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Benthic Insects and Fish of the Doubs River System

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Benthic Insects and Fish of the Doubs River System Hydrobiologia 490: 63–74, 2003. 63 © 2003 Kluwer Academic Publishers. Printed in the Netherlands. Benthic insects and fish of the Doubs River system: typological traits and the development of a species continuum in a theoretically extrapolated watercourse J. Verneaux, A. Schmitt, V. Verneaux & C. Prouteau Laboratoire de Biologie et Ecophysiologie -EA 3184 USC INRA. Pôle Hydrobiologie. Institut des Sciences et Techniques de l’Environnement, Universit´e de Franche-Comt´e, Place Leclerc, 25030 Besançon cedex, France Tel: 0381-665-737. Fax: 0381-665-734. E-mail: [email protected] Received 19 June 2001; in revised form 7 August 2002; accepted 20 November 2002 Key words: running waters, insects, fish, typology Abstract Using the distribution patterns of benthic insects (198 species) and fishes ( 29 species) from 11 tributaries and the main channel of the Doubs River drainage basin (French Jura), the authors have tried to establish whether there is an organization of species into discrete, identifiable communities. Principal Component Analysis (PCA) was used to identify whether a continuum existed, and also used to select 50 least-disturbed sites, which were used to define a theoretical watercourse. The density classes of each species were projected on this longitudinal gradient and each species response was characterised by two typological traits: its typological preferendum (tp) and its typological amplitude (ta), thus creating a synthesis of ecological characteristics. In the typological index given in the appendix, the 210 species which form a biological templet are listed in alphabetical order with their tp and ta values. These typological species traits are useful contributions to a database for running waters biomonitoring. Introduction Secondly, the observed succession of species along the downstream course of streams and rivers has long The present work, dealing with typological and bio- been recognised as a means of classifying water- logical qualitative assessment of running waters, was courses (Shelford, 1911; Le Roi, 1913). Fish, different developed following two observations. groups of invertebrates and algae, mosses and higher Firstly, the accelerated transformation of running plants have all been used to identify longitudinal or waters makes interpretation of contemporary biolo- altitudinal regions (zones) in riverine ecosystems al- gical data increasingly difficult. Various authors argue though the validity of pre-established zones has been for the establishment of biological references in every questioned (Macan, 1963; Hynes, 1970; Hawkes, drainage and geographical area (Verneaux, 1973; Fit- 1975). It has been generally concluded that the gen- tkau & Reiss, 1983; Verneaux, 1984, 1994; Moss et eral change in species communities along the rivers al., 1987; Reynoldson et al., 1997, Mouthon, 1999). tends to be not zonation but transition. According to Norris & Thoms (1999) have argued that the ‘poorer the River Continuum Concept, which includes more conditions are usually indicated by a loss of taxa’, and continuous than sectioned species distribution pat- that fish and benthic insects, especially Plecoptera, terns (Maitland, 1966; Ulfstrand, 1968; Verneaux Ephemeroptera and Trichoptera, are frequently con- & Rezzouk, 1974; Vannote et al., 1980; Gendron sidered as suitable indicators. Using reference bio- & Laville, 1997), the running water ecosystem is logical data and comparative analyses, temporal and viewed as an upstream-downstream gradient that can spatial changes in biological structures can be evalu- be modified by various ecological disturbances, of ated. natural or anthropic origin, that induce discontinuit- 64 Table 1. The 12 studied rivers of the Doubs river system. 117 general temperature pattern of the water reaches max- sampling sites - S = springs imum values in the months of July and August during Watercourse River site Length Altitude Drainage area the low-water period . Detailed descriptions of the (code) (km) (m) (km2) geographical area and the various rivers were given Doubs S 1–29 453 940–172 7700 previously (Verneaux, 1973) and we have cited here Drugeon S30–35 33 930–805 185 only main characteristics of the rivers and the code of Dessoubre S36–40 29 600–387 560 the sampling sites (Table 1). Allaine S41–48 34 605–350 230 Audeux S49–53 25 560–280 230 Cusancin S54–57 9 325–280 360 Loue S58–74 122 543–197 1900 Lison S75–79 25 410–293 290 Data and methodology Furieuse S80–88 18 575–251 100 Cuisance S89–96 34 375–205 180 Doulonnes S97–99 7 254–210 20 The data of the water survey, periodically published Clauge S100–105 29 260–195 135 by the Environment Boards, describe a deterioration in the water quality of the Doubs River system dur- ing recent decades (Agence de l’Eau, 1996, 1999). This disturbance was especially obvious in the sal- ies (Verneaux, 1973; Chessel et al., 1987; Angelier, monid rivers which were in good condition in the 2000). Even though it may appear that species arrange 1970s (Verneaux, 1973). The disappearance of many themselves in discrete communities, it is considered species (Bacchi, 1994) led to the scarcity of ref- that conditions and communities vary gradually along erence sites with biotic indices equal to or above a theoretical watercourse or along a model water- 18/20 (IBGN, AFNOR, 1992; Observatoire Régional course, without discontinuities. de l’Environnement, 1995). Because of this general Based on the species recorded at the selected ref- change and given our aim which implies recourse to erence sites along the Doubs River and its tributaries, reference sites, the field data collected from 1970 to the above observations allowed us to establish whether 1976 were selected for study. A major part of the same there was some predictable organization in the species data was recently used by Doledec et al. (2000) to de- found. In order to establish a database which can be scribe the functional diversity of communities in terms useful for comparative analysis, this study presents: a of their biological traits. These data have been updated general biological structure in the form of a species to include recent taxonomic changes. continuum, a theoretical running watercourse, com- The initial density matrix is composed of 227 posed of reference sampling sites from different rivers, rows (198 species of aquatic Insecta including 43 and two typological species traits used to establish a Plecoptera, 46 Ephemeroptera, 109 Trichoptera, 28 general distribution pattern of species. species of fish and one Cyclostoma) and 117 columns (117 sampling sites from the Doubs river and 11 trib- utaries, Fig. 1). Species densities were weighted using Sampling sites five classes of relative density. These classes were ad- apted to the highest values observed in the sample for The Doubs river and its 11 tributaries form a 832 km- each genus of invertebrates and for each species of long network which extends from the Saône river plain fish (Verneaux, 1973). This weighted method, which to the Swiss mountains, in a 7700 km2 drainage basin highlights structures using factorial analyses, has been that overlaps the northern Jura and the southern Vosges justified by other authors (Chessel et al., 1987). Of mountains (Fig. 1). the sites differentiated by factorial analysis, only the The hydrologic conditions in the basin are the sites which had a biotic index (IBG Verneaux, 1982; result of the alternating permeable and impermeable IBGN AFNOR, 1992) above or equal to 18/20 were Jurassic strata and many karstic structures. Except for considered as non-or slightly disturbed from 1970 to some true springs, most springs are karstic outlets. 1976 and were thus selected as reference sites. These Climatic conditions are mostly continental, some- sites had the highest relative species richness along what influenced by oceanic inputs, altitude, orienta- the entire upstream-downstream (U-D) gradient. All tion and morphological characteristics of the valleys. the species included in the matrix were present in the The river flow shows torrential characteristics and the sample of the reference sites. 65 Figure 1. The drainage area of the Doubs river system – distribution of the 117 sampling sites. Al: Allaine, Au: Audeux, Ce: Cuisance, Cl: Clauge, Cu: Cusancin, De: Dessoubre, Dl: Doulonnes, Dr: Drugeon, Fu: Furieuse, Li: Lison. To take into account the density differences (Jack- species were therefore defined using the projection of son, 1973), the weighted data were analysed by their density classes on the theoretical U-D gradient. Principal Component Analysis (PCA) applied to the Two descriptors were used to summarize the dis- covariance matrix using ADE 4 software (Thioulouze tribution of each species on the gradient: its mean et al., 1997). position, or preferendum (tp) and its typological amp- The environmental parameters were not taken into litude (ta). In agreement with Chevenet et al. (1994) account in establishing an environmental gradient, and Mouthon (1999), we chose the mean point of so that preconceptions about their effects on biolo- abundance rather than the point of maximum abund- gical distributions could be avoided. The theoretical ance. Thus, the specified ta values are equal to half the upstream-downstream gradient represented by a suc- difference between the limits of the abundance distri- cession of reference sites, was determined only by us- bution of each species and the tp values refer to the ing the biological data. Following the same principle, centers of gravity of the same abundance distributions Mouthon (1999) divided the longitudinal gradient ac- (Fig. 4). cording to the groups of river sites which were defined using a hierarchical ascending classification (HAC). In the present study, the distances between reference Results sampling sites were evaluated by their angular dis- tances, which provides a valid solution when PCA is Reference river sites and virtual watercourse used (Legendre & Legendre, 1984). The angles were measured from the radius joining the center of the axes The sites situated in peripheral positions in the plane to the virtual spring given by the species richness curve of a PCA (Fig.
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