi for

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