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Annual microplankton cycles in Villefranche Bay, Ligurian Sea, NW Mediterranean



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.

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

1976). During calm periods. The station is located at the entrance of the Bay of Villefranche (43°41Ј10ЉN. 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. 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. The entire chamber bottom was always examined using the Utermöhl technique. In the laboratory. is especially interesting due to the high number of studies performed in this area which facilitates comparison. Figure 1). regularly sampled since 1957 (Etienne et al. 7°19Ј00ЉE. and fortnightly in winter and summer. the wind stress can modify the circulation pattern (Nival and Corre. seasonal cycle and dynamics of microplankton succession in 1998 and 1999 with previous studies.m. Millot. METHOD Study site and sampling The Bay of Villefranche is located in the northern part of the Ligurian Sea (NW Mediterranean Sea. 1975).. 1999). and (ii) the evaluation of the relationship between hydrographic conditions and the causes of the interannual variability. water column depth ~80 m) which is open towards the sea and is not protected much from the wind (Nival and Corre. Deeper waters reach the surface when the strong eastward wind regime is persistent (Nival et al. 100 ml subsamples were allowed to settle (48 h) in composite chambers.. 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. 1991). 1976). characterized by a low chlorophyll concentration (Goffart et al.JOURNAL OF PLANKTON RESEARCH VOLUME  NUMBER  PAGES ‒  [or Northern current.. During the events of eastward wind. The present work focuses on (i) the comparison of composition. It is formed from the merging of two independent currents flowing northwards on both sides of Corsica Island.5 Km 7° 18' 41" Fig. the Western Corsica Current (WCC) and the Eastern Corsica Current (ECC or Tyrrhenian Current).. Phytoplankton analysis was carried out weekly during late spring and autumn when changes were more apparent. sensu (Millot 1999)]. allowing deeper waters to ascend the slope. Sampling was performed at a permanent coastal station called Point B. The Bay of Villefranche. 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. Conductivity–Temperature–Depth (CTD) and water sampling were performed weekly between 10 and 11 a. 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. The NW Mediterranean Sea with an inset showing the Bay of Villefranche-sur-Mer and the sampling station Point B (43°41Ј10ЉN. Organisms that France Sp n ai Italy Nice Monaco Villefranche/mer N Villefranche sur Mer 15 m 43° 41' 57" Mediterranean Sea 10 0 0m Cannes 43°30' 10 60 120 m 7° a Se n a uri g i L 00 m 7° 30' 0. 7°19Ј00ЉE). surface waters circulate towards the open sea and subsurface waters are upwelled. Water column profiles of temperature and salinity were recorded with a Seabird SBE25 CTD. 1. 1988. In Villefranche Bay.  m 1 30 m 5 m Station Marine B .

the decrease in the surface abundance values was followed by an increase in the abundance at 50 m depth (November–December. whereas in 1999 for the corresponding period the average  ..F.47°C) during 1998.37°C (minimum 12. probably due to the lower winter temperatures in 1999. Generally. The comparison of the observed abundance between the surface and 50 m depth showed that dinoflagellates are mainly surface-dwelling organisms. temperature was 13. It coincided with the deep (50 m) diatom peak. GÓMEZ AND G. with maximal surface values in winter. 1994). 1 m).84°C). The annual cycle of silicoflagellates fluctuated inversely when compared with the annual cycle of the bulk microplankton. 1963). R E S U LT S Hydrographic conditions The average water column temperature from January to April 1998 was 14. and silicoflagellates) at the surface and 50 m depth are shown in Figure 3. ciliates. an increase in the diatoms 50 m Data analysis The phytoplankton species diversity was estimated according to Shannon’s formula (HЈ = –Σpi log2 pi. Mixing events. 1998) was used in order to define groups of co-occurrent microplankton species. This can also affect the non-loricated ciliate’s quantification (Stoecker et al. For the statistical analyses. diatoms. nonloricated ciliates and tintinnids. The statistical program PASSTEC developed at the Observatoire Océanologique of Villefranche was used for calculations (Ibañez and Etienne. The lowest values were observed in winter. which are supposed to have similar ecophysiological characteristics. Dinoflagellates were concentrated mostly in the surface layer. coinciding with lower temperatures observed during this year. when diatoms were the taxa responsible for abundance fluctuations. it increased in the cases of low microplankton concentrations. 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. In late summer 1999. The abundance of naked ciliates ranged between 0. 1984).3 on a Euclidean matrix of distance (Legendre and Legendre. in late summer (August–September) and in October and November. The surface abundance of ciliates increased during the spring and during autumnal diatom maxima (more pronounced in 1999). In autumn 1999. 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. GORSKY ANNUAL MICROPLANKTON CYCLES IN NW MEDITERRANEAN SEA could not be identified to the species level were assigned only to genera. Figure 3).13°C (minimum 13. but only at 50 m depth. Gyrodinium or Amphidinium and suprageneric groups such as naked ciliates. In winter 1999 their abundance was higher than in 1998.. Seasonal cycles of the main microplankton groups Total microplankton abundance and the main groups considered (dinoflagellates. n1 = number of individuals of one species and N the total number of individuals) (Margalef. However. Colonies (usually diatoms) were considered as one unit for abundance estimations and small cells mainly belonging to the flagellate group were not considered. where p = n1/N. especially in February. induced by wind during the stratification period.. temperatures were lower and the 24ºC isotherm extended to 18 m only (Figure 2). temperatures in 1999 were the highest in the NW Mediterranean since 1995 (Romano et al. 2000). The highest abundance was recorded in May 1998 (up to 4 ind ml–1). 1998. From spring to autumn they were detected only under the thermocline (50 m depth). for example diatoms as Navicula or dinoflagellates as Gymnodinium. Dominance and diversity indices were computed using taxa identified at species level only. 50 m depth. In summer 1998. Although the counting error of the abundance estimations was generally lower than 20% (Lund et al.. resulted in lower values of the stratification index. The stratification of the water column started earlier in 1998 than in 1999 (Figure 2). In 1998 the spring maximum was not observed at the surface as usual. the microphytoplankton abundance remained low from December to March with a rapid increase in May.2 and 1 ind ml–1 throughout the year. January 1999. 1998). 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. The distribution of tintinnids showed large fluctuations with some of the maxima occurring shortly after a diatom peak (August 1998. Microphytoplankton species dominance was calculated according to the formula ␦ = 100 (n1 + n2)/N. According to Romano et al. The inclusion of supraspecific and heterogeneous taxa has probably resulted in overestimated similarity values. temperature values in the surface layer were exceptionally high (24ºC isotherm reached 27 m depth).. 1967). this group displayed low abundance during winter. A hierarchical flexible cluster analysis with b = –0. 1958). The average microphytoplankton abundance during the period studied was ~1 ind ml–1. In both years.

Composition. Guinardia striata and Pseudo-nitzschia spp. In addition to these taxonomic groups. followed by Leptocylindrus (four species) and Guinardia (three species) (Table I). minimus. L.02 0. mainly composed of 25 genera of dinoflagellates (80 species) and of 37 diatom genera (65 species). Seasonal variations of the temperature and density at the mouth of the Villefranche Bay (NW Mediterranean Sea) from January 1998 to January 2000. followed by the genus Protoperidinium (nine species). the Ceratium group with 18 species is the most important.JOURNAL OF PLANKTON RESEARCH VOLUME  NUMBER  PAGES ‒  1998 1999 JFMAMJJASOND JFMAMJJASOND 0 Depth (m) 20 40 60 80 0 Temperature (°C) Depth (m) 20 40 60 t Density (␴ ) 0. the silicoflagellate  .04 0. 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). diversity and succession More than one 150 taxa (Table I). 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. 2.03 0. dominance. Leptocylindrus danicus. Within the dinoflagellates.01 0 JFMAMJJASOND JFMAMJJASOND 1998 1999 Fig. Dinophysis (seven species) and Prorocentrum (six species) (Table I). is the most important. with nine species. were identified. Regarding species number. the genus Chaetoceros. The dominant bloom-forming diatoms are Dactyliosolen fragilissimus.

Rhizosomonas setigera (Pav) Patterson et al. Pseliodinium vaubanii Sournia Pyrocystis elegans Pav Pyrocystis fusiformis Wyville-Thomp. Dadayiella ganymenes Entz Dictyocysta elegans Jörg Dictyocysta spp.. didymus Ehr C. GÓMEZ AND G. and Strombidium-like cells (Rassoulzadegan. oceanicum (vanHöffen) Balech P. Histioneis 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. Asterionellopsis glacialis (Cast) Round Asteromphalus sp. fibula. rostratus Lauder C. Gyrodinium spp. Prorocentrum compressum (Bailey) Abé Prorocentrum dentatum Stein Prorocentrum micans Ehr Prorocentrum rotundatum Schiller Prorocentrum triestinum Schiller Protoperidinium claudicans (Paulsen) Balech P. pellucidum (Schütt) Balech P. this assemblage was composed of  . trochoidea (Stein) Balech ex Loeb III Pronoctiluca spinifera (Lohm) Schiller Prorocentrum balticum (Lohm) Loeb III Synedra spp. danicus Cleve C. Micracanthodinium sp. we were not able to make precise taxonomic identifications. Nitzschia longissima (Brév ex Kutz) Ralfs in Prich Odontella mobiliensis (Bailey) Grun Paralia sulcata (Ehr) Cleve Pleurosigma spp. Corythodinium constrictum (Stein) Taylor Corythodinium elegans (Pav) Taylor Corythodinium frenguellii (Rampi) Balech C. crassipes (Kof) Balech P. Cylindrotheca closterium (Ehr) Reim and Lew Dactyliosolen fragilissimus (Bergon) Hasle Detonula pumila (Castracane) Gran Diploneis spp. divergens (Ehr) Balech P. 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. densus Cleve C. curvisetus Cleve C. horridum var. Pterosperma sp. Codonella spp. Favella sp. GORSKY ANNUAL MICROPLANKTON CYCLES IN NW MEDITERRANEAN SEA Table I: Taxonomic composition of microplankton in the Villefranche Bay Dinoflagellates Dinoflagellates Diatoms Table I: Continued Diatoms Podolampas spinifera Okamura Amphidinium spp. For ciliates. hebetata Rhizosolenia hebetata f.F. 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. these species were progressively replaced in the surface layer by an assemblage composed of dinoflagellates. messanensis Castracane C. Eutintinnus fraknoii Daday Eutintinnus lusus-undae Entz Epiplocylis spp. Erythropsidinium sp. Ornithocercus heteroporus Kofoid Ornithocercus magnificus Stein Oxytoxum longiceps Schiller Oxytoxum milneri Murray and Whitting Oxytoxum scolopax Stein Phalacroma doryphorum Stein P. Pyrocystis lunula Schütt Pyrocystis obtusa Pav Pyrocystis robusta Kofoid Scaphodinium mirabile Margalef Scrippsiella cf. 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. Rassoulzadegan and Sheldon. tortissimus Gran Coscinodiscus spp. Heterodinium sp. 1986). Lohmanniella sp. rotundatum (Clap and Lach) Kof and Mich Podolampas elegans Schütt Podolampas palmipes Stein Amphora spp. diabolus (Cleve) Balech P. Both species generated the highest degree of dominance ( January at 50 m depth). lorenzianus Grun C. dadayi Pavillard C. Navicula spp. Melosira sp. depressum (Bailey) Balech P. mediterraneus (Per) Hasle Leptocylindrus minimus Gran Licmophora spp.. Codonellopsis spp. decipiens Cleve C. 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. Mesodinium rubrum Lohman Rhabdonella spiralis Fol Salpingella acuminata Jörg Salpingella decurtata Jörg Salpingella spp. In winter 1998. Towards the summer. compressus Lauder C. Thalassiothrix longissima Cl and Grun Ciliates Ascamphelliella sp. minusculum Pav (=Minuscula bipes Lebour) P. Proboscia alata (Brightwell) Sundström Pseudo-nitzschia delicatissima ‘complex’ Pseudo-nitzschia sp. The dominant phytoplankton species and dominance index for both years are summarized in Figure 4. ‘Nitzschia seriata complex‘ Pseudosolenia calcar-avis (Schultze) Sundström Rhabdonema adriaticum Kütz Rhizosolenia bergonii Per Rhizosolenia hebetata Bailey f. Thalassionema frauenfeldii (Grun) Hallegraeff Thalassionema nitzschioides Grun ex Mereschkowsky Thalassiosira spp. Gonyaulax spp. steinii (Jörg) Balech Protoperidinium spp. although the observed species consisted mostly of Halteria sp. In May 1998. Amphorides quadrilineata Clap and Lach Climacocylis sp. peruvianus Brightwell C. tessellatum (Stein) Loeb and Loeb III Dinophysis acuminata Clap and Lach Dinophysis caudata Saville-Kent Dinophysis cf. 1977. Gonyaulax verior Sournia Gymnodinium spp. Ditylum brightwellii (West) Grun Grammatophora spp. punctata Jörg Dinophysis sacculus Stein Dinophysis schroederi Pav Dinophysis tripos Gourret Dinophysis spp. the dominant species were Thalassionema frauenfeldii (which dominated until July at 50 m depth) and D. Laboea sp.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. 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.

fibula T. frauenf. Chaetoceros spp. frauenf. furca / P. fibula / T. Proboscia alata and Dactyliosolen fragilissimus bloomed at the surface. micans Gymno. seriata/ L. frauenfeldii D. closterium Ch. D./Gyrod. frauen. Mesodinium rubrum and Gonyaulax sp. micans / C. striata/Bacteriastrum P./P. balticum. seriata 0 20 40 60 80 % 0 20 40 60 80 100 % Fig. 4. J Gymno. sp Gym. followed by P. fragilissimus. frauenf.1 Diatoms 3 2 9 colonies ml-1 Naked ciliates 1 0.fib. T. frauenfeldii D. furca T. closterium T. fragil. G. frauenf/ D. furca and P. fragilis. fibula J F M A M J J A S O N D J F M A M J J A S O N D P. balticum M C.4 ind ml -1 0. frauenf. frauenfeldii / Gymnod.JOURNAL OF PLANKTON RESEARCH VOLUME  NUMBER  PAGES ‒  1998 10 8 6 1999 1998 1999 J FMAMJJASONDJFMAMJJASOND JFMAMJJASOND JFMAMJJASOND 1 m Microplankton total 50 m Silicoflagellates 0.peruvian. D. frauenfeldii dominated again. brightwelli T./ D./ T. striata T. In late spring the assemblage of Ceratium and Prorocentrum was progressively replaced by small dinoflagellates belonging to the genera Gymnodinium and Gyrodinium. frauen. fibula/T. J Gyrod. Ceratium furca (2. affinis L. frauenfeldii D. / Gonyaulax Gymnod.5 1 0 0 Dinoflagellates 1 0.3 0./ C. D. / Gyr. fragil. / Chaetoceros sp T. fibula / T. In summer C.curvisetus P. minimus / D. fibula/Gymnodinium T. frauenfeldii / D./Chaeto./ Cyl. minimus/Ch. / P. frauenf. Ch. danicus / D. Abundance (ind ml–1) of total microplankton. D Gymno. rotundatum L. seriata T. fibula / Gymnodinium A P. Leptocylindrus danicus and L. the abundance of tintinnids (mainly Amphorides quadrilineata and Salpingella spp. fibula / Gymnodinium F D. fibula D. frauenfeldii / Gymnod.  . fibula T./Gym.) became high. silicoflagellates. fibula / Gymnodinium A C. T. fibula / C. danicus Gym.7 bits ind–1) for this year in an assemblage dominated by Gyrodinium and Gonyaulax. fragil. frauenf. furca T. micans Gymno. micans dominated. / B. D. fibula / T. C. non-loricated ciliates and tintinnids at the surface and 50 m depth during 1998 and 1999 in the Bay of Villefranche.5 0 5 ind ml-1 Tintinnids 0. / T. micans sp/ Gyr. In early autumn the water column was dominated by Chaetoceros spp. fibula and T. Two weeks after this Ceratium furca/Prorocentrum micans maximum./ Pleurosigma M D. J D. fibula was the dominant species until April. fibula 1 9 9 8 1 9 J 9 A 9 S O N D J D.1 0./Gym./ P.2 4 Abundance (ind ml-1) 2 0 0. frauenfeld/ D. fibula / Navicula D. frauenf. J Chaetoceros P.4 ind ml–1) and Prorocentrum micans.26 bits ind–1)./ Gymno. danicus T. frauenfeld. Surface 0 20 40 60 80 % 50 m depth 0 20 40 60 80 100 % T. In early May. Gym. frauenfeldii / D. fragilis. fibula / Gymn. frauen. T. elegans S Gym. fibula / T. T. frauenf. In winter 1999. They were associated with the highest value of microphytoplankton diversity index (4. Gym. fibula D. micans / C. fibula / Gymnodinium M D. frauenf. frauenfeldii / P. Ceratium fusus and also other typical red tide species (Dinophysis sacculus/acuminata. peruvian. D. fibula / Gyrodinium F D. L.05 J F M A M J J A S O N DJ F M A M J J A S O N D 0 J F M A M J J A S O N DJFMAMJJASOND Fig. alata / D. D. tripos D. danicus / D. Dominance (%) index and dominant species at the surface (1 m) and 50 m depth in 1998 and 1999 in the Bay of Villefranche. dinoflagellates./ T. followed by Gymnodinium sp. frauenf. frauenfeldii D. frauenfeldii L. fibula D. trochoidea A Gymno. seriata/ Bacteriastrum P. In September. minimus dominated at both depths. fibula/T. peruv.45 ind ml -1 0./ Cyl. Finally in November 1999 (after the autumnal maximum). fragilisimus L./ T. frauenfeldii / D. frauenfeldii / D.peruvian. and Bacteriastrum delicatulum were the dominant species. O Ch. D. furca / P.). fibula / T. danicus / T. seriata D. and by tychoplanktonic diatom species such as Navicula sp. Towards the winter. micans Gonyaulax / Gyr. fibula / Gymnodinium D. the diversity index reached its highest value (3. N Ch. frauenfeldii / Gymnod. furca / D. furca / P. Chaetoceros sp/ G. and Pleurosigma sp. fibula T. / S. diatoms. frauenf. / Gyrod. Dinophysis caudata. / P. 3. frauenfeldii / D. / L. / T. Guinardia striata. frauen. fibula / T./ D. fragilissimus M C. At the end of August an assemblage composed of D. fibula/T. frauenf/ D. T. furca Gyr. seriata/ Ch. T. frauen.

May 1998). frauenfeldii has been cited as the most frequent winter species in Monaco (Pavillard. Lins da Silva (Lins da Silva. More recently. Diversity (bits ind–1) and abundance of total microplankton in the Bay of Villefranche at the surface and 50 m depth in 1998 and 1999.8ºC. This group also includes diatoms such as Hemiaulus hauckii and Bacteriastrum elegans. 1937). Nival (Nival.5 2 1. 1987). 1965) reported a maximum of 5. This abundance is similar to the maximum value observed (also in March) during this study (0.. Our results show that silicoflagellates reach their highest abundance when the winter temperature is low.. GORSKY ANNUAL MICROPLANKTON CYCLES IN NW MEDITERRANEAN SEA 1998 5 4 3 2 Microplankton abundance J F M A M J J A S O N D J F M A M J J A S O N D 1999 4.F. 1994). In the Bay of Villefranche.8 cells ml–1. 1913. During the rest of the year the silicoflagellates are only observed in subsurface waters (Travers and Travers.5 silicoflagellate cells ml–1 and Rassoulzadegan (Rassoulzadegan. fibula in the Bay of Villefranche with the highest abundance in winter. Ceratium furca and P. fibula are again clearly separated from the rest of the groups.5 3 2. 1968). 1932) and in other Mediterranean areas (Friligos and Gotsis-Skretas. 1999). fibula.5 3 2. GÓMEZ AND G. during this study the temperatures were always higher than 12. 1965) observed a seasonal pattern of D. Among the diatoms... Another group is composed from the most frequent species of bloom-forming diatoms. In surface waters. whereas T. Banyuls (Dangeard. when they form an important fraction of the phytoplankton population (Tolomio et al. At the permanent station Point B. At the surface one group is composed mainly of dinoflagellates. At 50 m depth. spring and autumnal peaks coincided with the increase of the diversity index values (cf. as it was in 1999. frauenfeldii dominated the phytoplankton community. frauenfeldii and D. At 50 m depth. showing the higher abundance in winter. Several groups of species can be separated in the dendrogram (Figure 6). 1968). DISCUSSION Microphytoplankton distribution During winter. D.45 cell ml–1). 5. T. 1979) 1. However.5 1 50 m 4 3. One composed of Ceratium furca and Prorocentrum micans is present all the year but only with episodic maxima from spring to autumn. 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. They are frequent components of the Mediterranean deep flora (Kimor et al. the abundance maxima generally occurred following an increase of the diversity index (Figure 5). fibula and non-colonial individuals of T. composed of T. micans appear to be grouped with the rest of the dinoflagellates.  Diversity index (bits cell-1) . 1991) reported a maximum value of 0. frauenfeldii and D. 1987). This tendency is usual in many Mediterranean areas with a high development of silicoflagellates in winter. Nival (Nival. The second.5 2 1m 1 0 4 3 2 1 0 J F M A M J J A S O N D J F M A M J J A S O N D 1. Two groups appear to be clearly separated.5 1 Fig. is inversely related to the rest of the groups.4 cell ml–1 in March. Bougis and collaborators reported temperatures lower than 12ºC in February–March during the 1960s (Bougis et al. the results are similar but the separation between dinoflagellates and diatoms is less clear.5 4 3.

pentagonum Ch. 1976). elegans Ch. danicus Fig. according to the review by Velásquez and Cruzado (Velásquez and Cruzado. fusus D. danicus B. curvisetus Ch.0 C. curvisetus T. decipiens Ch. pellucidum P. following the diatom peak. For example in 1973. Bernhard and Rampi (Bernhard and Rampi. minimus 0. minimus L. 1979).8 1. triestinum and P. balticum. During spring. diatom blooms were associated with a temporal transition between mixed and stratified conditions generated by the increase in temperature (usually in May). 1979). flaccida Chaetoceros sp Ch. organisms with apparently the same nutritional requirements. In the NW Mediterranean. while dinoflagellates composed 28% of the average phytoplankton biovolume (Rassoulzadegan. The most frequent species are Prorocentrum micans. The presence of quasi-perennial species decreases the specific diversity index in the surface waters between January and March of 1998 (Figure 5). rotundatum Ch. our data showed that diatoms and dinoflagellate species represented. candelabrum surface 1m Gonyaulax sp B. In summer. hauckii L. 1971). elegans Pseudo-nitzschia sp C. fusus and P. 1960. alata D. Dendrogram of the Euclidean distance between the abundant species at the surface (1 m) and 50 m depth. tortissimus T.4 3. carriense relative abundance of dinoflagellates in the NW Mediterranean microphytoplankton community augments from west eastwards. spinifera H. Prorocentrum (P. tortissimus P. The C. hauckii G. Ceratium furca and Ceratium fusus (Halim. Rassoulzadegan. pellucidum P. P. The usual presence of congeneric assemblages of Ceratium (C. micans. In the surface waters of the Bay of Villefranche. diatoms and dinoflagellate species represented. This assemblage is also typical in other Mediterranean areas (Revelante and Gilmartin. fragilissimus D.. the position of both species in the dendrogram does not appear associated with the diatom or the dinoflagellate group (Figure 6). P.2 4. micans were associated with diatom maxima and also with periods of dinoflagellate dominance. caudata G. 1995). furca P. frauenfeldii P. micans C. As a consequence. fusus L. C. furca. 51% and 36% of the total phytoplankton species. alata L. frauenfeldii Pseudo-nitzschia sp Chaetoceros sp D. 1967) reported 31% of diatoms and 48% of dinoflagellates. peruvianus G. respectively. respectively. depresum D. dinoflagellates became the dominant component of the microphytoplankton community (Figure 4). fibula . fibula C. pentagonum). Even genera  P. In the Gulf of Genova. fragilissimus Ch. diatoms represented 8%. diabolus H. decipiens L. In Villefranche. micans G. According to Halim (Halim. The composition and abundance of dinoflagellates in this study did not show significant differences from previous observations in the area. striata C. rotundatum) and Gonyaulax spp. fusus and C. Thalassionema frauenfeldii is a perennial species developing successfully in winter when other diatoms are absent (Travers. C. 6. striata Ch.6 2.JOURNAL OF PLANKTON RESEARCH VOLUME  NUMBER  PAGES ‒  3 4 5 6 Euclidean distance 1 2 50 m P. flaccida C. reveals that the interspecific competence is not clear. 43% and 52% of the total specific composition of microphytoplankton. rotundatum L. 1960). the dinoflagellates reach their maximal values in summer (up to 4 ind ml–1) with peaks in May and June–July. peruvianus P. furca P.

Many species of Gyrodinium and Gymnodinium present an unclear trophic behaviour (Novarino et al. 1967).. the subsurface maximum observed after the decrease in the surface abundance could be related to the motility of this group. Leptocylindrus and Rhizosolenia species (i.16 cell ml–1) and Salpingella sp. succeeded the microplankton maxima (after the autumnal bloom of 1999). 1989). Dinoflagellates are typically surface-dwelling organisms.. There is no clear relationship between the tintinnid abundance and the seasonal cycle (Vaqué et al.. 1978). Bustillos-Guzmán et al. 1981. The dominance of these species in nutrient-impoverished environments could be related to their trophic versatility. Figure 2). Stoecker et al. However. Dominant species in the Bay of Villefranche such as C. large tintinnids such as Amphorides quadrilineata (0. 1995) and with episodic development of Dactyliosolen.. 1981) is observed following the nanoplankton maxima in Villefranche (Lins da Silva.. have often shown a phagotrophic behaviour (Kimor. This tendency is related to the nutritional interdependence with several groups of phytoplankton (Kimor and Golandsky-Baras.. 1994) and the advection of water masses prevent the development of red tides. However. Kimor et al. 1990. Phytoplankton may respond to environmental changes by rapid population increases in the superficial layer. 1998) and in early autumn (Berland et al. Ibañez and Rassoulzadegan (Ibañez and Rassoulzadegan. During the summer and after the C. Microzooplankton distribution The specific seasonal distribution of naked ciliates is not detailed in this study... During summer in the Bay of Villefranche. 1999). (BustillosGuzmán et al. 1978–79). episodes of massive algae proliferation. Zweifel et al.3 µM. 1968. 1992). despite the presence of some dinoflagellate species capable of forming red tides. This environmental forcing can inject nutrients into the nutrient-limited surface layer and generate high biomass. 1983. The abundance of dinoflagellates did not reach the values normally reported in red tide events ( Jacques and Sournia. GORSKY ANNUAL MICROPLANKTON CYCLES IN NW MEDITERRANEAN SEA such as Ceratium or Prorocentrum. 1999). The average abundance of naked ciliates in this study was ~1 ind ml–1. In this study. The presence of these large diatom and dinoflagellate species can be explained by the presence of symbiotic cyanobacteria (Taylor. Smalley et al.F. 1997). peaking in May and November (Bernard and Rassoulzadegan. such as Chattonella. 1997). they may feed on the organic matter pool. August 1999. the Ceratium–Prorocentrum assemblages developed first. 1976. 1994). this phenomenon was not detected. 1977). Vertical migration may allow the uptake of new nutrients in deeper waters during the period of surface nutrient depletion (Eppley et al. considered as autotrophic because of the presence of photosynthetic pigment (Lessard and Swift. Abboud-Abi Saab. 1982. During summer stratified conditions.e.. 1993). 1997). Given that they succeed peaks of phototrophs.. small dinoflagellates of the genera Gyrodinium and Gymnodinium became dominant. 1991).. Ferrier-Pagès and Rassoulzadegan. late summer events could overlap with the autumnal bloom (such as in 1998). Their population dynamics may be more related to the prey (bacteria or detritus) distribution than to the changes in the physical and nutritional conditions. An increase in large ciliate (>50 µm) populations ingesting microplankton cells (Smetacek. furca are heterotrophic (Mikaelyan and Zavyalova. in the range of the abundance reported in previous studies (Rassoulzadegan and Gostan. Blasco. 1973. have been reported in the Bay of Villefranche during summer (Trégouboff. furca/P.. During the stratified period in Villefranche Bay. It is probable that the poor nutritional conditions prevailing in the Bay with low nitrate concentration (<0. followed by the Gyrodinium–Gymnodinium group. 1977) reported that Gymnodinium reaches its maximum abundance towards mid-June. 1963) and in other areas (Koray and Özel. The isolated peaks in the abundance of tintinnids and the sudden appearance of predetermined species can be  . chlorophyll concentrations (Dolan et al. GÓMEZ AND G. 1993). 1986). strong wind events were associated with diatom (fucoxanthin) maxima (Bustillos-Guzmán et al. Temperature-dependent distribution of different tintinnid species has been reported in Villefranche (Posta. 1999) or water column stability (Cariou et al. micans peak. Size and abundance of available prey may also influence the abundance and community structure of tintinnids. Experimental studies showed that primary production in the Bay was limited by phosphorus in summer (Thingstad et al. 1999. We reported tintinnid maxima in December–January and November (Figure 3) after the breakdown of the thermocline (Figure 2). 1961).. fusus and C. Rassoulzadegan. 1995) reported diatom maxima associated with wind events in August 1992. 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. however. 1981) and to the availability of large sized prey (Bernard and Rassoulzadegan. 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.

1990. In March–April. 1967 Estrada..  .17 and 0. in (Claustre et al. In Table IV. whereas the autumnal maxima were more prominent. these phenomena can be less relevant and the trophic character is gradually more oligotrophic. the average chlorophyll a concentration is 0. Villefranche Bay can represent an intermediate position between the adjacent zones (Gulf of Lions–Gulf of Genova). At the beginning of the last century. Near Banyuls. This feature surprised him after his studies on the usual spring bloom in the Gulf of Lions (Pavillard. Aug. Eastward. 1956 Zingone et al. a first maximum is composed of pico-nanoplanktonic autotrophs followed by a second maximum in May composed of microphytoplankton.JOURNAL OF PLANKTON RESEARCH VOLUME  NUMBER  PAGES ‒  explained by the seasonal recruitment from cysts as a consequence of a particular environmental condition (Paranjape.. unimodal cycles were reported with a marked maximum in January. chlorophyll peaks reached 3 µg l–1 ( Jacques. 1996 The biomass descriptors are the chlorophyll a concentration or the phytoplankton abundance. 1975 San Feliu and Muñoz. Oct Feb Feb. 1969 Ignatiades. 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. whereas in the Gulf of Genova the maximum was reported in September–October.32 µg l–1 (ranging between 0.. Dec Mar. In some cases the peak of large diatoms was not observed. 1995). 1996 Mura et al. Oct Mar. Oct May. 1967 Margalef and Ballester. 1975 San Feliu and Muñoz. When compared with other Mediterranean coastal areas (Table II). 1937). 1987. Sep. and in spring and late summer in enclosed coastal ecosystems (Cebrián and Valiela. 1997). 1999 Ignatiades. Seasonal cycles of phytoplankton The seasonal pattern of phytoplankton in temperate coastal areas usually shows two annual peaks. 1957 San Feliu and Muñoz. Aug Jan. Lefevre et al. Oct Feb. 1975 Margalef and Castellví. Kamiyama and Tsujino. 1996 Azov..64 µg l–1) and is considered within the range of oligotrophic values (Bustillos-Guzmán et al. May. 1986 Lakkis and Novel-Lakkis. Oct Mar. compared the hydrographic conditions between a typical diatom bloom year [1986. 1970. Nov Mar Feb. The phytoplankton biomass has been evaluated using different estimators such as chlorophyll. Oct Apr. 1993). we have compiled studies on seasonal cycles performed in the Bay of Villefranche.. In Villefranche Bay. 1999). it seems that the bimodal annual cycle is not the usual trend in the Ligurian Sea and Gulf of Lions (Table III). Nov Mar. 1970) and 6 µg l–1 (Ibarra. 1961 1962 1970 1966 1965–66 1978–79 1992 1993 Tolomio et al. 1996). 1984 Siokou-Frangou. fluorescence. Bustillos-Guzmán et al. Oct Apr.. 1995 Herrera and Margalef. Generally the spring bloom of phytoplankton in Villefranche develops in two phases.. Aug 1988–89 1967 1975–1976 1984–85 1983 1973–75 1955 1984–1988 1956–57 1957. 1981 De Angelis. the phytoplankton abundance can be related to the strong winter northerly winds that generate intense mixing processes associated with high primary production (MEDOC Group. the phytoplankton is seasonally influenced by the Rhône river outflow plume (Leveau et al. Pavillard reported the lack of the spring diatom bloom in some years during his study near Monaco. 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.1980 Mura et al. Oct Mar. Soto et al. In the Gulf of Marseille. In the Gulf of Lions. 1981).. in late winter and autumn in open coastal ecosystems. In the coastal Gulf of Lions (Banyuls).. Nov Mar. particles (using a Coulter particle counter) and the abundance of several taxonomic groups. Aug Mar.

. Oct Jun.. 1937 Pavillard. 1984 Bay is not included.F. 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. 1981 Travers. 1970 Neveux et al. The authors concluded that the conditions of 1992 favoured the development of nanoflagellates versus diatoms. Astraldi and Gasparini. 1967 Bernhard and Rampi. 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. 2000) and with a rapid increase of nutrients in the surface waters (Martin et al. When the light and stabilization became optimal for diatoms. 1999). Dec Jun. Bustillos-Guzmán et al. and a non-typical diatom bloom year [1992. in (Claustre et al. 1995)]. 1994). Oct Feb/Sep. The variable intensity of the Corsica channel transport is correlated with the winter NAO index. According to Astraldi et al. 1969 Fabiano et al.. 1999). Years with high transports (and low or negative winter NAO indices) are associated with a remarkable development of the spring diatom bloom.. 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.. 1962. precipitation is associated with the increase of wind stress (Andersen and Prieur. Interannual variability of microphytoplankton: comparison with previous studies The Mediterranean climate is under the continental influence of Europe and Asia.. 1999). In an oligotrophic environment this phenomenon may generate a significant interannual variability. 1995). Higher transports are associated with negative NAO and less water is transported in the Corsica channel during positive NAO years (Vignudelli et al. Oct Jan Jan Mar. 1992.. In March–April 1992 the irradiance increased when the water column was not stabilized. nutrient concentrations could support a regular spring diatom bloom despite the nanoflagellate population growth.. Aug. 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. Aug. 1938 Kane.. Sep. the nutrients had already been depleted by nanoflagellates. 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. Béthoux et al. whereas negative NAO years are associated with wet winters (Hurrell. The prominent interannual variability of the Ligurian current (Le Floch. Air temperatures were colder in 1986 than in 1992 (Fromentin and Ibañez. 1996).. When the intensity of the circulation is lower (positive NAO index) the level of the spring bloom is low (Table V). 1938 Jacques. Nov Oct–Nov Nov Feb. 1967 Kane. The Sahara Desert and the Atlantic Ocean favour thermal anomalies and changes of circulation patterns (Maheras et al. Oct Feb. in (BustillosGuzmán et al. 1967 Bernhard et al. In more eutrophic areas. Sep. The Mediterranean Sea is very sensitive to variations in heat or the water budget (Béthoux et al. 1988]. Sammari et al. Mar. May/Sep. The Corsica channel transport that feeds the Ligurian current (Northern current) presents a strong seasonality. 1963. 1973 Pavillard. 1989). positive values of the North Atlantic Oscillation (NAO) are related to dry winter conditions. 1975 Ibarra. GÓMEZ AND G. In southern Europe and the Mediterranean Sea.. During productive years the spring bloom develops earlier (late March) than during  . 1982. 1937 Bernard. 1967 Bernhard and Rampi. Oct May.. 1995) may influence the nature of the spring phytoplankton bloom intensity on the northern coast of the Ligurian Sea. 1999). In the Ligurian Sea.

Sep. The decrease in precipitation due to the NAO (Hurrell. Aug Mar. whereas the total phytoplankton carbon and the timing of the spring bloom were not correlated (Irigoien et al. Apr. 1996). 1993 Bustillos-Guzmán et al... 1994 Ferrier-Pagès and Rassoulzadegan. 1991 Fernex et al. Vignudelli et al. whereas during 1988 and 1989 this latter sequence did not occur (Fernex et al. Romano et al. During the period 1985–1987 in the Corsica channel. 2000). 1994 Ferrier-Pagès and Rassoulzadegan. Aug. 2000). Lins da Silva. 1999 (unpublished data) Mandleberg. May May Mar. 1994.. Santoreli et al. Thermal anomalies.. followed by high copepod concentration. in Sardinia the precipitation level recorded in 1985 was the highest since 1964 (Delitala et al. 2000). After the 1985–1987 period. 1995). 1991 Lins da Silva. In the Gulf of Genova an exceptional increase in the zooplankton abundance was also reported in 1987 (Licandro and Ibañez. 1995 Feb–Jun. 1997) could be associated with the progressive decrease in diatom abundance in the Mediterranean Sea. the highest levels of precipitation on the Mediterranean basin were recorded between 1985 and 1987 (Boukthir and Barnier. According to the model of Lacroix and Nival. Nov May. and Lins da Silva reported the development of an extraordinary diatom bloom in the Bay of Villefranche in 1986 (Claustre et al. 2000.. 1990. 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. transport was estimated to be the highest in the last 15 years. 1996. 1995 (unpublished data) Therond. 1999). 1995 (unpublished data) Senior. as in late summer of 1999 (Figure 2). 1994 Ferrier-Pagès and Rassoulzadegan. 1991). pointed out high phytoplankton abundance in late March 1987. Claustre et al.. Oct Mar. Aug. 1996 (unpublished data) Senior. Apr... Sep Mar Apr/Mar Mar. Aug.. 1995) and the damming of rivers (Milliman. 1995 Therond. 1996 Bernard and Rassoulzadegan. 2000). For example. In the English Channel. 1988/1989. are associated with unprecedented events of mass mortality of sessile invertebrates in the western Mediterranean Sea (Perez et al. During this period the winter air temperatures were the lowest since 1957 (Fromentin and Ibañez.. based on meteorological data from 1985. 1991 Lins da Silva. Jun May May Apr. 1999 1998 1999 Lins da Silva. 1998). From a study between 1987 and 1989. 1994. Fernex et al.0 Sverdrup (Astraldi et al. During the 1980s. May May (and Mar) Mar...JOURNAL OF PLANKTON RESEARCH VOLUME  NUMBER  PAGES ‒  Table IV: Interannual variability of biomass peaks (using different estimators) in the Bay of Villefranche Biomass peaks Biomass descriptor Period Reference Apr. up to 2. Sep Feb. Francour et al.. 1998). 1994 Bernard and Rassoulzadegan.. 1999 (unpublished data) This study This study the periods of lower production (May).. Mar. 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. Aug Apr. 2000). 1995.. Krahmann and Schott. May May May. Irigoien et al. From the analysis of sapropels. 2000). 1991 Lins da Silva. 1999 Jan–Jun. 1994 Selmer et al. winter transport in the Corsica channel decreased (higher values of the NAO index) and the intensity of the spring bloom declined. reported that the proportion of diatoms during the phytoplankton spring bloom was significantly correlated with the NAO index. 1996 (unpublished data) Mandleberg. Roether et al. and conversely the increase in nanoflagellates and dinoflagellates over  . 1995 Oct’95–May’96 Oct’95–May’96 Jan–Jun. 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. Nov May–Jun May. 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.

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