Marine Biodiversity in French Guiana: Estuarine, Coastal, and Shelf Ecosystems Under the Influence of Amazonian Waters

Gayana 67(2): 302-326, 2003 ISSN 0717-652X

MARINE BIODIVERSITY IN FRENCH GUIANA: ESTUARINE, COASTAL, AND SHELF ECOSYSTEMS UNDER THE INFLUENCE OF AMAZONIAN WATERS

LA BIODIVERSIDAD MARINA EN GUYANA FRANCESA: LOS ECOSISTEMAS DE ESTUARIOS, LAS COSTAS Y PLATAFORMAS BAJO LA INFLUENCIA DE LAS AGUAS AMAZONICAS

Luis Felipe Artigas1, Philippe Vendeville2, Marc Leopold2, Daniel Guiral3 & Jean-François Ternon3

1L.A.B.E.L. (UMR 8013 ELICO) - Université du Littoral Côte d’Opale - M.R.E.N. - 32, av. Foch – 62 930 Wimereux – France. Email: [email protected] 2Institut Français de Recherche pour l’Exploitation de la MER – IFREMER Délégation Guyane – Domaine de Suzini – BP 477 – 97 331 Cayenne - French Guiana 3L.E.L. (UR R053 ELISA) – Institut de Recherche pour le Développement – IRD Centre de Cayenne – route de Montabo – BP 165 – 97 323 Cayenne Cedex – French Guiana

ABSTRACT

Marine biodiversity in French Guiana is strongly influenced by the amagon River waters of the river Amazon, which constitute a major structuring factor for the estuarine, coastal, and shelf marine ecosystems. Moreover, the marked seasonal and interannual variabilities play important roles in the stability or fluctuations in the environmental parameters that influence biodiversity at the ecological, population, and genetic levels. Previous and ongoing studies of the marine and littoral biota relate mostly to commercial marine species, protected species in danger of extinction and, specially, to the biodiversity and functioning of local coastal and littoral ecosystems such as estuaries, mudflats, sandy beaches and, particularly, littoral mangroves. A more integrated approach involving local, regional, and international scientific collaboration is needed for a better assessment and understanding of marine biodiversity. Such studies would benefit from international cooperation that would allow the gathering of new information and the comparison of previous data, the organization of common oceanographic surveys, the homogenisation of analytical protocols, and also favour the exchange of scientists and postgraduate students for a real transfer of ideas, techniques, and know-how. Moreover, research on the comparative biodiversity of analogous littoral and marine ecosystems in different parts of South America would allow a more accurate estimate of marine biodiversity on a continental scale.

KEYWORDS: French Guiana, marine biodiversity, coastal systems, Amazon river waters, Intertropical Convergence Zone, ecological and conservation studies

RESUMEN

La biodiversidad marina en la Guyana Francesa está fuertemente influenciada por los aportes amazónicos, los cuales constituyen un factor estructurante de los ecosistemas estuarianos, costeros y de plataforma continental. Además, debido a las cambiantes condiciones meteorológicas y oceanográficas, la variabilidad estacional e inter-anual puede jugar un rol importante en la estabilidad o la modificación de los parámetros medio-ambientales que afectarían la biodiversidad ecológica, poblacional y genética de los ecosistemas locales. Los estudios llevados a cabo sobre la biología de las especies marinas y litorales se refieren principalmente a especies de interés comercial, especies protegidas en peligro de extinción, y sobre todo a las características del funcionamiento de ciertos ecosistemas costeros y litorales característicos como los estuarios, bancos de fango, playas de arena, y sobre todo los manglares. La idea de estudios o series de estudios mejor coordinados entre sí a nivel regional e internacional, emerge como una necesidad para una mejor estimación y comprensión de la biodiversidad marina. Estos estudios beneficiarían de una importante cooperación internacional que pemitiese comparar e integrar el conjunto de datos obtenidos en el pasado, llevar a cabo misiones oceanográficas conjuntas, mutualizar y homogeneizar los protocolos analíticos, y favorecer el intercambio de técnicas y estrategias por intermedio de intercambios de científicos y estudiantes en formación de postgrado. Finalmente, estudios comparativos sobre la biodiversidad de sistemas costeros análogos sudamericanos permitirían de alcanzar una mejor estimación de la biodiversidad marina de este continente.

PALABRAS CLAVES: Guayana Francesa, biodiversidad marina, sistemas costeros, aportes de aguas Amazónicas, Zona Intertropical de Convergencia, estudios ecológicos y de conservación.

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INTRODUCTION marine species have also been investigated as well as the biological colonisation of the mudflats. Until recently, most of the marine research As well as this large scale institutional approach, programs developed in French Guiana dealt with local naturalist and ecological associations are sedimentology and geomorphology (e.g. involved with more specific studies (e.g. marine Moguedet 1977, Bouysse et al. 1977, Prost 1989, turtles). This paper reviews, firstly, our present Pujos & Froidefond 1995) or fisheries (e.g. Bullis/ knowledge of the structure of the marine, coastal, Thompson 1959 ; Morice Warluzel 1968 , Abbes and littoral ecosystems, in order to assess local et al. 1972, Bonnet et al. 1975, Venaille, 1979, and regional marine biodiversity in French Dintheer et al. 1989a, Blanchard et al. 2000 ; Rivot Guiana, and then discusses possible perspectives et al. 2000). In 1997, however, the first integrated on short and medium term bases. program (PNEC: National Program of Coastal Environment) supporting coastal oceanographic CLIMATIC AND OCEANOGRAPHIC CONDITIONS research in French Guiana (Fig. 1) was set up with the participation of several French research insti- French Guiana has a typical wet equatorial tutes and universities (Clavier 1996). This climate, driven mainly by the seasonal meridional program focuses mainly on the characterization migration of the Intertropical Convergence Zone and quantification of the influence of the Amazon (ITCZ). In its southern position, the ITZC cau- river on the continental shelf and littoral zone of ses northeast trade winds in French Guiana from French Guiana, in terms of coastal oceanography January to July while, for the rest of the year (physical and bioge-ochemical characteristics in when it is in its northern situation, the country relation to living resources), geomorphology experiences southeast trade winds. There are 2 (mudflat displa-cements along the coastline), and periods of high rainfall, May - June (the princi- mangrove ecosystems (particularly those pal one) and January- February (secondary). The colonizing the stabilized mudflats). The ultimate dry season lasts from July to December when sta- objectives of the PNEC program are to establish ble south-eastern trade winds are dominant. The a better monitoring system of economically flow of the rivers of French Guiana is similar to exploited living resources (fishes and shrimps) that of the rainfall being maximum in May-June as well as improve economical management and and minimum generally in November, when the ecosystem preservation of the coast where about flow rate of the Amazon River is considerably 90% of the population of French Guiana live. lower. The tide in French Guiana is semidiurnal, At present, in addition to fisheries studies and with an amplitude of up to 2.5 meters (spring fish and shrimp stock assessments carried out as tides, mesotidal regime). With its 320 km of parts of the IFREMER programs, ongoing ma- coastline from the Oyapock estuary in the east rine research is concerned with 4 different spa- to the Maroni estuary in the west, French Guiana tial scales: 1) the continental shelf (the ‘‘CHI- is included in the Amazon-Guianas ecotone that CO’’and ‘‘GREEN’’ oceanographic projects - extends from the Amazon River mouth to the PNEC), 2) the littoral the ‘‘ELISA’’ coastal (0- Orinoco River mouth and is considered to be part 20 m depth) project of the IRD, the PNEC of the “Guianas Coastal Province” (Longhurst program, and the Cayenne Island ‘‘LITEAU’’ 1998). The coastal waters of this province are program 3) estuaries (e.g: the Kaw River estuary- highly turbid, specially north of the Amazon PNEC and ELISA-IRD, programs), 4) islands mouth, and extensive mudflats occur and migrate (Grand Connétable Island Natural Reserve). along the coast (Froidefond et al. 1988). Indeed, Three complementary strategies are being from January until July, the meso-scale coastal employed: 1) observations and measurements on current (North Brazil Current, NBC) and its ex- various spatial scales (continental shelf, mudflats, tension, the Guianas Current (GC), flow north- estuaries), 2) the development of a hydrodynamic westwards, carrying the low-salinity and nutrient- numerical model for the continental shelf and and sediment-rich water coming from the coastal zone, and 3) the analysis of satellite data. Amazon along the sorelines of French Guiana, The nutritional requirements of certain targeted Suriname, Guyana and Venezuela (Fig. 2). During

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the period of NBC retroflection, more saline and lowering of the sea surface temperature but less turbid surface waters cover most of the con- provides additional nutrient enrichment, would tinental shelf. These 2 well differentiated le generated by the direction of the prevailing hydroclimatic situations, together with a marked wind and the geostrophic slope of the spatial and temporal variability, represent isopycnals associated with the Guianas major constraints for the coastal ecosystems Current, which is enhanced by the outflowing and their biota. Moreover, constant upwelling, Amazon water (Cadée 1975). that is not accompanied by a pronounced The “SABORD 0” cruise in May 1996 was the first

FIGURE 1. French Guiana Coastal and Shelf areas (from Frouin 1997). Protected areas are shown in rectangles.

FIGURA 1. Areas costeras y de plataformas en Guyana Francesa (Frouin 1997). Las áreas en rectángulos son protegidas.

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FIGURE 2. Surface circulation in the tropical Atlantic Ocean (from Johns et al. 1998, in Ternon et al. 2002).

FIGURA 2. La circulación superficial en el Oceáno Atlántico tropical (Johns et al. 1998, en Ternon et al. 2002).

devoted to the French Guiana continental shelf. A shelf is currently being developed (C. Chevalier, IRD- steep salinity gradient found at about 10 m depth Marseille) and will eventually be linked to a showed the presence of a low salinity surface layer. biological model (M. Baklouti, IRD-Marseille). The Geochemical measurements made on the surface preliminary “CHICO 0” cruise confirmed (Ternon water showed that it was of Amazonian origin (low et al. 2001, 2002) that the hydrodynamics of the

salinity / high silicate / low CO2). The area acts as a French Guiana continental shelf present remarkable

sink for atmospheric CO2, unlike the equatorial ocean features caused by the presence of a large area of which is generally considered to be a global source low salinity water at the surface (8 to 10 m depth),

of CO2 (Ternon et al. 2000). The “CHICO” project, mainly from the Amazon, over most of the shelf which started in 1999 within the framework of the (out to 120 km from the coast). Four water classes “PNEC” program, consists of physical and bio- were identified using radiometric measurements at geochemical oceanographic cruises over the whole wavelengths corresponding to those of the SeaWIFS continental shelf during 3 periods with different sensors and could thus represent water masses of characteristics: February (the start of the wet season; well differentiated biogeochemical characteristics no NBC retroflection; increase in the Amazon flow), (Froidefond et al. 2002). May (the wet season; start of NBC retroflection; maximum Amazon flux), and November (the dry season; NBC retroflection; minimum Amazon flux). THE HISTORY OF BIOLOGICAL OCEANOGRAPHY IN FRENCH Physical, chemical and biogeochemical parameters GUIANA are being determined from the coast (20 m depth) out to the shelf break (200 m depth), on an east to Knowledge about the marine resources of French west transect across the continental shelf. At the Guiana followed the development of fishing same time, cruises for fish and invertebrate species activities over the continental shelf and slope biology (‘‘GREEN’’) are conducted over the same (Frouin 1997, Bernadac et al. 1999). The first area. A numerical model of the dynamics over the studies occurred after the second world war, at

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the end of the 1940s, with the compilation of an conducted surveys of red snapper populations inventory of marine fish species from artisanal (Perodou 1994, Rivot 2000; Rivot et al. 2000). coastal fishery landings (Puyo 1949). During the Similar surveys (IFREMER RESUBGUY) in 1950s, when the American shrimp fisheries in the 1993-94 updated incidental shrimp catch Gulf of Mexico extended to Central America and assessment in the 15 to 60 m depth area in terms of then to north South America, trawling surveys communities and assemblages as well as providing with R.V. Oregon covered both the northern further spatial distribution information (Guéguen Brazil and Guianas areas and were concerned 1993a, b, Nérini 1994, Rosé 1994, Rosé & Achoun mainly with shrimp species, but included the 1994, Moguedet et al. 1995). associated fish species. Between 1954 and 1958, In 1994, another trawling survey (the EPAULARD ORSTOM (IRD) scientists studied the benthic cruise) was carried out in the littoral area from the and demersal continental shelf fauna between 15 coast out to 15 m depth and focused on shrimp and and 100 m depths (Durand 1955, 1959). Later, juvenile fish population distribution (Vendeville during the 1960s and 1970s, ships from the USA, 1995) and, in 1999, a preliminary survey (GREEN Japan, Korea, Venezuela, and Cuba operated off 0 cruise, PNEC project) was made. the Guianas and northern Brazil for shrimp and Research in the infralittoral area began in 1971 with red snapper fisheries (Venaille 1979) and this led the study of a coastal marsh in the western part of to an improvement in the knowledge of the French Guiana and the biology of the young stages demersal and benthic fauna. The local ISTPM of the shrimp Penaeus subtilis; this was in relation laboratory, founded in 1971 (IFREMER), carried to the ORSTOM (IRD) project for the development out trawling surveys between 20 and 1000 m of shrimp aquaculture in the country (Rossignol depths with the R.V. Thalassa (Abbes et al. 1972), 1970, 1972a, b, c). After the introduction of rice the R.V. Phaeton (1975) and the Deana T (1976- cultivation in the area in 1984, a short study was 1978). It also participated in scientific cruises conducted by ORSTOM and IFREMER to evaluate organised by other countries (Dragovich et al. its impact on marine resources (Lointier & Prost 1978), Kaiyo Maru in 1973 (Japan) and Oregon 1986, Dintheer et al. 1985). Between 1988 and mid II from 1972-1976 (NMFS-USA), and, in 1975, 1994, numerous data and observations on the young carried out its own trawling in the littoral area stages of shrimps (postlarvae and juveniles of between 5 and 15 m depths, focusing on seabob, Penaeus subtilis, Xiphopenaeus kroyeri and a coastal penaeid shrimp (Bonnet et al. 1975). Macrobrachyum sp.), in relation to the variability The first primary productivity measurements of shrimp recruitment in the fishery, were collected (CICAR Dutch cruises) in both the Suriname and in several estuaries (Vendeville & Lhomme 1997). French Guiana shelf zones were also made during this period (Cadée 1975). FISHERIES IN FRENCH GUIANA Surveys during a Japan Marine Fishery Resource Research Center (JAMARC) program conducted The continental shelf shrimp fishery in French off Suriname and French Guiana from 1979 to Guiana started to develop with the arrival of US 1983 resulted in the identification of 452 species trawlers at the end of the 1950s and Japanese fishing of bottom fish, 146 species of crustaceans, and boats a few years later. The fishing fleet was largest 143 species of molluscs trawled in the area from in 1983 and then declined steeply until 1991. 10 to 1000 m depth (Aizawa et al. 1983, Takeda Nowadays, the shrimp fleet is exclusively French and Okutani 1983). This survey program also and has 63 trawlers. With the departure of the included an analysis of the shrimp species caught foreign fishing companies, shrimp production was incidentally and a resource assessment directed to the European market where smaller (Vendeville 1984). shrimps were commercially acceptable, and the The IFREMER PENJU (1987-88) and ORSTOM fishing activity tended to move into shallower JUVECREV (1989-90) trawling surveys both areas (Moguedet et al. 1995, Béné 1996, Charuau focused on the spatio-temporal distributions of et al. 2000). The two main species being shrimps (Dintheer & Rosé 1989, Vendeville & exploited are Penaeus subtilis (the brown shrimp) Lhomme 1997). In the same periods, IFREMER and Penaeus brasiliensis (the pink spotted

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shrimp), but Penaeus schmitti (the white shrimp) sediments of the deep slope from 456 to 818 m depth and Penaeus notialis (the pink shrimp) are also (Guéguen 1991 a, b, 2001). In earlier trawling surveys encountered in mixed catches from the area. The (JAMARC; 1979-83) on the Suriname and French coastal shrimp Xyphopenaeus kroyeri (the sea-bob) Guiana continental slopes, 68 species of Decapod is sometimes kept as a bycatch product for the lo- and 63 of Molluscs were identified (Takeda & cal market. The maximum shrimp landings occur Okutani 1983). between February and June. Since 1999, however, the fishery has markedly declined, production DEMERSAL AND BENTHIC CONTINENTAL SHELF falling by 30% from 1998 to 2000 (IEDOM 2001). ORGANISMS The same decrease has been observed in the neighbouring countries and is, in part, linked to Four studies based on trawling surveys on the unfavourable hydroclimatic conditions affecting continental shelf concerned the demersal and recruitment (Charuau in FAO/WECAFC 2001a, sometimes the benthic macrofauna: 2001b, 2002, Dintheer Kalaydjian 2002). · In the first detailed study of demersal and benthic The continental slope shrimp fishery started in 1988. continental shelf macrofauna, Durand (1959) It is seasonal and very small (2 to 4 boats) and identified 21 Echinoderm species, 38 Crustaceans, concerns 2 species: Plesiopenaeus edwardsinus (the 34 Molluscs, and 133 Osteichtyan species. From scarlet prawn) and Solenocera acuminata (the the analysis of the 400 stations sampled, the orange shrimp) (Guegen 1991 a, b). Landings have author concluded that there was a strong link been decreasing since 1988. between the fauna, the depth, and the substratum The second significant continental shelf fishery is quality. that of the red snapper (Lutjanus purpureus). Other · An analysis of the 4 Japanese surveys conduc- species (Rhomboplites aurorubens and Lutjanus ted on the continental shelf of French Guiana, synagris) are also found but less commercially based on 144 sampling stations and using 143 exploited. This fishery is mainly practiced by taxonomic references, pointed out the Venezuelan long line trawlers. predominant role of the depth and, secondarily, About 30 species are commonly exploited by the the nature of the sediment cover, in determining small scale coastal fishery. The principal types are the composition of the demersal assemblages weakfishes (Cynoscion acoupa, C. virescens, (Vendeville 1984). Plagioscion squamosissimus), sharks (Syphyrna · The results of the 1993-94 RESUBGUY surveys spp., Carcharinus spp.), sea catfishes (Arius proops confirmed the roles of depth and substratum in and A. parkeri), Tripletail (Lobotes surinamensis), determining the fish and invertebrate biodiversity snooks (Centropomus spp.), Tarpon (Megalops between 20 and 60 m depths (Nérini, 1994; atlanticus), mullets (Mugil spp.), Crevalle Jack Moguedet et al. 1995; Guéguen 2000). The results (Caranx hippos), stingrays (Dasyatis guttata), and revealed 3 distinct faunal assemblages: 1) a littoral Giant grouper (Epinephelus itajara). (or shallow-water) community present from 0 to 30 m depth in coastal brackish waters of salinities < 25‰ characterized by muddy bottoms; 2) a MARINE MACROFAUNAL POPULATIONS middle-shelf community distributed from 30 to 50 DEMERSAL AND BENTHIC CONTINENTAL SLOPE ORGANISMS m depth in turbid marine waters characterized by muddy bottoms; and 3) a lower-shelf community Of the 40 Decapod (Crustacea) species observed in deeper marine waters of low turbidity (depths during 4 surveys conducted over the continental > 50 m) characterized by sandy bottoms with slope between 200 and 900 m depth from August occasional rocks. In an attempt to simplify this 1990 to July 1991, 2 deep-water shrimps have been distribution pattern, Moguedet et al (1995) the target of the industrial fishery since 1988: the proposed to distinguish between only 2 major orange shrimp Solenocera acuminata (Soleno- types of demersal communities, coastal and deep- ceridae) and the scarlet prawn Plesiopenaeus sea (Table I). Most of the 110 different edwardsianus (Aristeidae). The shrimp A. macrobenthic species observed in that study had antillensis was also widely distributed in the muddy already been encountered in the 1980-81 Japa-nese

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surveys, including 8 Decapod Crustaceans (5 comprised more than 80 % of the total captures Penaeidae, one Palinuridae, one Portunidae and one obtained from 0 to 60 m depth: Dasyatis america- Calappidae), 2 Cephalopods (one Loliginidae and na, Orthopristis ruber, Macrodon ancylodon, one Octopodidae), one sea turtle (Cheloniidae), and Stellifer rastrifer, Upeneus parvus, Arius 99 demersal fish, mainly 9 Sciaenidae, 5 Ariidae, 5 grandicassis, Arius rugispinis, Lutjanus synagris, Serranidae, 4 Lutjanidae, 6 Carangidae, and 9 Chloroscombrus chrysurus, Cynoscion simileis, Pomadasyidae. Despite the high species richness Gymnura micrura and Xiphopenaeus kroyeri. of the benthic and demersal macrofauna, most The most recent study carried out on the species showed a low count, and only 12 of them macrofauna was the preliminary survey cruise,

TABLE I. Summary of the specific composition of the principal demersal communities on the French Guiana continental shelf (from Moguédet et al. 1995 and Frouin 1997).

TABLA I. Resumen de la composición específica de las comunidades demersales principales en la plataforma continental de Guyana francesa (de Moguédet et al. 1995 y Frouin 1997).

Demersal communities Families Species

Coastal 0 – 30 m depth Sciaenidae Cynoscion virescens, Nebris microps, Macrodon ancylodon, Stellifer rastrifer Ariidae Bagre bagre, Arius parkeri, Arius rugipinis Batrachoididae Batracoides surinamensis Clupeidae Dasyatidae Dasyatis americana Penaeidae Xyphopenaeus kroyeri, Penaeus subtilis Gimnuridae Gimnura micrura Sciaenidae 25 – 30 m depth Penaeidae Penaeus subtilis Pomadasyidae Genyatremus luteus, Pomadasys corvinaeformis Engraulidae Anchoa spinifer, Anchoviella lepidentostole Deep sea 30 – 60 m depth Lutjanidae Lutjanus purpureus, Lutjanus synagris Gerreidae Eucinostomus argenteus Carangidae Chloroscombus chrysurus, Selene vomer Pomadasyidae Orthopristis ruber Bothidae Syacium papillosum Mullidae Upeneus parvus

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GREEN 0, realised in April 1999. The role of (Arius grandicassis, Bagre bagre), with increasing bathymetry as the principal structuring factor of the numbers of Sciaenidae (Cynoscion virescens, benthic and demersal ecosystem, already identified Macrodon ancylodon, Paralonchurus elegans, in previous studies as mentioned above, was P.brasiliensis ), and Dasyatidae (Dasyatis confirmed and defined by the observations made americana). While the nature of the sediment layer during the GREEN 0 cruise (Le Loeuff & Von Cosel is the principal structuring factor of the benthic and 2000; Vendeville et al. 2000, 2002), which took place demersal communities of the French Guiana conti- in the middle of the wet season and the period of nental shelf, in shallow regions, the brackish coastal maximum discharge of both the local estuaries and water has a great influence on the littoral the Amazon River. The importance of the nature of communities. In the latter ecosystem, nychtemeral the substrate was pointed out and the nychtemeral rhythms would be the third structuring factor. rhythm was also defined as a structuring factor. The Therefore, the principal causes of the temporal salinity gradient probably does not have a direct variability of fishery resources have to be defined influence on the spatial distribution of the benthic in terms of the strict spacial limits of this ecosystem. and demersal populations because the low salinity Definitely, 3 adjacent ecosystems seem to interact is localised in the upper layer. in controlling nutrient inputs and the biological During the CHICO 0 cruise, an inorganic nutrient cycle, and determining the productivity of the layer was observed near the sea bottom, where system: the continental shelf, the littoral zone, and shrimps were abundant. This may in part be the local rivers. explained by the remineralisation by heterotrophic bacteria of the organic matter arriving on the bottom DEMERSAL AND BENTHIC LITTORAL ORGANISMS as waste from shrimp fishing. As many as 107 benthic species (Cnidaria, Polychaeta, Crustacea, During the 1994 littoral trawling survey, Mollusca, and Echinodermata) were then identified, “Epaulard”, carried out between the coast and as well as 139 demersal species belonging to 3 15 m depth, 52 species were observed in the zoological groups (Cephalopods, Chondrichtyans, catches at the 81 stations sampled: 8 Crustaceans, and Osteichtyans). An analysis of the specific including 5 shrimps, two Penaeidae (Penaeus richness and abundance of these communities subtilis and Xiphopenaeus kroyeri), one showed that only a few species in shallow sea-beds Palaemonidae (Nematopalaemon schmitti); are characterized by high abundances, whereas sometimes very abundant), one Hippolytidae numerous species identified in the deep sea-bottoms (Exhippolysmata oplophoroides) and one are characterized by weak abundances. Four types Sergestidae (Acetes sp.), two crabs, and one sto- of communities were distinguished, 3 of them being matopod. associated with the principal regional Penaeid Molluscs were represented by only one gastro- shrimps: 1) Deep communities of the ‘‘clear wa- pod, Bursa bufo. ters’’ at 55 to 85 m depths, in mid and coarse-grain Three species represented Batoid fish, two spe- sands and coral reefs: shrimps (Penaeus cies of Dasyatidae and one of the Gymnuridae. brasiliensis), soles (Gymnachirus nudus), coral reef Among the 40 bone fish species encountered, the fishes, red snappers (Lutjanus purpureus), and main families were Ariidae (7 species), Sciae- grunts (Orthopristis rubber); 2) Mid-depth (55 – nidae (9 species), and Carangidae (5 species). 65 m) communities, in fine-grain sands and rocks: Most of the catches concerned young individu- lane snappers (Lutjanus synagris), grunts als. AFC and CAH analysis showed that bottom (Orthopristis ruber) and searobins (Prionotus quality and hydrology (temperature and salinity) punctatus); 3) Mid-shelf (45 – 55 m) communities, were structuring factors of these communities in muddy sands of fine and very fine grains: shrimps and, in particular, the role of Guianese river (Penaeus subtilis), some Sciaenidae (Micro- plumes in the distribution of these species pogonias furnieri, Stellifer rastrifer, Cynoscion si- (Vendeville 1995; Vendeville & Lhomme 1997). milis); 4) Littoral communities of ‘‘turbid waters’’ Most of the knowledge on coastal fish diversity at 15 to 35 m depths, of mud and muddy sands: comes from a study on small scale fishery landings ‘‘seabob shrimp’’ (Xiphopenaeus kroyeri), Ariidae (Blanchard et al. 2000).

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FISH AND INVERTEBRATES IN THE INFRALITTORAL AREA A REVIEW OF MARINE ICHTHYOLOGICAL DIVERSITY IN (COASTAL MARSHES AND ESTUARIES) FRENCH GUIANA

In 1971-72, preliminary investigations were Marine fish species are not systematically stud- conducted in a western coastal marsh near Mana in ied in French Guiana and it is difficult to quan- order to introduce brown shrimp (Penaeus subtilis) tify fluctuations in the populations. IFREMER aquaculture. Biological data were collected on the (Cayenne) focused its studies mainly on the ex- juveniles and postlarvae of the brown shrimp and ploited fish (targeted species and incidental aquatic fauna (Rossignol 1972c). A later study by catches). Species associated with rocky substrates IFREMER in 1985 concerned mostly the brown remain badly identified, as are pelagic species shrimp (Dintheer 1986). A brief study in 1994 and the small benthic species of the littoral. In discovered that the marsh had evolved and that its particular, some uncertainties persist with regard function as a shrimp nursery had been greatly to certain families such as the Scianidae reduced (Le Cour Grandmaison 1995). The (Plagioscion genera), Mugilidae, Serranidae, presence of predators of the brown shrimp Engraulidae, Clupeidae, and Gobiidae. (Mugilidae, Tarpon atlanticus, and Centropomus The data available on the ichthyological fauna undicimalis) and of some juvenile fish (Achirus comes from various sources, not all published, achirus, Batrachoides surinamensis, and Hemi- and are enriched by information gathered during gramus ocelifer) was noted. scientific surveys conducted by French and for- A study of the life-cycle, recruitment, and young eign institutions throughout the 20th century (see stages of the brown shrimp was carried out from earlier paragraph on the history of biological 1988 to 1994. Bi-monthly samples were collected oceanography in French Guiana). Fourmanoir in several estuaries in order to assess the seasonal (1971) provided the first record of and interannual variations in its postlarval and Carcharhinidae species in French Guiana waters. juvenile abundances. The postlarvae entered the The atlases of the freshwater fish of French estuaries on the flood tide and left on the ebb tide. Guiana published by the MNHN (Paris) (Keith The recruitment of postlarvae occurred all the year et al. 2000; Le Bail et al. 2000, Planquette et al. round on the French Guyana coast with between 2 1996), and some INRA publications (Le Bail et and 4 intensive periods, and it was concentrated al. 1984a, b, Rojas-Beltran 1984, Boujard & Rojas- more in the western area. Data on other shrimp and Beltrán 1988) allow an approximate estimation of prawn species were also collected (Xiphopenaeus the number of brackish water fish species found kroyeri, Nematopalaemon schmitti, Acetes sp., and in the littoral to be made. Information is at present Macrobrachium sp.). being compiled by M. Léopold (IFREMER) on The Kaw estuary was also sampled, between fishes of the Guiana continental shelf. This list August 2000 and July 2001, for brown shrimp could be updated and supplemented in the next together with its associated fish community few years if the project of the Ichthyological In- (Vendeville 2002). The sampling showed 3 ventory of French Guiana led by the MNHN (Paris) seasonal peaks of brown shrimp postlarval materializes, and also if the planned GREEN abundance: in September, a weaker one in cruises (PNEC program) take place. December with the third, also weaker, in A 2 nautical mile margin around the Connétable February-March. These periods of abundance Islands (4°51’N, 51°26’W) to protect the bird coincided with those which had been observed population constitutes the only real marine re- in other estuaries (Vendeville & Lhomme 1997). serve of French Guiana. These small islands are The transport of the postlarvae during a tidal colonized by a large community of sea birds (frig- cycle, as characterized in other estuaries, was ates, sterns, noddis, and gulls) that feed on cer- also observed in the Kaw River estuary and tain fish species. No study has yet been made of extended to juvenile fish coming from the coastal its marine fauna but a survey could take place zone. soon of the numbers of the giant grouper, Epinephelus itajara, there.

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PLANKTONIC BIODIVERSITY IN COASTAL productivity and competition with heterotrophic WATERS bacteria for nutrients are in progress.

THE ECOLOGY OF THE MICROPLANKTON IN COASTAL AND THE BIODIVERSITY OF THE PHYTOPLANKTON IN THE ESTUARINE WATERS MARINE AND BRACKISH WATERS OF FRENCH GUIANA

The coastal waters of French Guiana are subject to Only a few surveys of the phytoplankton species large inputs of dissolved and particulate matter from have been made. The first data for the area came the Amazon River, from the local hydrological sys- from the adjacent zones (Teixeira & Tundisi 1967; tem, and also from the extensive mudflats by Margalef & González-Bernáldez 1969) and, only resuspension processes (Froidefond et al. 1988). Of later, from 1976 to 1979, were the microplankton these, the Kaw River carries most of the drainage of local marine and brackish waters studied from the Kaw swamp. Near the littoral mudflats, (Paulmier 1993). the high levels of suspended particulate matter limit Skeletonema tropicum was prevalent in the estuarine light penetration, modify the N/P mineral balance zone with Chaetoceros subtilis var. abnormis, due to the adsorption of phosphate (Guiral & Dytilum sol, Thalassionema nitzchioides, Plenecassagne 2002) and reduce phytoplankton Coscinodiscus jonesianus, C. aculeatus, C. oculus- growth (DeMaster et al. 1991). A zone of high chlo- iridis, C. curvatulus, and Odontella mobiliensis also rophyll and primary production has been observed present. Resuspended benthic diatoms (pennate outside the turbid coastal waters on the continental forms) and thycopelagic (centric and pennate) forms shelf (Cadée 1975, Ternon et al. 2001, 2002). Artigas together with sessile or pedonculata types were & Guiral (2001, 2002) carried out a comparative characteristic of the estuarine waters. The most study of the phytoplankton and bacterial biomasses representative taxa were the Naviculaceae and present during 2 extreme seasonal conditions. At Nitzschiae-Navicula sp., Entomoneis alata var. the end of the wet season (July 2000), the high lev- pulchra, Nitzschia sp., Nitzschia longissima, N. els of degraded pigments in the riverine and transi- sigma var. intercedens, N. circumsuta, Bacillaria tional zones suggested the presence of large amounts paradoxa. Some Cyanobacteria (Lyngbya and of allochtonous matter from the flooded lands and Oscillatoria) were also occasionally present in the Kaw swamp, and forest marshes and wetlands. samples. The Dinophyceae were rare except for The would have been in an advanced state of degra- Scrippsiella sp. dation and thus stimulated bacterial production lead- The coastal zone includes the archipelagos ing to large numbers of these microorganisms asso- (Connétable Is., Rémire Is., Salut Is.). Although the ciated with the very low chlorophyll a concentra- hydrographical parameters have wide ranges tions. Transport from the mudflats and young man- (salinity varying from 12 to 35‰ for instance), this groves seems to contribute to the high pigment con- zone is more stable and highly productive, especially centrations measured, in July 2000, in the mudflat the surface waters. Pelagic diatoms are predominant littoral waters. The resuspension of benthic diatoms in the phytoplankton, though the Dinophyceae are and/or their grazing by benthic organisms might also important during certain seasonal periods. explain the prevalence of large forms and degraded Skeletonema tropicum is responsible for major pigments. In November, mean total estuarine chlo- blooms in May that are often coupled with lower rophyll concentrations were up to 10 times higher numbers of Ceratium fusus (Dinoflagellate). than in July, and were correlated, along the estua- Coscinodiscus asteromphalus, Thalassionema rine gradient, with bacterial abundances. The high- nitzschioides, Odontella mobiliensis, Chaetoceros est photosynthetic rates in coastal waters were ob- subtilis var. abnormis, and Cerataulina dentata served in November, probably due to the combina- (Diatomophyceae) are also present in this area, as tion of senescence and the grazing of a large are Noctiluca scintillans, Ceratium lineatum, and phytoplankton bloom enhancing and diversifying the Protoperidinium oblongum (Dinophyceae). Dytilum pathways of organic matter transfer between sol is dominant in August, associated with Nitzschia photoautotrophs and heterotrophic bacteria. New pungens, Chaetoceros appendiculata, and Ceratium studies of phytoplankton species cycles and their lineatum. In October, Coscinodiscus jonesianus is

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the dominant species, associated with nated by Skeletonema associated with Odontella. Coscinodiscus karstenii, Chaetoceros These species were also detected in in the sub- appendiculata, Thalassio-nema nitzschioides, surface layer of shelf waters, associated with the Diplopeta bomba, and Ceratium lineatum. genera Denotula, Hemiaulus, Chaetoceros, Rhizo- The shelf province (>50 m depth) is dominated by solenia (Diatoms), and Ceratium furca and C. fusus a single diatom, Thalassionema nitzschioides and (Dinophyceae). Very low cell abundances charac- a cyanobacterium, Oscillatoria (Trichodesmium) cf. terized the wet period. The majority of the fresh thiebautii; the other secondary species belong to water genera were represented by different different pelagic genera: Proboscia (Rhizosolenia) Desmiidae (Closterium, Pleurotaenium, alata, Pseudosolenia (Rhizosolenia) calcaravis, Micrasterias torreyi, M. furcata, M. abrupta, and Hemialus sinensis, and Climacodium fraufel- M. laticeps, Staurodesmus, Staurastrum, Xan- dianum. Dinoflagellates are also characteristic of thidium, Desmidium, Cosmarium, Sphaerozosma, this system, especially Ceratium tripos var. Spondylosium), Chlorophyceae (Eudorina, atlantica, C. massiliense, C. trichoceros, C. vultur, Ankistrodesmus falcatus, Botryococcus braunii), C. contortum, etc. Paulmier (1993) indicates that Euglenophyceae (Lepocinclis, Traochelomonas the dominant species in the coastal area from De- hispida, Euglena, Phacus), and Dinophyceae cember to July are the “Skeletonema-Thalassio- (Ceratium, Peridinium). nema-Coscino-discus-Ceratium society” which corresponds to Group II of Margalef & González- ZOOPLANKTON IN THE ESTUARINE AND COASTAL WATERS Bernáldez (1969) who characterized a similar OF FRENCH GUIANA phytoplanktonic coastal community in Venezue- lan waters. This is temporarily followed by a Protozoans make up an important part of the “Coscinodiscus association”. pelagic biomass and are normally flagellates, The oceanic area is dominated by the species dinoflagellates and diatom consumers, such as Nitzschia pungens and the genus Rhizosolenia. the Sarcomastigophora, classes of the Rhizopoda This association corresponds to Group II as de- and Actinopoda represented by the genera fined by Dandonneau (1971) for the Ivory Coast Granuloreticulosia (Foraminidae), Radiolarians waters, confirming the ubiquity of the tropical and Acantharians, and also the Ciliophora Atlantic species. Even if the phytoplankton is comprising the Spirotriches (Tintinnidae). The dominated by Diatomophyceae (Chromophytes) genera Tintinnopsis, Favella, and Coxliella are and by Dinoflagellates (Pyrrhophytes), other taxa common in estuarine and littoral waters, whereas such as Euglenophytes (Euglena spp.), and Rhabdonella, Epiplocylis and Eutintinnus are Chlorophytes (Spirogyra spp., Closterium spp.) more oceanic (Paulmier 1993). In the estuarine are often present in brackish estuarine and litto- zone, zooplankton population’s maxima are in ral waters. The Chrysophyceae is particularly May, July, and December and, very often, consist important with Phaeocystis (Order: Prymne- of holoplanktonic larvae, such as copepod nauplii siales) represented by 2 species, P. pouchetii and and adults of the genera Oithona, Cyclops, and a larger form which is very abundant under cer- Paracalanus (P. cf. crassicornis). The Cladocera tain hydrochemical conditions. The Cyano- e.g. Bosminia probably originate from fresh- bacterie of the Schizobacteria group also peri- water. The Rotifers and especially the Tin- odically constitute a significant biomass in pre- tinnidae, Tintinnopsis karajacensis, Tintinnopsis oceanic neritic waters and in the littoral area. sp., Codonaria fimbriata, and Favella Maia de Oliveira (2000) studied the phytoplank- panamensis are the other major representatives ton in 2 estuarine systems (the Sinnamary and of the zooplankton. The composition of the Iracoubo Rivers). The effect of tidal changes on zooplankton is mostly homogeneous from the the phytoplankton diversity of the Kaw River coastline out to 50-60 m depth. Oithona sp., estuary, were reported by Vaquer (2000). At neap Acartia tonsa, the nauplii of various copepods tides, the dominant genera were large and Tintinnopsis sp., are the most frequent species Coscinodiscus, Thalassionema, and Odontella, of this area. The zooplankton is very much more whereas at spring tides, the community was domi- diversified in the oceanic area and includes pelagic

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molluscs. Two size-classes were considered by LITTORAL BENTHIC COMMUNITIES AND Lam Hoai & Rougier (2000, 2002) in their study of POPULATIONS OF FRENCH GUIANA the zooplankton: microzooplancton (40-150 µm) and mesozooplancton (>150 µm). LITTORAL MANGROVE ECOSYSTEMS: THEIR STRUCTURE The mudflat station is different from the 2 other AND ECOLOGICAL FUNCTION stations of the Kaw River estuary, in the channel of the river itself and in the Paul Emile creek When the massive migrating mud waves, which which drains a mature mangrove. During the wet stretch 1,100 km along the northeast coast of South season and high flow period, the Kaw River America, from the mouth of the Amazon River to estuary becomes an extension of the riverine that of the Orinoco River, settle out, they produce ecosystem. The drainage waters from the Kaw the mudflats that are progressively colonized by swamp are characterized by very low abundances mangroves which extend along almost all of the and biomasses of zooplankton with a strong 320 km coastline of French Guiana (Fig. 3; Prost predominance of Rotifers, in terms of abundance. 1989), occupying a band from a few meters to about Sixty-one taxa corresponding to 27 taxonomic 15 km wide (Vignard 2001). The mudflats are rap- groups have been identified. Tintinnopsis is the idly covered by mats of microalgae that find the principal genus present in microplankton assem- light conditions optimal for their growth. These blages. The dominant species of Copepods were microalgae, dominated by colony forming diatoms, Paracalanus crassirostris, Oithona ovalis, and have the ability to migrate into the sediments, ac- Acartia giesbrechti whereas the dominant cording to the tidal and daily irradiation cycles. Tintinnids were Codonellopsis and Favella in Progressively, a benthic community of Nematods, neap-tides and Tintinnopsis in spring-tides. The Tanaidacea, and Foraminifera becomes established pelagic copepods, Paracalanus crassirostris, within the mudflat (Vignard 2001, Guiral 2002). A Acartia giesbrechtii, and Oithona ovalis, number of species use the tidal cycles to benefit provided 63% of the total biomass in neap-tides from these resources: limicole birds and crabs at as opposed to only 31 % in spring-tides. Rotifer low-tide, and numerous fish and postlarval and ju- populations were represented in neap-tides by venile shrimp at high water. Despite high levels of Synchaeta and Trichocerca but were practically species biodiversity in the ecosystem as a whole, absent in spring-tides. On the other hand, the the number of plant species is relatively few in the Chaetognatha Sagitta frederici constituted a ma- coastal mangroves, only 4 indigenous genera be- jor part of the biomass in spring tides. The ing represented-Avicennia, Rhizophora, Lagun- detection of rare taxa such as Pseudodiaptomus cularia and Conocarpus (Dodd et al. 1998). Five marshi, Ostracods, Oikopleura, Cletodidae, species of mangrove trees have been identified: the Diosaccidae, Canthocamptidae, the hydromedusa red type (Rhizophora mangle and R. racemosa, Obelia, the Copepod Eucalanus, Decapod larvae, Rhizophoracea), the grey-black type (Laguncularia Polychaeta larvae, Anthomedusa, Isopod larvae and racemosa, Combretacea), and the white type Nematods varied greatly between flow and ebb tides (Avicennia germinans, Verbenaceae). A very rare and day and night periods. This hydrological context species, Conocarpurs erecta (Combretaceae), is for the wet season is radically different from that located at only one site in French Guiana. These of the dry season and the reduced flow of the river may be separated into coastal and estuarine man- when the estuary is regularly subjected to tidal in- groves (de Granville 1986). The white type pre- trusions of saline water. This period is characterized dominates in coastal mangroves, but recently de- by relatively significant estuarine biomasses of posited mudflats are colonized firstly by the grey- zooplankton whereas populations and biomasses black type and only later by A. germinans. The ge- upriver are considerably smaller. This seasonal nus Rhizophora dominates upstream from the es- enrichment of the estuarine zone is thus related tuaries. In parallel with the changes in hydrography, directly to the penetration of the littoral waters a sequence can be recognised in the shoreline-main- which fertilize and increase the local productivity land direction: young trees growing on the even, and also transport the zooplankton communities fluid muds in the intertidal zone, then mature for- present in both the littoral and coastal waters. ests of large trees (medium density and low mortal-

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ity). On the older alluvia are the senescent stages al. 2003) is to analyse the process of mangrove of the trees (high mortality). The aim of the ongo- colonization and to identify its role within the ing mangrove study (PNEC program, Fromard et coastal food webs.

FIGURE 3. Migration of the mudflats along the Guianese Littoral. Situation in 1988 (from Prost 1989, in Frouin 1997).

FIGURA 3. La migración de las marismas a lo largo del litoral de Guyana; la situación en 1988 (de Prost 1989, en Frouin 1997).

THE PHYSICAL CHEMISTRY OF THE LITTORAL MUDFLAT cies of fish and 34 species of crabs and shrimps. WATERS AND ITS ECOLOGICAL IMPLICATIONS FOR Some of them live in these habitats permanently, BIODIVERSITY while others, e.g fish and crustaceans, temporarily visit the mangroves to feed (Hogarth 1999), to pro- As already mentioned, in the dry season, ‘‘marine’’ tect themselves from predatory pressure (Primavera waters contribute to the transport of phosphate into 1997), or to acheive part of their life cycle. Thus, the rivers, whereas, in the wet season, the ebb tides various species find that these systems provide the carry particulate suspended mineral and organic best conditions to ensure part of their development matter out to sea (Vignard 2001, Guiral 2002); in cycle: shrimps (Primavera 1998), postlarval and this context, a large part of the phytobenthos could juvenile fish, and the adult stages of crabs (Jones also be moved within the water column. 1984). The young stages of these species are mainly The water draining from the coastal mangroves is detritivores (Rojas-Beltrán 1986) and are therefore rich in nutrients and suspended matter and tends to at the base of the trophic chain of the estuarine sys- be brackish though its salinity varies greatly, de- tem. Significant populations of Penaeus subtilis and pending on the rainfall. The fauna which live in Plagioscion sp. of small average size and the ab- these habitats is thus characterized by euryhaline sence of sexually mature individuals indicate the species. According to Rojas-Beltrán (1986), the role of the mudflats as nurseries for these species. mangroves in French Guiana shelter some 100 spe- These two taxa make up the largest trophic level,

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their juvenile stages being omnivores capable of sequences were related to Gelidibacter algens feeding on detrital material produced by the sur- (Cytophaga-Flexibacter-Bacteroides group), rounding mangrove. The adults of Platystachus Actinobacteria, Sulfitobacter and Ruegeria spp. cotylephorus and Potamotrygon hystrix constitute (K-proteo-bacteria), all of which are chemo- a secondary trophic level, the primary being carni- organotrophs, consistent with abundant labile vores. Lastly, the adults of Plagioscion sp. and organic carbon. Frequent resuspension, Centropomus sp. are intermediate carnivores (Rojas- reoxidation of bottom sediments, and re-deposi- Beltrán 1986). Of the 22 species found in this mudflat tion, coupled with the entrapment of system, 9 are considered to be permanently estua- allochthonous organic particles including organ- rine, seven mainly freshwater, and six mainly ma- isms, create transient environmental complexity rine. Based on the stage of maturity and the diets of and promote high microbial diversity. Further work the taxa in samples described by Odum & Heald on the microbiology and biogeochemistry of these (1972) and (Rojas-Beltrán 1986), it would appear that mobile muds is being planned. 41% of the captured species would be shrimp con- The productivity and adaptation of the photosyn- sumers when adults (Caranx sp., Lutjanus sp., thetic biofilms that colonize the littoral mudflats Centropomus sp., Arius sp., Plagios-cion sp., Strong ecological interactions, including compe- Platystacus cotylephorus, Potamotrygon hystrix, tition, exist between microphytobenthos and man- Gobioides sp., and the crabs, Callinectes sp.). grove vegetation. Photosynthetic biofilms, one mainly consisting of cyanobacteria and another of PROKARYOTE DIVERSITY AND AN ANALYSIS OF COMMUNITY diatoms, were sampled on a mudflat of the Kaw STRUCTURE IN THE MOBILE MUD DEPOSITS OF FRENCH estuary (Guiral et al. 2001, Sygut 2002). Four GUIANA sediment samples were taken from the Kaw mudflat: one from the central part of the mudflat Venkateswaran et al. (1998) were the first to have in the course of being colonized by a phytobenthic isolated a bacterium (Shewanella amazonensis) community including diatoms and cyanobacteria, from Amazonian shelf coastal muds in the north- another from the upper sector colonized by a ern Amapá region where shore-attached mud phytobenthic mat, very largely dominated by waves are first formed. Madrid et al. (2001) have Chlorophyceae, a third from the central part of this since provided a molecular census of sector dominated by mobile pennate diatoms within prokaryotes in a shore-attached coastal mud the sediment, depending upon the irradiance, and wave, 3-4 m thick, migrating to the northwest the last from the lower part, where there was only over a relict mud surface. These authors exam- one settlement of very large centric diatoms, ined the relationships between the inferred mi- inherited from of the oceanic area. A fifth sample, crobial community and a range of seabed sedi- taken from the zone of a mature Rhizophora and mentary and geochemical properties, and its Avicennia mangrove, was of a sediment very potential as an indicator of hydrodynamic proc- partially colonized by cyanobacteria and actively esses. A sequence analysis, extracted from a exploited by crabs. New studies are planned to subsurface sediment layer (10-30 cm) of 96 non- determine the factors which control the deve- chimeric sequences showed the majority of the lopment of autotrophic biofilms and their microbes to be bacteria. The diversity was high, productivity. with 64 unique sequences, and Proteobacteria were dominant. Bacterial sequences belonged to DIATOM DIVERSITY IN THE MUDFLATS AND MANGROVES the Cytophaga-Flexibacter-Bacteroides group, OF THE KAW RIVER ESTUARY Actinobacteria, Planctomycetes, Cyanobacteria, low-GC Gram-positive, K, Q, and N subdivisions The diatom communities in 85 samples of surface of Proteobacteria. A sizeable fraction of se- sediments taken from habitats ranging from mo- quences from the Kourou-Sinnamary library are bile mud to stabilized mangroves of the Kaw River normally found in water column populations, estuary during 3 cruises-at the beginning of the dry reflecting the frequent entrapment of suspended season (July 2000) and in the rainy season (Janu- debris into the underlying sediments. Dominant ary 1999 and December 2000)- were studied by

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Sylvestre et al. (2002). 178 taxa belonging to 51 gen- genus Navicula characterize the settling areas era were identified. The diatom communities were which are subject to higher variations of the en- dominated by periphytic species; only 34 species vironmental parameters. This study demonstrated were identified as holoplanktonic forms. The that the penetration of oceanic waters has a big tychoplanktonic species were represented by 2 types influence on the ecosystems of the Kaw estuary, of Synedra in 3 samples. All of the identified spe- purely freshwater habitats occurring well up- cies were characteristic of saline conditions. The stream from the estuary. strictly marine forms were all planktonic. Most of the periphytic forms were species observed in FORAMINIFERAL BIODIVERSITY littoral as well as in continental saline situations whereas the diatom communities were all char- In the 37 surface sediment samples collected in acteristic of coastal systems. Five species January 1999 and July 2000 for the study of the (Coscinodiscus centralis, Cymatodiscus microfaunal biodiversity in the mangrove swamps planetophora, Cyclotella stylorum, Gyrosigma and mudflats of French Guiana where strong sea- peisonis and Thalassionema nitzschoides) were sonal variations occur, 44 species were identified present in all of the samples. (Debenay et al. 2002). The main factor regulating The periphytic forms were mostly of the genera the distribution of foraminiferal assemblages is the Gyrosigma, Pleurosigma and Nitzschia; these live hydrodynamics of the estuary, characterized by the both on and in the mud, their migration into the influence of both saline coastal water and low cal- sediment depending upon sediment resuspension cium fresh water, with major seasonal differences. and the irradiance conditions. A freshwater spe- The coastal end member is dominated by the cal- cies (Eunotia spp.) was observed in only one sam- careous species Ammonia tepida, A. parkinsoniana, ple taken upstream of the estuary. The samples and Cribroelphidium spp. whereas the continental taken on the unstabilized mudflat contained the (freshwater) end member is characterized by the typically marine planktonic species, agglutinated species Miliammina fusca and Coscinodiscus centralis, Cyclotella stylorum, and Trochamminita irregularis. The calcareous species Thalassionema nitzschoides, which are charac- penetrate the estuary, even as far as the mangrove teristic of the Atlantic Ocean. These are found, forests, during the dry season but totally disappear however, not only on unstabilized mudflats also from the estuary during the rainy season. The other in mature mangroves and are indicators of the factor affecting the distribution of the Foraminifera penetration of oceanic waters far into the estu- is their vertical elevation which determines the pe- ary. The samples taken on the fluid mudflat and riod of aerial exposure and the colonization by the slikke were dominated by Gyrosigma spp., mangrove trees. Foraminifera are very rare on or which possesses a high degree of mobility within even absent from exposed mud banks containing the sediment. Finally, in the samples from the cracks, but begin to grow as soon as young mangrove swamps stabilized by vegetation, Avicennia are present. Both the canopy and the de- Nitzschia spp., considered to be mobile epipelic tritus of the mangrove forest protect the sediment forms, and Navicula spp., characteristic of muddy from heating and drying and, thus, increase in sa- shores, were predominant. The mature mangrove linity caused by the sun and wind. The distribution samples were characterized by a greater specific of the agglutinated species in the mangrove swamps richness and a greater diversity than those from is, as a result, closely related to the presence of the the other locations. The parameters such as sa- mangrove trees. linity, nutrient concentrations, and pH have a great influence on species diversity. This study MEIO- AND MACRO-BENTHIC BIODIVERSITY IN THE LIT- suggests that the motile epipelic species TORAL MUDFLATS OF FRENCH GUIANA (Gyrosigma spp. and Nitzschia spp.) are indicators of fluid mudflats, with a gradation from the The meiofauna of the Kaw mudflats are largely oceanic influenced area mostly dominated by dominated by Nematods (Ragot 1999, Boucher Gyrosigma spp. to progressively more stable ar- 2000). The “marine slikke” zone has significant eas dominated by Nitzschia spp. Species of the populations of adult and nauplii stages of

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Harpacticoidae Copepoda (Crustacea) and related to the highly turbid littoral and estuarine Turbellariae (Annelida). Due to their high indi- waters. vidual biomasses, the insect larvae found within the inner mudflats represent 50% of the total THE CARCINOLOGICAL FAUNA OF A FRENCH GUIANA biomass. The importance of the Tanaïdaceae at MUDFLAT COLONIZED BY MANGROVES some locations on the Kaw river mudflat and the “estuarine slikke” of the mudflat were confirmed The extremely rapid establishment of a mangrove by a comprehensive meiobenthic survey that in- makes it possible to follow, on a very short time cluded the young stages of this group. scale (3 to 5 years), the progressive transformation The Nematods were the numerically dominant of an unstabilized mudflat into a mangrove group of the meiobenthic fauna. A total of 66 inhabited by the carcinological fauna specific to this species were identified with a mean of 16 spe- type of habitat. The interactions between vegetation cies per sampling site and a maximum of 28 spe- and crabs is demonstrated by the occurrence of cies in the mature Rhizophora mangrove. The species which depend first on the shade provided Monhysteridae family was the most abundant and by the trees and then on the production of leaves this characterizes the nematofauna as a commu- and litter which constitute their food (Amouroux nity of non-selective detritivores. Two species of & Tavares 2002): Goniopsis sp. feeds on the young this family represented over 25% of the nematods foliage of trees and Ucides sp. consumes analyzed. The second community is represented decomposed leaves, whereas Aratus pisonii depends by the specific epistratum grazers, Dicromodora on the primary production of Rhizophora trees and and Metacrhomadroides. These genera are the epibiontes of their stilt roots. adapted to the grazing of diatoms and are domi- The gradual disappearance of Uca maracoani is nant on the marine slikke. related to changes in its habitat. Initially, the loss Justou (1999), Glevarec (1999) and Clavier of mud without vegetation narrowed its surface and (2000) have reported the presence of 26 the mud covered with vegetation became compacted. macrobenthic taxa in the Kaw mudflat area. The Uca maracoani is now, therefore, confined to the Tanaïdacea Crustacea is the most abundant taxa, sloping edges of the river. Its place has been occupied representing 75% of the total abundance, fol- by other Uca spp. and also by Ucides cordatus. The lowed by the Polychaeta and Oligochaeta settlement of Rhizophora at the edge of the river allows (Annelids). The communities were found to be the development of Aratus sp. The population densities low in diversity though the populations covered of the various species increase as the mangrove a wide range of abundances; this structure is com- develops. The appearance of Ucides sp. and the anas- mon in an environment subject to a major physi- tomoses of the burrows of Uca sp. in 2001 were indi- cal constraint. It was possible, as a result, to dif- cations of increasing bioturbation. The survey of this ferentiate between the stations of the stabilized mudflat made it possible to show a process of faunal mudflat, the mangrove and mudflat at the young succession and colonization of the different habitats stage of tree colonization, the mudflat marine near the littoral mudflats. There are plans to continue slikke, and the areas colonized by the dense com- this research, as part of the PNEC program, over a munities of Tanaidaceae. The total biomass was longer period, to supplement it with a study of the very low over the mudflat (a factor of 10 less river margins colonized by Uca sp., and to follow it than the mean biomass) except within the drain- up with a senescent mangrove area. age channels. The macrobenthic biomass was, on the other hand, high in the estuary border slikkes STUDIES ON PARTICULAR PROTECTED SPE- and, especially, in the mangrove zones. CIES CONSERVATION STRATEGIES The small-sized macrobenthos was dominated by deposivores, mostly little Peracarid Crustacea and The coast of the Guianas is a mix of vast mudflats, Capitellidae Annelida. Suspensivores were very narrow sandy beaches, mangrove swamps, wet rare and only some Cirripedae (Crustacea) and savannas, and brackish water creeks. The mangrove Mytilidae (Mollusca) were observed on floating forests of the Guianas are among the most impor- branches. The low numbers of suspensivores are tant and least degraded in the world and are home

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to millions of North American migratory birds. Off- Guianese and WWF scientists to better under- shore, the strong Guianas Current flows east to west stand turtle biology. In French Guiana, sea turtle carrying the nutrient-rich mud of the Amazon and conservation activities are coordinated by the the local smaller rivers and creeks. In a continual DIREN (the Ministry of Natural Resources for process of erosion and rebuilding, the Guianas Cur- French Guiana). Partners working with DIREN rent seasonally creates and then washes away huge include associations such as SEPANGUY and mudflats and wide stretches of sandy beach (Fig. Amerindian organizations such as Kulalasi. The 3). This process is important because the Guianas Amana Nature Reserve, WWF, ONCFS, Kwata, provide nesting sites and feeding grounds for some Université d’Orsay and CNRS are also active part- of the largest remaining populations of 4 species ners in local sea turtle conservation. of sea turtle. Mangroves provide critical habitats Leatherback (Dermochelys coriacea), green (Che- for many marine species such as shrimp, tarpon, lonia mydas), olive ridley (Lepidochelys olivacea), bonefish, and rays. Within the Guianas, the main and hawksbill (Eretmochelys imbricata) turtles nest threats to biodiversity are the removal of man- on the beaches of French Guiana. With the exception groves; over-hunting of turtles, birds, and other of the hawksbill, for which there are not enough data, mammals; incidental catches of turtles by shrimp the nesting seasons are well-known: green turtles trawlers and fisheries activities, the poaching of (January-May), olive ridleys (May-September), and turtle eggs; over-fishing; increasing pesticide and leatherbacks, almost all the year round but divided fertilizer use in expanding rice fields; bauxite min- into two distinct nesting seasons, whit a minor peak ing operations; and land clearance for livestock. In in December/January and a major peak during April- addition to the known threats, there is the possibil- July. ity that other industries, such as sandmining, The primary nesting sites in French Guiana are lo- goldmining, and oil development may also be di- cated at Awala-Yalimapo, along the Point-Isère/ minishing the biodiversity of this ecoregion. Organabo coastline, and at the urban sites of Kourou and Cayenne. The Awala-Yalimapo Beach (Amana STUDIES OF THE BIOLOGY AND ECOLOGY OF MARINE Nature Reserve) is historically considered the most TURTLES (FROM THE WWW SITE OF WWF ©) important single nesting site in the world for the leatherback sea turtle; during the primary peak sea- Together with partners at STINATSU (the Foun- son (May-June), more than 200 of them may nest there dation for Nature Conservation), indigenous com- in a single night. It is also an important nesting site munities in Suriname, and KAWANA in French for green turtles, though this species has not been for- Guiana, the WWF has been supporting sea turtle mally monitored. Olive ridley turtles (on average, less conservation in the Guianas for over 20 years. than 10 nests per year) and hawksbill turtles appear to Since 1999, with the assistance of WWF Guianas, be only occasional nesters. WWF-France has gradu- this regional approach has gradually taken shape. ally refocused its activities on to the protection of nest- In recent years, local organizations and commu- ing females during the leatherback egg-laying season nities have begun to play an even more pivotal (the Kawana Campaign, Talvy et al. 2001), in col- role in the conservation and management of these laboration with Kulalasi (an Amerindian organization species (Talvy et al. 2001, Fretey et. al. 2002); based in Awala-Yalimapo) and the Amana Nature recently, they have completed a Regional Sea Reserve staff. Turtle Conservation Strategy that will provide the Beaches along the Pointe-Isère/Organabo coastline regional program with a framework and goals for shift with changes in the shoreline structure, mak- future conservation work. In all 3 countries, sea ing consistent monitoring difficult. Nevertheless, the turtle conservation activities include research and area is known to be an important nesting habitat enforcement, and developing ecotourism and al- and is considered to be the main nesting area for ternatives to unsustainable fishing practices and green turtles in the country. the harvesting of turtle meat and eggs. Classic Finally, there are 2 important “in-town” sea turtle research techniques, such as surveys, and new nesting sites, in Kourou and Cayenne, in French research techniques, like satellite tracking, PIT- Guiana. They nest on 4 beaches, covering a distance tagging, and genetic research, are enabling of about 4 km, in the vicinity of Kourou. In Cay-

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enne, nesting sea turtles are currently using 6 main shrimp populations, which has greatly reduced a sites, spread over about 12 km. These beaches are in major food supply for the dolphins. an urban setting with buildings and other man-made structures often only a few meters away from the THE BIRDS OF THE LITTORAL ZONE OF FRENCH GUIANA nests. Two local organizations, KWATA and SEPANGUY, are monitoring these sites. Their re- More than 200 of the 700 species of birds which ports show the importance of these sites for olive populate French Guiana are essentially or com- ridleys and leatherbacks (Talvy et al. 2001). pletely habitants of the littoral wetland areas (Hansen & le Dreff 2002), and more than two-thirds of the THE DISTRIBUTION, HABITAT, AND CONSERVATION STATUS 30 birds cited in the “Red book of the endangered OF THE WEST INDIAN MANATEE IN FRENCH GUIANA species of the French overseas regions” (Thiollay 1988) live in these wet coastal systems. The wild- The manatees, Trichechus manatus, may be less life of the littoral of French Guiana, which has been abundant than in the recent past (de Thoisy et al. less studied than that of the rainforest, presents a 2002), but are still present and regularly sighted all very particular evolutionary dynamic which con- along the coast and in estuaries, up to 80 km inland stantly modifies its structure. from the open sea. The main habitats for the mana- Three of the 10 species of ducks (Anatidae) observed tees are estuarine mangroves. At the moment, their in French Guiana are strictly migratory Anas discors, secretive behaviour, fairly undisturbed estuarine A. americana, and A. clypeata (Hansen & le Dreff habitats, species scarcity, and protection laws cre- 2002). Dendrocygna autumnalis and Anas ate an optimistic outlook for the future of the mana- bahamensis are nesting species in French Guiana. tee populations in French Guiana. The manatees are The great majority of the Anatidae are located on also encountered on nearby rocky shores (Cayenne the western part of the coast and particularly in the Island, Malmarouny, Ilet-la-Mère and Rocky point Mana lagoon. This particular location has been pro- at Kourou) and up to 80 km upstream in some of tected by creating the Amana Regional Natural Re- the larger rivers as the Maroni, the Approuague and serve, as well as the Kaw swamp Regional Natural the Oyapock. As the alluvial coastal plain is quite Reserve. narrow, their habitats are naturally restricted. The emblematic red ibis, Eudocimus ruber, is a protected species along the whole of the French Guiana A STUDY OF SOTALIA FLUVIATILIS (CETACEA, DELPHINIDAE) coast and lives at the mouths of rivers or in the coastal IN COASTAL WATERS marshes. Not less than 24 species of herons are present in French Guiana, occupying various habitats such as The results of a bibliographical study coupled with mudflats and littoral lagoons, freshwater marshes and field work (direct observations and the collection forest creeks. The most abundant species are Egretta of data since 1997) on the distribution and possible thula and E. caerulea. These species share their habi- threats to Sotalia fluviatilis in the waters of French tats and nesting sites with 6 other species, e.g. Egretta Guiana have been reported by S. Bouillet (2002). tricolor. Egretta alba is frequently observed in marshes These dolphins are observed along almost all of the and rice plantations as well in the littoral. Ardea cocoi littoral area. They seem to be present all through is the most numerous representative species of the the year towards the east of the French Guiana coast Ardeidae in South America. but seem to be more frequent in the dry season Among the rarest species, the pink “spatule”, Ajaia towards the west; this could be connected with the ajaja, is present at littoral sites of difficult access occurrence there of less turbid water and better food and Mycteria americana is mostly observed in stocks. They are rarely observed at the mouths of coastal lagoons. Some small groups of pink flamin- and within the principal rivers (Oyapok and Ma- gos (Phoenicus ruber) are regularly seen coming roni) where river traffic is rather heavy, but more from Suriname and Amapá State in Brasil. frequently within the mouths of the secondary rivers Nycticorax nycticorax, N. violacea, and Cochlearius (Larivot and Mahury) and around the “du Salut” cochlearius are small Ardeidae species that live in and “Connétable” Islands. Habitat destruction seems dense mangroves and reproduce with other Ardeidae to be less important than the 3 year decline in the and the red ibis.

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Some birds, not strictly marine, inhabit the littoral search teams coming from Metropolitan France, freshwater marshes; typical of these are Ixobrychus together with some local scientists, temporarily involucris, I. Exilis, Botarus pinnatus, and Zebrilus based in French Guiana, working for various re- undulatus. On the other hand, Butorides striatus is search institutions (IRD, IFREMER, CNES, a very common green heron that occupies different Institut Pasteur, Université Antilles Guyane - UAG, parts of the coastal plain, as does Bubulcus ibis. CNRS, etc.), Governmental Agencies (DIREN, Twenty-seven species of North American limicolous Natural Regional Parks) or different types of Non- birds have been recorded in French Guiana. “Small” Governmental Organizations, national (KWATA, limicolous birds include most of the sandpipers, GEPOG, SEPANGUY, etc.) and international Calidris pusilla being the most abundant, and then (WWF). These scientists were or still are involved Charadrius semipalmatus. Recent new censuses in projects of limited duration (PNEC, PNRZH, of limicoles were carried out by ONFCS and GEPOG. LITEAU, etc.). Therefore, except for the programs Small species arrive during their post-nuptial migra- that dealt with species of particular commercial tion between August and October. Thereafter, a part interest or those already identified as endangered of the community migrates towards the Brazilian species, the present review describes the outcome coast and, in January-February and April, they re- of mainly short-term studies which focussed on turn to the Arctic. Some of the large limicoles ob- specific parts of the different ecosystems under served are Tringa flavipes and T. melaneuleuca, precise hydroclima-tological conditions. There is, Arenaria interpres and Calidris canutus. therefore, a need for a more generalized long-term The rocky islands in the coastal zone of French series inventory of the existing data (including Guiana provide refuges for marine birds such as geographical distributions) and a more detailed Fregata magnificens that travel up to 100 km away assessment of marine biodiversity at the ecologi- from the coast, following the shrimp fishing ves- cal, population and genetic levels. In addition, sels, as do seagulls. Nearly 500 couples of F. the available information needs to be compared magnificens are observed all through the year in with and, indeed, extended to the neighbouring the “Grand Connétable” Marine Natural Reserve. countries of the “Amazon-Guiana Ecotone” or The numbers of the seagull, Larus atricilla, in marine “Guianas Province”, from the mouth of the French Guiana represent 20% of the Caribbean popu- Amazon River to that of the Orinoco River and lation and the most southerly community of this the southern Caribbean Sea. A large reservoir of species. existing expertise could be called upon for carry- The Cayenne stern, Sterna eurygnatha, is one of ing out these studies, if there were greater finan- the most endangered species in the “Grand cial and logistical support from national French Connétable” Natural Reserve, as very few nesting scientific institutions (Universities, CNRS, IRD, colonies are known. An international cooperative IFREMER, MNHN, INSU, and CNES). The conservation program has been established with creation of new academic careers to deal with Aruba Island off the Venezuelan coast which hosts the environment, planned for the near future by a similar sized colony of this stern. Sterna fuscata is the local university, UAG, together with the for- represented by 250 couples on “Grand Connétable” mation of a local unit of the National Research island. Like Anous stodilus, this species often flies Council (CNRS) and the renewal of IRD and away from the highly turbid littoral areas as far as IFREMER research groups in French Guiana, 90 km offshore. The royal stern, Sterna maxima, is involving a significant number of local and met- often seen associated with the Cayenne stern on ropolitan French scientists, are elements that the “Grand Connétable”. should ensure the success of projects of medium- term duration involving also neighbouring countries, financially and scientifically supported by DISCUSSION AND PERSPECTIVES national and inter-regional programs. At present, only the ECOLAB network, initiated in 1993, is The information about marine biodiversity in improving coastal studies in all disciplines French Guiana which has been described in this through the communication and collaboration paper is the result of studies carried out by re- between scientists from northern Brazil, French

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Guiana and Suriname. Cooperation in marine re- edges, characteristic habitats of the littoral) would search between an institute in French Guiana and allow the development and application of common another in Venezuela has also been proposed. strategies for the assessment and management of All these ongoing initiatives could be strengthed the marine biological richness of the continent. within the CoML Program. First of all, some studies This could begin with different strategies at local, could be carried out at the ecological level for a interregional, and international levels, within a me- better understanding of the functioning and energy dium-term project such as the CoML program (10 and material fluxes in the different estuarine, litto- years), which envisages two or three projects as are ral, coastal, and open sea ecosystems already indi- currently being developed in French Guiana. The vidually studied in previous and ongoing programs. first need would be to compare capabilities in or- Existing data, at different levels and scales, need, der to see how much each participant can achieve first of all, to be brought together in a common once the goals and time scales have been defined. Geographical Information System data base. The aim would be to identify what local studies Coastal oceanographic cruises covering the entire can be made fairly quickly in order to prepare the area, supported by satellite imaging, would then be different stages in the research strategy during the a good strategy for integrating local studies, though next 10 years and thereafter. the protocols used for data gathering would need to be standardised in order to compare the measurements and estimations of stocks and fluxes made ACKNOWLEDGMENTS in the different countries of the region. A major research effort is required to increase and We would like to thank all of the colleagues men- coordinate field observations of the littoral, not only tioned for the use of their published and/or unpub- by scientists but also by the local populations that lished data, and their long-distance or close sup- know and benefit from these ecosystems. In this port, in the achievement of this review. Special way, an improved inventory of the known and still thanks go to J.P. Debenay and F. Sylvestre for their unknown marine species, from microbes to turtles, data and comments. Thanks are also due to Dr. G.H. dolphins, and birds, could be produced. For this, it Sala for allowing Dr. L.F. Artigas to work at the would be essential to cross fertilize the capabilities IRD Cayenne Center since 2000, and to all the tech- of all of the scientists involved in the area. It is also nical and administrative staff of the IRD Cayenne important to stress that most classifications are for their help. Dr. B. de Thoisy of the KWATA As- changing rapidly as microscopic (SEM, TEM) and sociation in Cayenne is gratefully acknowleged for biochemical and genetic probe techniques improve his information concerning the conservation actions our knowledge and ability to discriminate between that are currently being undertaken in French different species and taxa in general, and between Guiana. Finally, we want to express our gratitude microbial organisms in particular. Moreover, assess- to the local and national coordinators of the PNEC ments of genetic diversity could be made by means Program for their support. of international cooperation, in order to establish the in situ diversity of communities and populations. This would allow the accurate identification, clas- REFERENCES sification, and incorporation into an inventory of the organisms present at different locations and ABBES, R., Y. ALDEBERT, D. DOREL, C. LEROY, R. LEMEN, J. during the various seasons. The exchange of scien- PRADO & C. SAINT-FELIX. 1972. Reconnaissance des fonds de pêche de la Guyane Française. Cam- tists and of students involved in postgraduate stud- pagne de la THALASSA dans la région Antilles- ies and research should be facilitated in order to Guyane, 15 juin - 1er septembre 1971 - Science et improve the global vision of the personnel study- Pêche, Bull. Inst. Pêches marit. 210: 1-22 ing marine biodiversity in South America. Finally, AIZAWA, M., T. ARAI, E. FUJII, T. INADA, K. MATSUURA, T. comparative research on the biodiversity of analo- MIYAKE, K. SASAKI, Y. SATO, T. SHIMIZU & T. UYENO. 1983. In: Fishes trawled off Suriname and gous littoral and marine ecosystems in South French Guiana. Uyeno T., K. Matsuura E. Fujii America (large river coastal plumes, upwelling (Eds). JARMAC: 1-519 zones, estuaries, narrow or extended shelves, shelf AMOUROUX, J.M. & M. TAVARES. 2002. La faune

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