Ecological and chorotogical variations of macrophytic vegetation in acid rivers of Armorican Massif and Northern Vosges (France)
For the study of the vegetation of acid rivers in France, two areas have been investigated by the authors : Armorican Massif and Northern Vosges (fig. 1). This paper has two aims : - assessing the general trends of macrophytic vegetation : floristic composition and plant communities, with special reference to the influence of physical and water quality parameters, - pointing out and interpreting the variations observed between the two regions (chorological differences), and in the watercourses (ecological variations). Floristic data and phytosociological surveys (from bryophytes to angiosperm macrophytes) were collected along 50 m. stretches (at least) by the stratified sampling method in the case of large watercourses, and by systematic surveys for smaller ones. Consecutive 50 m. long sketches were used to measure the effect of localized pollution. (The inventory is as precise as possible). A distinction was made between the aquatic communities and river-bankside ones. At the same time, physical parameters were measured or observed (depth, light conditions, width,...), certain water-quality parameters were measured (pH, conductivity,...) and water samples were collected for further analysis. The data collected are gathered and compared in a table (table 1) showing differences between aquatic and subaquatic species, and between sunlit and shaded stretches. Trophic tendencies of most species are given according to either our observations or the literature. The main ecological feature is the streaking longitudinal variation of aquatic species, which agrees with the zones described by ILLIES and BOTOSANEANU : crenon, rhitron and potamon. In sunlit areas the succession is dominated by the following species : Potamogeton polygonifolius, Juncus bulbosus, Glyceria fluitans,... -> Ranunculus spp., Callitriche hamulata, Myriophyllum alterniflorum,... ->Potamogeton crispus, Elodea canadensis,... Areas in the shade are characterized by bryophyta; the succession is : Scapania ondulata ->Rhynchostegim riparioïdes, Fontinalis antipyretica As for the aquatic species, a longitudinal succession of bankside species exists : Sparganium erectum, Carex paniculata, Carex rostrata ->Phalaris arundinacea ->Phalaris arundinacea, Typha latifolia The macrophytic vegetation in acid rivers of Western Europe shows a great homogeneity, marked by the constancy of a floristic list, in which some species (Callitriche hamulata, Myriophyllum alterniflorum, Potamogeton polygonifolius, Potamogeton alpinus, Scapania undulata, etc.) only grow into acid waters, while others species (Callitriche platycarpa, Sparganium emersum, Elodea canadensis, Potamogeton crispus, etc.) grow also in calcareous waters. This basic unit is submitted to ecological and chorological variations. Ecological variations are due mainly to changes of the water quality. They are illustrated by two examples : - two river systems in the Northern Vosges where a general survey shows longitudinal variations of communities without pollution or assessing different kinds of pollution, - a pisciculture pollution with observations of species distribution on consecutive sketches. An increase of water trophic level leads progressively from an oligotrophic vegetation (marked by Potamogeton polygonifolius, Juncus bulbosus, etc.) to a meso-eutrophic community (characterized by Callitfriche obtusangula, Nasturtium officinale, etc.). Four communities are distinguished in Northern Vosges (table 2) and characterize water quality (both pollution and trophic level) : A : Potamogeton polygonifolius, Juncus bulbosus, Sphagnum spp. community, with the following main water-quality parameters : conductivity : 40-80 μS, phosphates : <20 ppb P, ammonia : <20 ppb N; B : Potamogeton polygonifolius, Ranunculus peltatus, Callitriche hamulata community, (40-80 μS, 20-40 ppb P, ammonia : <40 ppb N) C : Ranunculus peltatus, Callitriche hamulata, Potamogeton alpinus, Myriophyllum alteriflorum community, (60-120 μS, 40-120 ppb P, ammonia : <200 ppb N) D : Callitriche obtusangula and Nasturtium officinale inside C community list, (80-130 μS, 40-300 ppb P, ammonia : -> 500 ppb N). Two vegetation maps show the distribution of these communities (fig. 2). The succession of communities is related to water quality and fastened sequences are compared to normal ones. Without any pollution, A and B communities caver more than 5 km along the upper streams; under these conditions, the entire sequence A to D develops to 15 km, while with much anthropic pollution, it may occur within less than 3 km. A heavy pollution leads to the leads of communities as observed with the change tram A to C (without any B) due to a camping-site. A localized organic pollution by a Breton pisciculture is responsible for water-quality changes (table 3): ammonia, nitrites, phosphates and conductivity increase. The succession of mineral nitrogen forms is assessed (fig. 3). In the N-NH4+ and N-NO2- peak areas, the eutrophic species increase (Callitriche spp., specially Callitriche obtusangula, Amblystegium riparium) and the oligotrophic ones decrease (Scapania undulata) (fig. 3). Tissue analysis of Ranunculus penicillatus and of Callitriche spp. does not show any enrichment in nitrogen in polluted areas: the ecological response is mainly an increase of nitrophilous macrophytes (Callitriche spp.). The influence of water quality is more obvious in the Northern Vosges for most of the oligotrophic communities grow in forested sketches, white in Brittany these conditions are rare and eutrophication occurs in all the streams, even in the source areas. Floristically, neutrophilous or ubiquitous species are numerous. Some of them indicate either eutrophication or pollution : Amblystegium riparium and perhaps Octodiceras fontanum for bryophyta, Callitriche obtusangula, Potamageton perfoliatus and Sparganium. emersum (long leaves form), for aquatic species, Nasturtium officinale, Glyceria maxima for helophytes. Physical parameters also lead to variations. The succession of swift and slow areas leads to changes in the communities. Thus, in Brittany, Oenanthe crocata and Ranunculus penicillatus are associated on the riffles, while Nuphar lutes and Sparganium emersum characterize the pools, even in the upper parts of the streams where there are watermill ponds. The relationships between local hydrodynamic features and species distribution are discussed with regards to the stretches around the pisciculture area. Main chorological variation is an enrichment of atlantic areas with numerous eu-atlantic or mediterraneo-atlantic species : Porella pinnata and Amblystegium fluviatile as bryophyla, Oenanthe crocata, Apium inundatum, Ranunculus omiophyllus as spermaphyta. As there are few or no species restricted to medio-european areas, Atlantic communities are often richer than eastern ones. But some invading species as Elodea nuttallii are spreading all over the country from North-Eastern France. Ecological trends of certain species (Sparganium emersum, Ranunculus peltatus, Potamogeton polygonifolius) can change between the two biogeographical areas. So, Sparganium emersum grows in the upper oligotrophic zones in Northern Vosges while it characterizes lower meso-eutrophic zones in Brittany. In Brittany, Ranunculus peltatus un exist in eutrophic areas when the stretches are far from the sea. Potamogeton polygonifolius is very rare in swift streams in Brittany. Therefore, macrophytes permit, as benthic invertebrates do, to get a river biodiagnosis. They are easily observed and mapped so they can be used for a general survey.