Offprint Botanica Marina Vol. 43, 2000, pp. 223—231 © 2000 by Walter de Gruyter • Berlin • New York

Mass Development of Marine Benthic Sarcinochrysidales (Chrysophyceaes.l .) in Corsica

L. Hoffmann3*, C. Billardb, M. Janssens3, M. Leruth3 and V. Demoulin3 a University of Liège, Institute of Botany (B. 22), Sart Tilman, B-4000 Liège, Belgium b University of Caen, Laboratoire de Biologie & Biotechnologies Marines, Esplanade de la Paix, F-14032 Caen, France * Corresponding author

A mass development of benthic Sarcinochrysidales (Chrysophyceae s.L), especially of Chrysoreinhardia gi­ raudii comb. nov. (basionym: Tetraspora giraudii) and Nematochrysopsis marina comb. nov. (basionym: Tribonema marinum), is reported from the Corsican coast. Morphology, habitat and seasonal development of these taxa as well as of the less regularly observed Chrysonephos lewisii and of some Chrysophyceae s.l. associated with Chrysoreinhardia giraudii sheaths are presented.

Introduction The benthic algal vegetation of the Bay of Calvi (north-western coast of Corsica) has been monitored Large-scale mucilage accumulation events due to dif­ since the late sixties. Since 1987, great changes have ferent types of planktonic or benthic algae have occurred in the structure of this vegetation. Indeed, plagued different seas in recent years (Vollenweider the Cystoseira communities which dominated all the and Rinaldi 1995). Pelagic mucilage formation is well rocky substrates until 1987 have been subject to a known in the North Sea where mucilage from Phaeo­ dramatic regression of their cover and biomass (Hoff­ cystis (Haptophyta) colonies can produce great quan­ m ann et al. 1988, Janssens et al. 1993 and unpub­ tities of foam accumulating along the coasts lished observations by Demoulin and coworkers). At (Lancelot 1995). In the Adriatic Sea several diatom the same time several benthic macroalgae (Janssens species [especially Cylindrotheca closterium (Ehren­ et al. 1993), especially nitrophilous species such as berg) Lewin et Reimann, Chaetoceros affinis Lauder Cladophora prolifera (Roth) Klitzing and Colpomenia and Skeletonema costatum (Greville) Cleve] are re­ sinuosa (Mertens et Roth) Derbès et Solier, as well as sponsible for large-scale pelagic mucilage production epiphytic microalgae (e. g. Bangiophyceae) (Hoff­ (Degobbis et al. 1995, Mingazzini and Thake 1995), m ann et al. 1994) show a clear progression. In addi­ as has also been observed along other European tion, massive development of mucilaginous benthic coasts (e. g. Boalch and H arbour 1977 off the coast microalgae was observed, the most conspicuous com­ of south-west England). Benthic mucilaginous aggre­ ponents of these mucilages are palmelloid and fila­ gates have mainly been described from the Tyrrhen­ mentous Sarcinochrysidales (Chrysophyceae s.l.). ian Sea (Rinaldi et al. 1995, Diviacco 1992, Giaccone The order Sarcinochrysidales was erected by 1992, Cinelli 1992, Rinaldi 1992, Tnnamorati 1992, Gayral and Billard (1977) and included marine Sartoni and Sonni 1992, Tnnamorati 1995). The most Chrysophyceae characterised by having motile cells frequently reported taxa from these benthic commu­ similar to those of brown algae. On ultrastructural nities are Tribonema marinum J. Feldmann (formerly and phytochemical grounds the homogeneity of this considered to be a member of the Tribonematales, group has been questioned (Billard 1984, see also Bil­ Xanthophyceae), Acinetospora crinita (Carmichael ex lard et al. 1990), followed by O ’Kelly (1989) and Sar­ Harv.) Sauvageau (Ectocarpales, Phaeophyceae), toni et al. (1994). Small subunit ribosomal RNA se­ Microcoleus lyngbyaceus (Klitzing) Crouan (Oscilla­ quences confirmed the difference between the Sarci­ toriales, Cyanophyceae; this name usually covers a nochrysidales V str. and the order Chrysomeridales variety of filamentous blue-green algae) and Lopho­ that still lacks formal description (Saunders et al. cladia lallemandii (Montagne) Schmitz (Ceramiales, 1997). A tendency exists to multiply classes among Rhodophyceae) (Giaccone 1992). The causes of these the heterokont algae and the Sarcinochrysidophyceae mucilaginous aggregates are still subject to discus­ has been provisionally introduced by van den Hoek sion. The development of the Phaeocystis blooms in et al. (1995), while Saunders et al. (1997) favour an the N orth Sea (Lancelot 1995) is apparently due to inclusion in the Pelagophyceae which may not con­ the increase of N relative to P and especially silica. vince those who want a strong morphological basis Mucilage development in the Adriatic Sea also seems for taxa. Several taxa have still not been studied for to depend largely on the N/P ratio, the physical envi­ some critical ultrastructural, chemical or molecular ronment and climatic conditions (Rinaldi et al. 1995). characters and a final decision seems premature to

Brought to you by | Artesis Hogeschool Antwerpen (Artesls Hogeschool Antwerpen) Authenticated | 172.16.1.226 Download Date | 1/10/12 10:02 AM 224 L. Hoffmann et al us. However, it should be clear that we are referring respectively. In 1989, however, the attention of one here to organisms belonging to the order Sarcinoch­ of the authors (V. D. ) was drawn to their abundance rysidales s. str. within the Chrysophyceae s. I. and possible link to the Cystoseira regression. Besides The purpose of this paper is to give an account of Acinetospora crinita, different diatoms and blue-green the benthic chrysophycean flora of the Corsican algae (especially from the genera Hydrocoleum and coast, especially of the Calvi area, together with an Pseudanabaena), form mucilaginous aggregates outline of the seasonal pattern and habitat of the which, however, are dominated by different species most frequently encountered species. of the order Sarcinochrysidales, the most frequently observed being Chrysoreinhardia giraudii and Nema­ tochrysopsis marina and more locally Chrysonephos Materials and Methods lewisii (Taylor) Taylor. Other Chrysophyceae s.l. are Field observations and collections of specimens were also regularly observed but do not form large aggre­ made by SCUBA diving at several locations on the gates. Corsican coast. Most observations were performed in Revellata Bay near the town of Calvi (north-west Chrysoreinhardia Billard, nom. nov. coast of Corsica) where the benthic algal vegetation Replaced synonym: Pulvinaria Reinhard, Notes de has been regularly monitored since the late sixties, la Société de Chercheurs en Sciences Naturelles de No- and especially since 1988, with repeated samplings at vorosisk, IX, 2: 248, 1885, non Pulvinaria Bonorden, fixed points. The points usually monitored numbered Handb. Mykol.: 272, 1851. 19, regularly distributed along the 12 km shoreline The genus name Pulvinaria used by Reinhard between the harbour of Calvi and the tip of the Punta (1885) is illegitimate because of the earlier homonyms Revellata ( Janssens et al. 1993). Routine observations Pulvinaria Bonorden as well as Pulvinaria Fournier were carried out until the rocky substrates ceased or (Farr et al. 1979, p. 1464). The new name Chrysorein­ down to 15 m depth, deeper on more occasional hardia is therefore proposed. This name has already dives. been suggested several years ago and has provision­ Light microscopical observations were carried out ally been used by some authors but had never been using a Leitz Dialux microscope; photographs, mea­ validly published. surements with a screw micrometer and camera lu­ Type species: Chrysoreinhardia algicola (Reinhard) cida drawings were made from living material. Mate­ Billard comb. nov. rial brought back to Caen was maintained in the en­ Basionym: Pulvinaria algicola Reinhard, Notes de riched seawater medium ES-Tris (Cosson 1973), at la Société de Chercheurs en Sciences Naturelles de No- room temperature under natural daylight illumina­ vorosisk, IX, 2: 248, 1885. tion. Living cells were examined and photographed with a Leitz Orthoplan microscope equipped with in­ In the Calvi area, the genus is represented by the spe­ terference optics. For biomass estimations, the muci­ cies: laginous algae were collected from a 100 cm2 surface. Dry weight was obtained after lyophilisation of the Chrysoreinhardia giraudii (Derbès et Solier) Billard collected material. The organic matter weight (ash- comb. nov. (Figs 3 6) free dry weight) was determined by subtracting from Basionym: Tetraspora giraudii Derbès et Solier, the original dry weight the weight of the ash remain­ Suppl. C. R. Acad. Sei., 8,I: pi. I, figs. 1 -6 , 1856. ing after combustion for 6 h in a muffle furnace at Synonyms: Phaeocystis giraudii (Derbès et Solier) 500 °C. Chlorophyll a concentration was determined Lagerheim, Pulvinaria giraudii (Derbès et Solier) after extraction in 90 % acetone according to Jeffrey Bourrelly (invalid combination: incomplete refer­ and Humphrey (1975). Herbarium specimens are ence). held at the Herbarium of the University of Liège Description: The mucilaginous light brownish colo­ (LG). nies are up to 15 mm in diameter (Fig. 6). Cells are hemispherical to spherical, 7-10 |im in diameter and Results and Discussion are embedded in a homogeneous non-stratified muci- Abundant mucilaginous aggregates have been ob­ served from May to July in different sites on the Cor­ Table I. Observations of macroscopic colonies of Chrysore­ sican coast since 1989. Some of the algae involved inhardia in the Calvi area. were present previously since Nematochrysopsis ma­ rina comb. nov. was already occasionally observed J F M A M J J A S o N D in the seventies (Demoulin unpublished observations) * * * * * and its occurrence recorded together with that of 1995 * * * * * * Chrysoreinhardia giraudii comb. nov. by Coppejans 1996 1997 * * * * * * * * * and Boudouresque (1983) as Tribonema marinum and 1998 * * * * * * * * * Pulvinaria giraudii (Derbès et Solieri Bourrelly,

Brought to you by | Artesis Hogeschool Antwerpen (Artesls Hogeschool Antwerpen) Authenticated | 172.16.1.226 Download Date | 1/10/12 10:02 AM Benthic Sarcinochrysidales (Chrysophyceae) in Corsica 225 lage (palmelloid stage) or in a stratified mucilage of 3 m by one of the authors (VD)(Coppejans and (gloeocystoid stage) (Fig. 5). Numerous granules are Boudouresque 1983), but was inconspicuous until present at the periphery of the cell. Cells contain two 1989. The development of C. giraudii is maximal at yellow-brown chloroplasts with pyrenoids and one or the end of spring and in summer (Table I). It gen­ several lipidie granules. Reproduction is by vegetative erally appears in macroscopically visible aggregates cell division and by zoospore formation. in May and disappears in September. In 1997 and Habitat and ecology: Chrysoreinhardia giraudii was 1998, the colonies remained in large numbers until first collected at Alga (not Algajola as erroneously November. From our collection data, C. giraudii ex­ reported) in Revellata Bay in June 1979 from the rhi­ tends to a depth of at least 38 meters, but is often zomes of Posidonia oceanica (L.) Delile, at a depth more sparse below 20 m. The maximum development

Figs 1—4. In situ underwater pictures. 1) Nematochrysopsis marina filaments on a rope; 2) Nematochrysopsis marina filaments developing on the leaves of Posi­ donia; 3) Chrysoreinhardia giraudii colonies covering a large concrete block; 4) Chrysoreinhardia giraudii colonies.

Brought to you by | Artesis Hogeschool Antwerpen (Artesls Hogeschool Antwerpen) Authenticated | 172.16.1.226 Download Date | 1/10/12 10:02 AM 226 L. Hoffmann et al is usually at shallow depths in relatively calm and Table II. Observations of macroscopic Nematochrysopsis in well lit situations in agreement with Verlaque’s (1987) the Calvi area. classification in the ‘groupe photophile infralittoral J F M A M J J A S O N D thermophile’. There it can form a layer several mm thick (Figs 3, 4) that replaces the macroalgae. In 1995 * * * * * * more exposed situations it may, however, start deeper 1996 * * * * * * * and have its maximum at depths of 15-20 m. Com­ 1997 * * * * * * * * petition with Nematochrysopsis marina, which is even 1998 * * * * * * more wave limited, seems to be a major factor in usu­ ally limiting this latter species to the first 10 meters. Dry weight: up to 400 pg cm~2; organic matter: up to unbranched, 13-15(17) pm wide. The cell wall is 110 |.ig cm~2, chlorophyll a: up to 0.25 pg cm~2. formed by H-pieces. Cells are 10-13 pm wide, Geographic distribution: Corsica [Bay of Revellata 15-30 pm long and contain 4 light yellow chloro- (Coppejans and Boudouresque 1983, this study), Pte plasts with stalked pyrenoids, numerous mucous Caldano, Punta Bianca, Bravone, Gulf of Porto, bodies and lipid droplets (Fig. 9). Porto-Vecchio (this study), Galena (Verlaque 1987)]; Habitat and ecology: The alga was first observed Mediterranean coast of France (Castagne 1851, by one of the authors (V. D.) on several occasions in Derbès and Solier 1856, Hamel 1930); Adriatic Sea the seventies in the Bay of Calvi and studied with (Feldmann in Magne 1959); Red Sea (Nasr 1941). Coppejans and Boudouresque in 1979 (Coppejans Comments: This benthic species, first described in and Boudouresque 1983). Since 1989 it has been reg­ the genus Tetraspora, and later transferred to the ge­ ularly observed. Continuous monitoring since 1995 nus Phaeocystis Lagerheim (Haptophyceae), belongs indicates that the filaments of Nematochrysopsis gen­ in fact to the genus Pulvinaria Reinhard (= Chrysore­ erally appear before Chrysoreinhardia in spring inhardia Billard) as suggested by Bourrelly (1957). (April-May) and that it is most abundant from mid- Bourrelly’s combination (1957: 373) was, however, May to July (Table II). Towards the end of Septem­ invalid according to article 33 of the International ber, macroscopically visible aggregates generally dis­ Code of Botanical Nomenclature (Greuter et al. appear, except in 1997, when it was still abundant in 1994), as no complete reference to the basionym was November. It grows on various macroalgae, in partic­ given. ular on Cystoseira balearica Sauvageau and species The type species of Chrysoreinhardia, C. algicola, of Dictyota, on the leaves of the seagrass Posidonia also forms mucilaginous colonies (1-2 mm in dia­ oceanica (L.) Delile (Fig. 2), animals (especially gor- meter) epiphytic on macroalgae such as species of gonians), but also on any hard substrates including Ceramium and Striaria and is closely related to C. artificial ones (metal cages, ropes, etc.) (Fig. 1). This giraudii, if not conspecific. Chrysoreinhardia algicola alga, which is easily disintegrated by waves, only is known so far from a single collection from the grows near the surface in very calm places. In most shores of the Black Sea (Reinhard 1885). If C. gi­ areas it only grows below 6 meters and even deeper raudii and C. algicola proved to be identical, the spe­ in very exposed situations. The optimum is usually cific epithet ‘giraudii’ would nevertheless have prio­ between 7 and 20 meters and it has been recorded rity over ‘algicola’. down to 42 meters. According to Sart oni et al. (1994), The third species of the genus is C. feldmannii this species does not tolerate summer sunshine, but (Bourrelly et Magne) Billard et Fresnel comb. nov. this is contrary to our observations which links its (basionym: Chrysobotrys feldmannii Bourrelly et rarer occurrence near the surface with wave sensitiv­ Magne, Rev. gén. Bot. 60: 684, 1953). Synonym: Pul­ ity. Indeed, in our network of stations this species vinaria feldmannii (Bourrelly et Magne) Billard et does not show a gradual decrease up to the surface Fresnel (Cryptogamie: Algologie P. 281, 1980). It as would be expected if light intensity was involved, forms almost monospecific stands on the mud at the but starts abruptly. The first occurrence is several base of halophytes or on porous limestone rocks in meters deeper in exposed stations as characterised by the supralittoral zone of the French Atlantic coast indicator species and minimal depth installation of (Billard and Fresnel 1980, Billard 1988). Cystoseira balearica as studied by Clarisse (1984). Furthermore, the development largely depends on Nematochrysopsis marina (Feldmann) Billard comb, the weather conditions; strong agitation of the sea nov. (Figs 1, 2, 9) can ‘clean’ the whole bay of the species in summer Basionym: Tribonema marinum Feldmann, Bull. with no later resettlement. Dry weight: up to Soc. Hist. Nat. Afrique du Nord 32: 56, 1941. 1,800 |ig cm~2. Organic m atter up to 800 pg cm~2; Synonyms: Nematochrysopsis roscoffensis Chade- chlorophyll a: up to 5 pg cm~2. faud, Nematochrysopsis roscoffensis f. giganteus Bil­ Geographic distribution: Corsica [Bay of Revellata lard. (Coppejans and Boudouresque 1983, this study), îles Description: The filaments are attached by a basal Lavezzi (Frick et al. 1996), Punta Bianca, Gulf of cell with a mucilaginous sole. Uniseriate filaments are Porto, Gulf of Lava, Porto-Vecchio (this study)]; Me-

Brought to you by | Artesis Hogeschool Antwerpen (Artesls Hogeschool Antwerpen) Authenticated | 172.16.1.226 Download Date | 1/10/12 10:02 AM Benthic Sarcinochrysidales (Chrysophyceae) in Corsica 227

Figs 5—9. Light microscopy. 5) Chrysoreinhardia giraudii cells (bar = 10 pm); 6) Chrysoreinhardia giraudii colony on Halopteris (bar =100 pm); 7) Unidentified Chrysophyceae (bar = 10 pm); 8) Chrysonephos lewisii filaments showing pseudodichotomous branchings (bar = 100 pm); 9) Nematochrysopsis marina cells (bar = 10 pm).

Brought to you by | Artesis Hogeschool Antwerpen (Artesls Hogeschool Antwerpen) Authenticated | 172.16.1.226 Download Date | 1/10/12 10:02 AM 228 L. Hoffmann et al diterranean coast of France (Feldmann 1941, Billard 20 m (Oscellucia ), epiphytic on Cystoseira at a depth 1988); Italy: Sicily (Giaccone 1992, Calvoet al. 1995), of 9 m, July 1997 (Bibliothèque). Tuscan Archipelago (Sart oni and Sonni 1992, Sart oni Geographic distribution: Corsica (Bay of Revellata, et al. 1993), Adriatic Sea (Sartoni et al. 1994); Spain this study); Italy: Tuscan Archipelago (Sartoni et al. (Ballesteros et al. 1986); Atlantic coast of France 1995); Florida, Bermuda Islands (Taylor 1951); Pa­ (Chadefaud 1947, Gayral and Lepailleur 1971, Bil­ cific Ocean and in the Caribbean (cited in Boddi et lard 1988). al. 1999). Comments: Tribonema marinum was described as Comments: The monotypic genus was established belonging to the Xanthophyceae because of the pres­ by Taylor (1951, 1952) on the basis of material col­ ence of H-pieces in the cell wall by Feldmann (1941), lected in Florida and the Bermuda Islands and has on the basis of material collected at a depth of been very rarely reported since. This is the second 20-25 m near Cape Béar (Pyrénées Orientales) where report for the Mediterranean Sea. According to Sar­ it formed mucilaginous tufts on different macroalgae, toni et al. (1995), C. lewisii disappears from the pho- especially Cystoseira spinosa Sauvageau. A similar tophilic algal communities during the wannest species was described five years later by Chadefaud months and is then present at a depth range of (1947) as belonging to a new chrysophycean genus 15-40 m. Nematochrysopsis (N. roscoffensis) on the basis of Chrysonephos is a close relative of the genus Nema­ material collected from aquariums at Roscoff, on the tochrysopsis with respect to its general habit and cy- north-western coast of France. Comparison of tological characters such as presence of chloroplasts Tribonema marinum recently obtained from the type with stalked pyrenoids in agreement with the diag­ locality (Billard unpublished), with a cultured strain nostic features of the order Sarcinochrysidales to of N. roscoffensis isolated from Normandy (strain which these two algae definitely belong, as already CHR8 of ALGOBANK, Caen Culture Collection) pointed out by Sartoni et al. 1995 and Boddi et al. confirms that the two names apply to the same spe­ (1999), rather than to the Chrysomeridales as pro­ cies, T. marinum being identical to N. roscoffensis f. posed in O’Kelly’s (1989) and Preisig’s (1995) classifi­ giganteus considered a large, Mediterranean form cations. (Billard 1988). The distinction between the two forms of Nematochrysopsis based on size differences only Other members of the Chrysophyceae associated with seems unnecessary at the moment and can be attrib­ Chrysoreinhardia giraudii: uted to different ecological conditions. As noted by Polypodochrysis teissieri Magne Billard (1988), large filaments of Nematochrysopsis collected from the Mediterranean can give rise in cul­ Description: Isolated cells are 10 pm wide and ture to smaller individuals. Furthermore, ultrastruc­ 15-18 pm high, with up to 5 open tube-like and up tural and biochemical observations on this taxon by to 100 pm long expansions. They have a single yellow Sartoni et al. (1994) clearly indicate that this alga chloroplast and lack a pyrenoid. does not belong to the Xanthophyceae but to the Sar­ Habitat and ecology: The species is regularly ob­ cinochrysidales. Tribonema marinum is thus transfer­ served in the mucilage of Chrysoreinhardia colonies. red to the genus Nematochrysopsis established by Geographic distribution: Corsica (Bay of Revellata, Chadefaud, the specific epithet ‘ marinum’ having pri­ this study), Atlantic coast of France (Magne 1975, ority over the later epithet ‘ roscoffensis’, the name of Billard pers. obs.). the species becoming Nematochrysopsis marina. Comments: This rarely reported monotypic genus was described by Magne (1975) on the basis of material collected from aquariums in Roscoff, but Chrysonephos lewisii (Taylor) Taylor (Fig. 8) its presence in the Mediterranean was suspected Description: The plants form delicate, whitish to (Magne 1975).Polypodochrysis teissieri is a distinc­ pale yellow tufts, up to 4 cm long. The mostly uniseri- tive marine organism currently placed in the Chryso­ ate filaments are attached to the substrate by an in­ phyceae s. I. conspicuous holdfast, with pseudodichotomous branchings especially above (Fig. 8). Filaments are Unidentified species (Fig. 7) 20-38 |im wide; cells 8-18 pm long, 8-11 pm wide surrounded by an up to 14 pm wide hyaline, non Description: The ellipsoidal cells are isolated or in stratified sheath. Cells have two, four-lobed, band­ small groups, embedded in a homogeneous non-stra- shaped, parietal yellowish chloroplasts with stalked tified mucilage (palmelloid stage) (Fig. 7) or in a stra­ pyrenoids. tified mucilage (gloeocystoid stage). Cells are Habitat and ecology: Bay of Revellata, August 8-10 pm long, 5-7 pm wide with generally a single, 1995: onPosidonia leaves at a depth of 30 m (Pointe), very deeply lobed yellow chloroplast containing a on Posidonia leaves at a depth of 20 m (Biblio­ stalked pyrenoid. Cells are smooth without periph­ thèque), epiphytic on Cystoseira at a depth of 25 m eral granules, but with one or several lipidie inclu­ (Pointe), epiphytic on Halopteris sp. at a depth of sions.

Brought to you by | Artesis Hogeschool Antwerpen (Artesls Hogeschool Antwerpen) Authenticated | 172.16.1.226 Download Date | 1/10/12 10:02 AM Benthic Sarcinochrysidales (Chrysophyceae) in Corsica 229

Habitat and ecology: The taxon is regularly ob­ gates which are considered nuisances by fishermen. served as small groups of cells in the mucilage of Its ecological importance in suffocating benthic com­ Chrysoreinhardia colonies. munities may, however, be superior to that of the Comments: This organism represents a new genus other algae described here. probably belonging to the Sarcinochrysidales and will be described in detail later. The mucilaginous aggregates found along the Cor­ Acknowledgements sican coast are of the same benthic type as the mu­ This study was carried out in the framework of sev­ cous developments in the Adriatic and Tyrrhenian eral FRFC grants and a ‘Action concertée’ program. Seas, where Acinetospora crinita and Nematochry­ LH is research associate of the Belgian National sopsis marina (= Tribonema marinum) are the most Fund for Scientific Research. We are indebted to important components of these mucilages. The Corsi­ C. O. Kelly, Bigelow Laboratory for Ocean Sciences, can flora is remarkably rich in Sarcinochrysidales and Maine, USA, for suggesting to CB the new name especially in Chrysoreinhardia giraudii which so far Chrysoreinhardia. We also express our sincere thanks has not been reported from the recent studies of ben­ to Mme M. Knoepffler-Péguy, Laboratoire Arago, thic mucilages on the Italian coast, whereas it is fre­ Banyuls-sur-Mer, for collecting ‘ Tribonema marinum’ quently encountered and in important biomasses on from the type locality in Cape Béar and to J. Fresnel, the Corsican coast. According to Sartoni (pers. University of Caen, for her gift of figures and stimu­ comm.), it may, however, be present. The difference lating discussions. may be due to the fact that Chrysoreinhardia giraudii remains benthic and does not form floating aggre­ A ccepted 14 D ecem ber 1999.

References Ballesteros, E., M. Y. Perera and S. Puchades. 1986. Contri- Castagne, L. 1851. Supplément au catalogue des plantes qui bució al coneixement algolögic de la mediterrània es- croissent naturellement aux environs de Marseille, 125 pp. panyola, VII. Fol. Bot. Misc. 5: 135-139. + 4 pi. Aix, imprimerie de Nicot et Pardigon. Billard, C. 1984. Recherches sur les Chrysophyceae marines Chadefaud, M. 1947. Une nouvelle Chrysophycée marine de l'ordre des Sarcinochrysidales. Biologie, Systématique, filamenteuse: Nematochrysopsis n. gen., n. sp. Bull. Soc. Phylogénie. Thèse de doctorat d’Etat, Univ. Caen, 223 Bot. France 94: 239 - 243. pp. Cinelli, F. 1992. Il “caso” Argentarola.In: La etisi del Med­ Billard, C. 1988. Les Sarcinochrysidales des côtes frança­ iterráneo in seguido alla fioritura de masse algali. Accade- ises. Bull. Soc. limi. Normandie 110—IIP. 61—69. mia internazionale si scienze e tecniche subaequee ustica; Billard, C. and J. Fresnel. 1980. Nouvelles observations sur Quaderno 9, Firenze: 33—36. le Pulvinaria feldmannii (Bourrelly et Magne) comb. nov. Clarisse, S. 1984. Apport de différentes techniques carto­ (Chrysophycée, Sarcinochrysidale) formant une ceinture graphiques à la connaissance de l’autoécologie de Cysto­ sur substrat meuble. Cryptogamie: Algologie 1: seira balearica Sauvageau, macroalgue marine domi­ 2 8 1 -2 9 2 . nante dans la région de Calvi (Corse). Lejeunia Billard, C., J. C. Dauguet, D. Maume and M. Bert. 1990. N. S. 113: 1 -2 4 . Sterols and chemotaxonomy of marine Chrysophyceae. Coppejans, E. and C.-F. Boudouresque. 1983. Végétation Bot. M ai: 33: 2 2 5 -2 2 8 . marine de la Corse (Méditerranée). VI. Documents pour- Boalch, G. T. and D. S. Harbour. 1977. Unusual diatom off la flore des algues. Bot. Mai: 26: 4 5 7 -4 7 0 . the coast of south-west England and its effects on fish­ Cosson, J. 1973. Influence des conditions de culture sur le ing. Nature, Lond. 269: 687—688. développement de Laminaria digitata (L.) Lam. (Phéo- Boddi, S., M. Bigazzi and G. Sartoni. 1999. Ultrastructure phycées, Laminariales). Bull. Soc. Phycol. Ft: 18: of vegetative and motile cells, and zoosporogenensis in 104-112. Chrysonephos lewisii (Taylor) Taylor (Sarcinochrysi­ Degobbis, D., S. Fonda-Umani, P. Franco, A. Malej, dales, Pelagophyceae) in relation to taxonomy. Eut: J. R. Precali and N. Smodlaka. 1995. Changes in the Pity coi. 34: 297-3 0 6 . northern Adriatic ecosystem and the hypertrophic ap­ Bonorden, H. F. 1851. Handbuch der allgemeinen Mykolo­ pearance of gelatinous aggregates.Hie Science o f the To­ gie, E. Schweizerbart, Stuttgart, xii + 336 pp. tal Environment 165: 4 3 —58. Bourrelly, P. 1957. Recherches sur les Chrysophycées: mor­ Derbès, A. and A. Solier. 1856. Mémoire sur quelques phologie, phylogénie, systématique. Rev. Algol., Mém­ points de la physiologie des algues. Stippi. C. R Acad. oire hors série(I): 1—412. Sei, 7: 120. Bourrelly, P. and F. Magne. 1953. Deux nouvelles espèces Diviacco, G. 1992. Aggregati gelatinosi in Mar Tirreno du­ de Chrysophycées marines. Rev. gén. Bot. 60: 684 — 687. rante Testate 1991: indagini sulla presenza e suU’evoluzi- Calvo, S., R. Barone and L. Naselli Flores. 1995. Observa­ one del fenómeno. In: La etisi del Mediterráneo in segu­ tions on mucus aggregates along Sicilian coasts during ido alla fioritura de masse algali, Accademia internazio­ 1991 — 1992.Hie Science of the Total Environment 165: nale si scienze e tecniche subacquee ustica; Quaderno 9, 2 3 -3 1 . Firenze: 19—32.

Brought to you by | Artesis Hogeschool Antwerpen (Artesls Hogeschool Antwerpen) Authenticated | 172.16.1.226 Download Date | 1/10/12 10:02 AM 230 L. Hoffmann et al

Farr, E. R., J. A. Leussink and F. A. Stafleu. 1979. Index onaux du CNRS, Ecologie des Algues Marines (Dinard, Nominorum Genericorwn {Plantarían). W. Junk, La 20—28 Septembre 1957), C N R S, Paris: 91 — 100. Hague, 3 vols, 1896 pp. Magne, F. 1975. Contribution à la connaissance des Feldmann, J. 1941. Une nouvelle Xanthophycée marine: Stichogloéacées (Chrysophycées, Stichogloéales). Cali. Tribonema marinum nov. sp. Bull. Soc. Hist. Nat. Afrique Biol. Mar. 16: 531-5 3 9 . du Nord 32: 56—61. Mingazzini, M. and B. Thake. 1995. Summary and conclu­ Frick, H., C.-F. Boudouresque and M. Verlaque. 1996. A sions of the workshop on marine mucilages in the Adri­ checklist of marine algae in the Lavezzi Archipelago, atic Sea and elsewhere. The Science o f the Toted Environ­ with special attention to species rare or new to Corsica m ent 165: 9 — 14. (Mediterranean). Nova Hedwigia 62: 119 — 135. Nasr, A.-H. 1941. Some new and little known algae from Gayral, R and C. Billard. 1977. Synopsis du nouvel ordre the Red Sea. Rev. Algol. 12: 57—76. des Sarcinochrysidales (Chrysophyceae). Taxon 26: O’Kelly, C. J. 1989. The evolutionary origin of the brown 2 4 1 -2 4 5 . algae: information from studies of motile cell ultrastruc­ Gayral, R and H. Lepailleur. 1971. Etude de deux ture. In: (J. C. Green, B. S. C. Leadbeater and W. L. Chrysophycées filamenteuses: Nematochrysopsis roscof­ Diver, eds) The Chromophyte Algae: Problems and Per­ fensis Chadefaud, Nematochrysopsis hieroglyphica spectives. Clarendon Press, Oxford, pp. 255—278. Waern. Rev. gén. Bot. 78: 61—74. Preisig, H. R. 1995. A modern concept of chrysophyte clas­ Giaccone, G. 1992. Nematotalli algali mucillaginogeni sulle sification. In: (C. D. Sandgren, J. P Smol and J. Kristi­ coste delia Sicilia e delle sue isole minori. In: La crisi ansen, eds) Chrysophyte Algae, Ecology, Phylogeny and del Mediterráneo in seguito alla fioritura de masse algali, Development. Cambridge University Press, Cambridge, Accademia internazionale si scienze e te ai ic lie subacquee pp. 46 — 74. ustica; Quaderno 9, Firenze: 9 — 18. Reinhard, L. 1885. Recherches algologiques. I. Matériaux Greuter, W., F. R. Barrie, H. M. Burdet, W. G. Chaloner, sur la morphologie et la systématique des algues de la V. Demoulin, D. L. Hawksworth, P. M. Jorgensen, Mer Noire. Notes de la Société des Chercheurs en Sci­ D. H. Nicolson, P. C. Silva, P Trehane and J. McNeill. ences Naturelles de Novorosisk IX (Odessa): 201—512. 1994. International code of botanical nomenclature (To­ Rinaldi, A. 1992. Aggregati mucillaginosi nei mari italiani. kyo Code) adopted by the Fifteenth International Bo­ I casi dell’Adriático e del Tirreno nel quadriennio tanical Congress, Yokohama, August-September 1993. 1988 — 1991.In: L a crisi del Mediterráneo in seguito cdla Regnum Veg. 131: xviii + 389 pp. fioritura de masse algali, Accademia internazionale si Hamel, G. 1930. Chlorophycées des côtes françaises. Rev. scienze e tecniche subacquee ustica; Quaderno 9. Firenze: Algol. 5: 1—54. 4 7 -6 0 . Holtmann, L., X. Detienne, A. GolTart and V. Demoulin. Rinaldi, A., R. A. Vollenweider, G. Montanari, C. R. Fer­ 1994. Studies of marine epiphytic algae, Calvi, Corsica. rari and A. Ghetti. 1995. Mucilages in Italian seas: the III. Variations in the populations of epiphytic Bangio­ Adriatic and Tyrrhenian Seas, 1988 — 1991. The Science phyceae. Cryptogamie:Algologie 15: 53—63. o f the Toted Environment 165: 165 — 183. Holtmann, L., S. Clarisse, X. Detienne, A. Goffart, R. Ren­ Sartoni, G. and C. Sonni. 1992. Tribonema marinum J. Feld­ ard and V. Demoulin. 1988. Evolution of the popula­ m ann e Acinetospora crinita (Carmichael) Sauvageau tions o f Cystoseira balearica (Phaeophyceae) and epi­ nelle formazioni mucillaginose bentoniche osservate phytic Bangiophyceae in the Bay of Calvi (Corsica) in sulle coste toscane nell’estate 1991. Inf. Bot. Ital. 23: the last eight years. Actes du Colloque “Recherches 2 3 -3 0 . océanographiques en mer Méditerranée”, Université de Liège. Bull. Soc: Roy. Liège 4 —5: 263—273. Sartoni, G., S. Boddi and J. Hass. 1995. Chrysonephos lew­ Innamorati, M. 1992. Mucillagini e fitoplancton. In: La isii (Sarcinochrysidales, Chrysophyceae), a new record crisi del Mediterráneo in seguito cdla fioritura de masse for the Mediterranean algal flora. Bot. M ar 38: algali, Accademia internazionale si scienze e tecniche su­ 121-125. bacquee ustica; Quaderno 9, Firenze: 61—80. Sartoni, G., F. Cinelli and S. Boddi. 1993. Ruolo di Innamorati, M. 1995. Hyperproduction of mucilages by Tribonema marinum J. Feldmann ed Acinetospora crinita micro and macro algae in the Tyrrhenian Sea. The Sci­ (Carmichael) Sauvageau negli aggregati mucillaginosi ence o f the Total Environment 165: 65 — 81. bentonici delle coste toscane. Biología Marina, Stippi. Notiz. S. I. B. M . 1: 3 1 -3 4 . Janssens, M., L. Hoffmann and V. Demoulin. 1993. Carto­ graphie des macroalgues dans la région de Calvi (Corse): Sartoni, G., L. Lazzara and S. Boddi. 1994. Ultrastructural comparaison après 12 ans (1978—79, 1990—91).Lejeu­ and biochemichal observations on Tribonema marinum nia N. S. 141: 1 -6 2 . J. Feldmann (Tribophyceae, Chrysophyta). Giorn. Bot. Ital. 128: 820-823. Jeffrey, S. W. and G. F. Humphrey. 1975. New spectropho- tometric equations for determining chlorophylls a, b, cl Saunders, G. W., D. Potter and R. A. Andersen. 1997. and c2 in higher plants, algae and natural phytoplank- Phylogenetic affinities of the Sarcinochrysidales and ton. Biochem, Physiol. Pflanzen 167: 191 — 194. Chrysomeridales (Heterokonta) based on analyses of Lancelot, C. 1995. The mucilage phenomenon in the conti­ molecular and combined data. J. Phycol. 33: 310—318. nental coastal waters of the North Sea. The Science of Taylor, W. R. 1951. Structure and reproduction of the Toted Environment 165: 83 — 102. Chrysophaeum lewisii, Hydrobiologie! 122—130. 3: Magne, F. 1959. Les conditions écologiques favorables aux Taylor, W. R. 1952. The algal genus Chrysophaeum, Bull. Chrysophycées marines benthiques. Colloques Internati­ Torrey Bot. Club 79: 79.

Brought to you by | Artesis Hogeschool Antwerpen (Artesls Hogeschool Antwerpen) Authenticated | 172.16.1.226 Download Date | 1/10/12 10:02 AM Benthic Sarcinochrysidales (Chrysophyceae) in Corsica 231 van den Hoek, C., D. G. Mann and H. M. Jahns. 1995. Thèse de Doctorat d’Etat, Univ. Aix-Marseille II. 389 Algae: An Introduction to Phycology. University Press, pp. Cambridge, xiv + 623 pp. Vollenweider, R. A. and A. Rinaldi (eds). 1995. Marine M u­ Verlaque, M . 1987. Contribution à l'étude du phytobenthos cilages. Proceedings of an International Workshop on d'un écosystème photophile thermophile marin en Médit­Marine Mucilage Phenomena. The Science of the Total erranée occidentale, (Etude structurale et dynamique du Environment 165: 1—231. phytobenthos et analyse des relations Faune-Flore).

Brought to you by | Artesis Hogeschool Antwerpen (Artesls Hogeschool Antwerpen) Authenticated | 172.16.1.226 Download Date | 1/10/12 10:02 AM