The dyked freshwater marsh of Bourgneuf (Loire-Atlantique, France) Preliminary study of the fish community. Species-habitat relationship. Evidence of main management problems
The dammed up marshes of the French Atlantic coast cover about 200 000 ha between River Vilaine and the « Bassin d'Arcachon ». Eighty eight % are managed with freshwater. They constitute original environments initially created for agriculture or for salt production, and they are now threatened by land abandonment within the next decade. Concurrently to aquaculture (in created or existing ponds), exploitation of the fish stocks in the ditchweb is likely to encourage a diversification of agricultural activities. Unfortunately, bibliographic analysis reveals the relative scarceness of research about sampling methods and qualitative or quantitative characteristics of these fish communities. This is quite surprising considering the importance of the ditchwed of this kind of environment outlined by several authors. In the Netherlands, BELTMANN (1984) assessed that there is a total of 400 000 km of ditches. In France, the littoral dyked marshes of the Atlantic coast couid comprise 20 000 km of ditches and about 24 000 ha of open water. The present work provides for the first data on the fish community of Bourgneuf marsh. The northern part of the marsh of Bourgneuf, 2 700 ha provided whith f reshwater, contains nearly every kind of landscapes found throughout the whole Breton-Vendéen marsh. The pattern of the ditch network strongly changes from a zone to another (fig.1) : presence of former salt pans in the western part, regular geometric shapes in the recently created polders next to the River Falleron, irregular ditchweb pattern in the eastern part. The average density of the ditch network is 91 m of ditches per ha, totalizing 234 km in the study area. The total surface of open water, composed of ditches and basins (former salt pans), covers 411 ha (over 15 % of the study site). Diversity of ditch types occurs at fine scales (<1 000 m2), they vary according to their widths (0,3 to 7 m), depths (average, 42 cm; SD, 20,4), thickness of silt layer (average, 43 cm; SD, 42) and their hydrophyte vegetation cover (average, 70 %; SD, 60 %). As a consequence of this heterogeneity, available habitats are scattered over the marsh (mosaïc distribution). A nested sampling (FRONTIER, 1983) was carried out to take into account this high heterogeneity : 5 sampling areas were selected randomly. In each one, 3 to 5 ditches were chosen according to their characteristics (see above). Sampling stations were delimited by 2 stop nets (5 mm mesh) settled 30 m apart, in order to avoid fish migration. Field work was conducted using « Heron » electric fishing material (see LAMARQUE et al., 1978). In each ditch-section, we carried out as many successive catches as necessary to apply the maximum likelihood weighted estimation method of CARLE and STRUB (1978). Nine to 19 stations were sampled at 5 periods, between 1987 and 1989. A total of 74 samples were collected. The fish community was composed of 21 species (table 1) and corresponded to the bream zone of Verneaux's classification (1977). The densities and biomass were quite high (on average 315 kg/ha and 11 460 fishes/ha) but very variable (0 to 2 120 kg/ ha and 0 to 39 300 fishes/ha). The catfish, Ictalurus nebulosus (170 kg/ha), the eel, Anguilla anguilla (47 kg/ha) and the tench, Tinca tinca (28 kg/ha), represented on average 77,5 % of the standing crop, but their spatial distribution was very irregular. These estimates are assumed to be reliable considering that the data used for the calculations were provided by a sampling design which permits to respect the basic assumptions of the removal method. (f) The population size could only change because of the fishings (no migration because of the stop nets; no recruitment/death because of the short duration of the fishing sequences). (ii) The standard sampling design permitted to reduce the variations of the catch probabilities between the successive removals. Several studies have shown that this removal method under-estimates by about 20 % the true size of the fish populations (e. g. BOHLIN and SUNDSTROM, 1977; MAHON, 1980). But they were based on Zippin's method, and the estimator of CARLE and STRUB (1978), that we used, was shown to be more robust (COWX, 1983; GERDEAUX, 1987). Nevertheless, we assume that the values presented in this paper provide for an approached information on the sizes of the studied fish populations. To assess the fish-habitat relationship, a correspondence analysis (fig. 2) was performed on the 74 samples X 17 species matrix (excluding the sticklebacks, Gaslerosteus aculeatus and Pungitius pungitius, which population size estimations failed because of their low catchabilities). Four groups of samples were ordinated according to their specific richness and the species they contained. Several habitat parameters were projected on F1-F2 factorial map (fig. 3). Hydrophyte cover, thickness of silt layer, water depth (fig. 3 and 4), which are directly controlled by human maintenance, appeared to be the major structuring habitat parameters for the fish community. In the deepest and less silted stations, the communities were rich (on average 11 species; group 4, fig. 2). Predators such as pike-perch, Stizostedion lucioperca, and perch, Perca fluviatilis, occurred, and cold water species were found, such as minnow, Phoxinus phoxinus, or chub, Leuciscus cephalus. When the silt layer was thicker and the water level was intermediate, the specific richness decreased (average, 6,2 species) and the community was either dominated by the cattish (group 2, fig. 2) or by the rudd, Scardinius erythrophtalmus (group 3, fig. 2), according to the importance of the aquatic vegetation cover. Habitats with thickest silt layers, shallowest waters and maximum aquatic vegetation cover contained the poorest communities (average 3,9 species) dominated by eel (group 1, fig. 2). There is also evidence that the diversity of the community has progressively decreased since 1987 (fig. 5). The most stenothermous species disappeared, and the importance of the catfish increased : it doubled between May 1987 and September 1989 (fig. 6). Although the eel is the species most adapted to this environment, we emphasize the diminution of its biomass (fig. 7). These phenomena could be partly due to the climate (cold winter in 1987, important swelling in January 1988 and 2 droughts in summers 1988 and 1989). But they are mainly caused by the water management policy which is intended to favour agriculture by keeping stable water levels (evacuation of swellings) and by preventing the freshwater part from the marine influence (collective sluice gates). This does mot permit an optimal breeding of the species that have to spawn on flooded meadows, neither a proper colonisation of the marsh by elvers.