the rhone river basin - aalto · 2017-01-05 · table 7.1 general characterization of the rh^one...

Chapter 7

The Rhone River Basin

Jean-Michel OlivierUMR CNRS 5023, Universit!e Lyon 1, 43

Boulevard du 11 novembre 1918, F-69622

Villeurbanne Cedex, France

Georges CarrelUR Hydrobiologie, Cemagref, 3275 Route

C!ezanne, CS 40061, F-13182 Aix-en-

Provence Cedex 5, France

Nicolas LamourouxUR Biologie des Ecosyst"emes Aquatiques, 3

bis quai Chauveau, CP 220, F-69336 Lyon

Cedex 09, France

Marie-Jos!e Dole-OlivierUMR CNRS 5023, Universit!e Lyon 1, 43



Florian MalardUMR CNRS 5023, Universit!e Lyon 1, 43



Jean-Paul BravardUniversit!e Lyon 2, Facult!e de G!eographie,

Histoire, Histoire de l’art, Tourisme, 5

Avenue Pierre Mend"es-France, 69676 Bron

Cedex, France

Claude AmorosUMR CNRS 5023, Universit!e Lyon 1, 43



7.1. Introduction7.2. Biogeographic Setting

7.2.1. General Aspects7.2.2. Palaeogeography

7.3. Physiography, Climate and Land Use7.3.1. Landforms and Geology7.3.2. Climate7.3.3. Land-Use Patterns

7.4. Geomorphology, Hydrology, andBiogeochemistry7.4.1. GeomorphicDevelopment of theMain

Corridor7.4.2. Hydrology and Temperature7.4.3. Biogeochemistry7.4.4. Bedload7.4.5. Nutrients and Pollution7.4.6. Heavy Metals and Organic

Micropollutants7.4.7. Pesticides7.4.8. Priority Substances of the Water

Framework Directive7.4.9. Artificial Radionuclides

7.5. Aquatic and Riparian Biodiversity7.5.1. Algae7.5.2. Macrophytes7.5.3. Floodplain Forests7.5.4. Aquatic Invertebrates7.5.5. Non-native Species7.5.6. Protected Species

7.5.7. The Rhone Groundwater System andits Obligate Fauna (Stygobionts)

7.5.8. Fish7.5.9. Fishes in the Swiss Upper Rhone

7.5.10. Fishes in Lake L!eman7.5.11. Fishes in the Downstream From

Geneva and French Upper Rhone7.5.12. Fishes in the French Lower Rhone7.5.13. The Apron7.5.14. Migratory Fishes in the Rhone7.5.15. Amphibians7.5.16. Reptiles7.5.17. Birds7.5.18. Mammals

7.6. Management and Conservation7.6.1. Economic Importance7.6.2. Flood Control7.6.3. Fishery7.6.4. Conservation and Restoration7.6.5. Restoration Activities and Potential7.6.6. EU Water Framework Directive

7.7. The Ain River7.7.1. Geomorphology7.7.2. Hydrology and Temperature7.7.3. Biogeochemistry7.7.4. Aquatic and Riparian Biodiversity7.7.5. Management and Conservation

7.8. The Saone River7.8.1. Geomorphology7.8.2. Hydrology and Temperature

Rivers of EuropeCopyright ! 2009 by Academic Press. Inc. All rights of reproduction in any form reserved. 247

7.8.3. Biogeochemistry7.8.4. Aquatic and Riparian Biodiversity7.8.5. Management and Conservation

7.9. The Durance River7.9.1. Geomorphology7.9.2. Hydrology and Temperature7.9.3. Biogeochemistry, Bedload and

Sediment7.9.4. Nutrients and Pollution7.9.5. Aquatic and Riparian Biodiversity7.9.6. Management and Conservation

7.10. Conclusions and PerspectivesAcknowledgementsReferences

7.1. INTRODUCTION

The Rhone links several valleys and rivers from the upperAlps with the Mediterranean Sea (Figure 7.1, Table 7.1).Although the same pre-Celtic name was given to the riverupstream and downstream of Lake L!eman, the present SaoneRiver and Rhone downstream of Lyon were considered bythe Greeks, Romans, and even recent historians as the maineconomic corridor between the Mediterranean Sea andNorth Sea (Braudel 1986). The Alps provided a strong eco-nomic and cultural unity to the Rhone throughout the lastthousand years. One manifestation of this unity is the use oflanguage since Roman Times; that being French around theMont Blanc massif. Cities such as Sion, Martigny, Geneva,Yenne, Aoste, Lyon, Vienne, Valence, Orange, Avignon andArles were founded by Romans or older groups and werepolitically and economically important to Rome. The Ro-mans were still unified before the short Napoleonic periodthat allowed the conquest of Italy in the late 1790s. In mod-ern times, Switzerland and France have distinguished theupper Rhone River between Lake L!eman and the headwatersin the Valais region of Switzerland and the ‘French UpperRhone’ between Lyon in France and Lake L!eman. The dis-tinction between these two entities on each side of LakeL!eman was reinforced by the harsh topography, poor com-munication along the river, and difficult conditions for nav-igation (Photo 7.1).

Since the late 19th century, embankment policies inValais, hydropower developments in France, and a decreasein traditional navigation and fishery have decayed commu-nication links even further. Today, the Rhone River is ahighly regulated river, viewed as a large waterway for com-merce. Forgotten is the fact that it represents about 60% ofthe freshwater inputs into the Western Mediterranean Sea.The Rhone is a major source of nutrients and particulatematter to the Mediterranean Sea (Alliot et al. 2003), and amajor factor in sole fishery yields from the Gulf of Lyons(Salen-Picard et al. 2002). The public and politicians areincreasingly aware of environmental problems and sociolo-

gists indicate a new interest towards integrating river envir-onments in everyday life, so called ‘river re-appropriation byriverine people’. Recent floods also show that the Rhone,although regulated, can still overflow its banks and pose arisk hazard. Energy production, navigation, irrigation, tour-ism, habitat and biodiversity protection and cultural activi-ties all must be taken into account by river managers andlocal decision-makers. The complex uses of the river havealso caused new initiatives between Switzerland and France.

In this chapter, we present the general physical, chemicaland biological characteristics of the Rhone River. Alltributaries on the Swiss side are alpine rivers and have beenused for water storage and hydroelectricity. Several tributar-ies from the French side play an important role in the Rhonedischarge and are included in this chapter, the rivers Ain,Saone and Durance in particular. The Is"ere River also is animportant tributary, but it was historically affected by regu-lation, channelization and industrialization and will not bediscussed in detail. Lake L!eman is an important feature ofthe river and its effects on the river downstream also arediscussed in this chapter.

7.2. BIOGEOGRAPHIC SETTING

7.2.1. General Aspects

The present morphology of the Rhone valley is a result of theRiss andW#urm glaciations that deepened valleys in the Alps.The post-glacial colonization of the upper Rhone above LakeL!eman by aquatic organisms is still poorly documented, buta possible connection between the Upper Rhone and L!emancatchment and the Rhine catchment is believed to explainfish colonization in the Upper Rhone. Today, the Rhoneflows through three major ecoregions (Alps, Western High-lands and the Mediterranean part of the Western Plains) thatencompass a large geographical gradient in altitude andlatitude.

7.2.2. Palaeogeography

TheUpper Rhone catchment lies inside the tectonic trench ofValais between the high mountains (>4000 m asl) of thecrystalline Pennine Alps in the south and the northern sed-imentary Bernese Alps. The folding of the Alpine Range andthe Jura Mountains during the late Tertiary and Pliocene wasfrom plate tectonics. The folds were pushed westward overthe north–south tectonic trough of the Saone and Rhone, andabutted to the west by the tectonic margin of the MassifCentral plateau from Hercynian folding of crystalline andmetamorphic rocks. The geological history of the Rhonebetween Lake L!eman and the Bas-Dauphin!e foreland ispoorly documented. The present river cuts across anticlinesthrough narrow trenches, following synclines before thesouthern Jura bend. To the west, the valley displays recentpostglacial features. The former river may have flowed

PART | I Rivers of Europe248

FIGURE 7.1 Digital elevation model (upper panel) and drainage network (lower panel) of the Rhone River Basin.

Chapter | 7 The Rhone River Basin 249

TABLE 7.1 General characterization of the Rhone River Basin

Upper Rhone Rhone Ain Saone Is"ere Durance

Mean catchment elevation (m) 1655 699 669 373 1463 1137Catchment area (km2) 8018 90 538 3713 29 498 11 865 14 322Mean annual discharge (km3) 5.77 53.61 3.88 14.92 10.63 0.82Mean annual precipitation (cm) 162.4 103.1 101.0 95.2 130.6 107.9Mean air temperature (!C) 4.3 9.2 9.2 9.6 6.5 8.8Number of ecological regions 2 3 1 1 3 2Dominant ("25%) ecological regions 2 48; 70 70 70 2 2; 48

Land use (% of catchment)Urban 3.6 4.3 3.6 4.9 2.9 1.3Arable 10.6 30.1 16.7 38.0 13.6 20.5Pasture 5.2 10.7 20.0 20.3 6.2 1.7Forest 23.6 36.7 53.8 34.4 34.4 38.0Natural grassland 16.8 10.7 4.0 1.4 19.3 23.9Sparse vegetation 32.5 6.0 0.1 0.0 22.9 14.1Wetland 0.1 0.6 0.5 0.1 0.0 0.0Freshwater bodies 7.6 0.9 1.3 0.9 0.7 0.5

Protected area (% of catchment) 1.4 9.5 1.1 0.4 31.9 12.2

Water stress (1–3)1995 1.2 1.1 1.0 1.1 1.0 1.02070 1.2 1.1 1.0 1.1 1.0 1.0

Fragmentation (1–3) 3 3 3 3 3 3Number of large dams (>15 m) 0 6 5 0 2 9Native fish species 17 35 27 30 21 27Non-native fish species 5 16 11 11 9 16Large cities (>100 000) 2 1 0 2 1 0Human population density (people/km2) 190 98 61 94 82 32Annual gross domestic product ($ per person) 26 721 24 262 23 298 24 410 25 506 22 787

For data sources and detailed explanation see Chapter 1.

PHOTO 7.1 Pfynwald in the UpperRhone valley in Switzerland (Photo: A.Peter). This is the largest dynamic sectionremaining along the Swiss Rhone.


across the Jura Mountains or entered the Is"ere valley throughthe present Lake Bourget.

The Saone-Rhone trough was a tectonic rift in the earlyTertiary, and was earlier connected to the Rhine rift. This riftshunted the Rhine River south towards Basel before creationof the present flow of the Rhine to the North Sea. Thenorthern part of the Saone-Rhone rift is presently drainedby the Saone River. The middle part of the rift, occupied bythe Bas-Dauphin!e foreland between Lyon and Valence, is alarge alluvial fan of sediments originating from the formerUpper Rhone and Is"ere catchments. It was incorporated intothe Alpine piedmont during the late Tertiary. The Rhone thenincised a course through the hard rocks of the Massif Centraland the soft rocks of the Alpine piedmont, alternating be-tween incised valleys and wide floodplain basins. The inci-sion of the river also was influenced by the lowering of theMediterranean Sea during the closure of Gibraltar Narrows,the so-called Messinian Crisis in the Pliocene between 5.3and 5.9 million years ago (Gargani 2004). The re-connectionof the Atlantic Ocean and Mediterranean Sea inundated theRhone gorge and deposited Plaisancian clay up to Lyon (1.8–4.9 million years ago). The level of the Mediterranean Seadecreased by about 120 m during the Quaternary glaciation,changing the longitudinal profile of the Rhone several times.The present lower Rhone, including the Camargue delta, wasbuilt up by sediments from tributaries and eroded fluvio-glacial terraces of the W#urm glacier that covered part ofthe Bas-Dauphin!e foreland.

7.3. PHYSIOGRAPHY, CLIMATE ANDLAND USE

7.3.1. Landforms and Geology

The Rhone is an 8th order 812 km long river, of which512 km flows downstream from Lake L!eman. Its catchmentcovers about 98 500 km2 with 90 500 km2 of this in France.It is mainly an Alpine river with 50% of the catchment above500 m asl and 15% above 1500 m asl. The river originatesfrom the Furka Glacier (2341 m asl), and drains the Valaisregion in Switzerland. Downstream from Lake L!eman, itcrosses the folded area between the Alps and Jura mountains,flowing north of the Bas-Dauphin!e alluvial fan. West of theJura, the Rhone receives the Ain River, a right bank tributary.In Lyon (163 m asl), it receives the Saone River that flowsfrom the Vosges mountains and drains the Bresse andDombes plains. Downstream of Lyon, the river flows south-ward to the Mediterranean Sea. Tributaries flowing from theAlps such as the Is"ere and Durance are longer relative to theshort steep rivers flowing from the Massif Central. The lon-gest western tributaries are the Ard"eche and Gard in thesouthern catchment. Although the western rim of the formerrift is made of acidic rocks, most of the catchment is com-posed of alkaline rocks of Jurassic and Cretaceous sedimentsand Tertiary and Quaternary deposits.

7.3.2. Climate

The Rhone catchment includes several climatic zones ofmostly oceanic influence with moderate precipitation in allseasons. Northern areas and mountains receive the highestprecipitation and altitude affects the duration and thicknessof snow cover. Eastern areas have summer storms of conti-nental influence, and cold winter temperatures occur in theSaone valley. The southern part has a Mediterranean climatewith hot and dry summers and rainfall mostly in spring andautumn. Total precipitation is around 600 mm/year in thesouthern valley, but rainfall can be intense (>600 mm indays) in the C!evennes Range (1600 m asl) in the southeastof the catchment in September and October. This area isinfluenced by warm, low-pressure air masses that flow overthe northern Mediterranean Sea. The Valais in Switzerland,some inner Alpine valleys such as the Upper Is"ere and ArcRiver, and the Rhone valley south of Lyon have lower rainfalland are influenced by northern ‘Mistral’ wind.

7.3.3. Land-Use Patterns

The Rhone catchment has a long history of human influencewith agriculture and livestock breeding being active #6000years ago. Archaeological studies have documented severalperiods of land clearance and land abandonment in the mid-dle Rhone valley (Van Der Leeuw 2005) that are associatedwith periods of soil erosion and of soil and forest recovery.The Roman and Medieval periods had warmer climates withextensive agriculture use in valleys and higher altitudes. TheLittle Ice Agewas a period of increasing population sizewithdensities averaging 100 inhabitants per km2 in mountainareas. High population densities occurred from 1840 to1870 before increased emigration to cities and lower popula-tions after the First World War. Highest deforestation in theAlps and Massif Central occurred during this populationmaximum. Today, over 30% of the Rhone basin is forestedand#80% of the production is used for timber. The types offorest range from coniferous (Abies sp., Picea sp.), hard-wood (Quercus sp., Fagus sylvatica), and poplar plantationsin alluvial valleys to fire-prone Mediterranean forest (mostlyPinus halepensis) developing on abandoned agriculturallands (Web site: EUFIRELAB, Laboratory forWildland FireSciences and Technologies in the Euro-Mediterranean Re-gion) (Curt et al. 2004).

Present land use in the catchment is diverse because ofpedologic, climatic and geomorphologic differences.Marked regional differences are associated with specificeconomic features and irrigation in alluvial valleys, in par-ticular around the Mediterranean area. Mountain areas havebeen used mostly for livestock breeding and dairy produc-tion, but have been widely reforested during the last century.World-renowned vineyards are common on well-exposedslopes (Dangreaux 2002). The plains and valleys are richagricultural areas, producing corn and cereals. Orchards,mostly apricot, peach and cherry, also have increased


because of new irrigation practices, and often extend intofloodplains. Perfume and medicinal plants are produced inthe southern Rhone, and walnuts are grown in the Is"ere Rivervalley (ACO ‘Noix deGrenoble’). Rice production occurs onthe Camargue delta, and fodder is an important irrigationcrop in the south (ACO of ‘La Crau’ fodder). The north–south orientation of the catchment influences irrigationdemands with a threefold difference between the upperSaone basin and the Mediterranean area. Agriculture used2.8 billion m3 of surface water and 196 million m3 ofgroundwater in 2001 (DIREN 2005).

7.4. GEOMORPHOLOGY, HYDROLOGY,AND BIOGEOCHEMISTRY

7.4.1. Geomorphic Development of the MainCorridor

The Upper Rhone in Switzerland drains steep headwatersand eventually flows through a large alluvial plain that entersLake L!eman. The average slope between the source andLake L!eman (164 km) is #0.9%. After the 1860 and 1861floods, high levees and spurs constricting the riverbed wereinstalled along the Upper Rhone. Between 1928 and 1956,the river channel was reshaped to be narrower and deeper,and the alluvial plain was dredged and developed (Meileet al. 2006). Another plan in the early 1980s (the so-calledHydro-Rhone project) aimed at developing hydropower witha series of 10 dams, but it was not implemented. The last‘Rhone correction’ was begun in 2000 after the large 1987and 1993 floods that threatened the valley. The main purposeis to widen the river and increase channel capacity, securelevees, and improve environmental quality (HYDRONAT2000).

The Rhone contributes 68% of the total water and pro-vides suspended sediments into Lake L!eman (Web site:EUROLAKES, Integrated Water Resource Managementfor Important Deep European Lakes and their CatchmentAreas). Lake L!eman is a warm monomictic lake of glacialorigin created by a moraine dam at its western end. It has twobasins: the eastern ‘Grand Lac’ (499 km2, maximum depth310 m) and thewestern ‘Petit Lac’ (81 km2, maximum depth76 m) (Web site of the International Commission for theProtection of Lake L!eman, CIPEL).

In Geneva, the Arve River enters the Rhone from theFrench Alps, delivering #500 000 tons/year of suspendedsediment. Downstream of Geneva, the Rhone flows througha gorge in the lower Savoy plateau, then it widens across anarrow alluvial plain built up by sediments from the Arveand Fier rivers and small tributaries from the JuraMountains.Upstream of Lake Bourget, the Chautagne and Lavoursmarshes are large peaty swamps originating in the earlyAtlantic period. The Basses Terres area at the Rhone bendnear the southern Jura Mountains has several preserved me-ander scars dating back to 7000 years BP (Salvador 1999,

Bravard et al. 2002b) and a meander belt dated from the lateRoman period (4th century AD). During the Little Ice Age,the rivermorphology changed frommeandering to braided inresponse to an increased bedload from the Alps and moreintense floods. The river downstream of Basses Terres isstable, straight and has a low gradient up to the confluencewith the Ain River. Downstream of the Ain, the Rhone passesthe area beyond the W#urmian front morain of 18 000 yearsago. Here is a wide, steep alluvial plain (slope of the riv-er = 0.1%) with low terraces dating back #10 000 years BP,Holocene meander scars, and the remnants of a large braidedbelt from the Little Ice Age. Archaeologists and geomor-phologists have reconstructed the past history of the rivernear Lyon, recording a braided belt dated 2800–2400 BP,Roman and Medieval meander scars, and a period of in-creased flooding during the 1st and 2nd century AD (Salva-dor et al. 2002).

Historically, the Upper Rhone in France was developedbecause it was used for navigation, mostly for the transportof stone and wood. After 1830, steamboats carried passen-gers between Lyon and the thermal resort of Aix-les-Bainson the eastern shore of Lake Bourget. Before the first rail-ways in 1848, this reach was further developed to improvenavigation. Completion of low embankments was completedin 1886 and river traffic slowly vanished. Public interestsmostly involved flood protection measures. The large floodsof 1840 and 1856 destroyed parts of Lyon that were locatedin the alluvial plain. The alluvial plains and Lake Bourgetwere recognised in 1857 as important water storage areas,and in 1858 a law forbid any embankments upstream of largecities such as Lyon. Consequently, all floodplains upstreamof Lyon are still flooded by high flows (285 km2), exceptareas affected by recent construction of hydropowerschemes.

The Rhone upstream of Lyon has been developed forenergy production periodically between the end of the 19thcentury up to the 1980s (Table 7.2). First, dams were builtnear large cities such as Geneva (1884) and upstream ofLyon (1899), with the largest diversion dam built in Europeat that time. Next, upstream gorges were dammed by theChancy-Pougny (1925), Verbois (1943) and G!enissiat(1948) dams. A nuclear power plant called Bugey was builtin the 1970s. An experimental fast-breeder reactor was runin Creys-Malville after 1986 but it is being dismantled.Last, following the high energy demand after the 1973oil crisis, the National Rhone Company (CNR) built aseries of hydropower developments: Chautagne (1981),Belley (1982), Br!egnier-Cordon (1984) and Sault-Br!enaz(1986). These include a diversion dam that raises theheight of the river and a power plant that uses the divertedflow. The old riverbed is used to accommodate flood flowsthat exceed the maximum operating flow of the plant(Photo 7.2). These large dams have impacted the river inmany ways, but especially in the amount of stored sedi-ments that must be flushed through the system at periodicintervals (Roux 1984).


Downstream of Lyon, the Rhone flows north and southtowards theMediterranean Sea. The geomorphic features areless complex along this reach because of no glacial influenceexcept during the G#unz glaciation. River slope averages

0.05% and is controlled partly by bedload transport, sedi-ment inputs from tributaries such as the Is"ere, and bedrockoutcrops. Donz"ere trough, south of Mont!elimar, is a narrowreach #1 km wide shaped during the Pliocenic Sea

TABLE 7.2 Hydroelectric development scheme along the Rhone River

Name Year ofcompletion

Fall (m) Reservoirlength (km)

Maximumdischargeprocessed(m3/s)

Maximumhydropowercapacity(MW)

Meanproduction/year(Gwh/year)

By-passedsectionlength(km)

Lock

Chippis 1911 88 – 65 41 245 NoLavey 1950 34–42 – 220 69 400 NoSeujet 1995 3 – 550 6 20 YesVerbois 1943 34 11.4 530 100 466 NoChancy-Pougny 1925 10.7 9 520 38 210 NoG!enissiat 1948 67 23 750 420 1700 NoSeyssel 1951 8.5 5 600 42 165 NoChautagne 1980 15 5 700 90 454 8.9 NoBelley 1982 17 5 700 90 449 17.2 NoBr!egnier-Cordon 1984 13.7 12 700 70 324 11.4 NoSault-Br!enaz 1986 9.7 28 700 45 245 1.8 NoCusset 1899 12.2 19 600 63 410 18 NoPierre-B!enite 1966 8 11.2 1380 80 535 9.8 YesVaugris 1980 6.7 19.5 1360 72 335 – YesP!eage-de-Roussillon 1977 12.2 15.7 1600 168 850 12 YesSaint-Vallier 1971 11.5 19.5 1650 120 700 3.7 YesBourg-l"es-Valence 1968 11.7 11.1 2300 186 1100 8.9 YesBeauchastel 1963 11.8 15 2100 192 1200 6.4 YesBaix-Le-Logis-Neuf 1960 12 8.8 2260 210 1200 8 YesMont!elimar 1957 16.5 9 1850 270 1600 13 YesDonz"ere-Mondragon 1952 22.5 28 1970 354 2140 29 YesCaderousse 1975 8.6 11.5 2400 190 840 5 YesAvignon 1973 10 12 2400 200 935 2.6 YesVallabr"egues 1970 13.5 18 2200 210 1300 6.3 Yes

PHOTO 7.2 Rhone River: Br!egnier-Cordon by-passed section (Photo: J.M.Olivier).


regression. The Rhone becomes wider as it flows across theValence and Orange plains. The geomorphology of the mid-dle Rhone between Lyon and Donz"ere are poorly documen-ted, and extensive economic development has stronglymodified the river landscape. The alluvial plains still showsome Holocene meander scars along their margins. Beforemajor developments in the 19th century, the Rhone was abraided river from Lyon to the sea and every tributary fromthe Massif Central or the Alps delivered large volumes ofcoarse sediments during floods. Erosion also was intense inthe deforested watersheds. During the Little Ice Age, braid-ing also dominated the Grand Rhone along the Camarguedelta to its outlet into the Mediterranean Sea (Arnaud-Fas-setta and Provansal 1999), and deposited sediments wereredistributed along the shoreline by storms. Bedload datafor the river indicate a reduction from 22 to 30 Mt/year atthe beginning of the 20th century to 7.4–9.6 Mt/year at theend of this century (Pont 1997; Pont et al. 2002; Antonelli &Provansal 2002). A decrease in annual flood frequency at theend of the Little Ice Age caused this bedload reduction, alongwith changes in land-use mentioned above.

The Camargue delta is a vast alluvial plain of#1450 km2

with numerous ponds and lagoons (Pineau and Sinnassamy2001). The substrate is a thick layer of silt-clay alluvium andsand covering a shingle base from the Pleistocene (2 millionyears). The delta is a naturally dynamic and relatively recentsedimentary construction (developed since the eustatic riseof the Mediterranean Sea ca. 7000–6000 BP). Between 1895and 1944, the delta area increased by #3.9 km2, then itdecreased by #1.7 km2 between 1944 and 1990, and it in-creased again by #0.2 km2 between 1990 and 2000.Changes of the littoral fringe are associated with seasonal

variation in sediment inputs from the river. The littoraladvances during high flows with high sediment loads andretreats during low flows with low sediment loads (Sabatier& Suanez 2003). The construction of coastal protectionstructures in the 1980s has limited coastal erosion (Provansal2003).

Downstream of Lyon, the Rhone has large fertile alluvialplains protected since the late 18th century in some placesand since the early 19th century in most areas. Farms andadjacent lands have been protected from floods by earthenlevees built on the margin of and in the braided belt itself, butthis did not prevent flooding from downstream. Cities likeAvignon were prone to flooding, despite thick city walls.The lower Rhone was developed for navigation after 1840using a complex set of low dykes that maintained constantdepth at low flow. Henri Girardon was the most notedengineer. Some river transport still occurred in the 20thcentury despite competition by two railway lines. In the early1950s, management of the river by the CNR changed to a‘triple target’; that is navigation, energy production and ir-rigation (‘Rhone Law’ of 1921). From 1952 to 1980, 12hydropower schemes were built between Lyon and the GardRiver, producing 12 500 GWh/year. Four other nuclear pow-er plants were built from 1980 to 1987: Saint-Alban (twoPWR units of 1300 MWE), Cruas-Meysse (four PWR unitsof 900 MWE) and Tricastin (four PWR units of 900 MWE)(Table 7.2).

The various river impoundments modified the naturalslope of the Rhone (Figure 7.2). A system of low dykesconstrained the width of the former braided Rhone into anarrow channel, while floods deposited sand and silt on rivermargins (i.e. on the former braided belt) that then became

PHOTO 7.3 Rhone River: Lone deBrotalet (restored side arm at Chautagne)(Photo: J.M. Olivier).


densely vegetated with willows, poplars and maples. Formerriver arms were affected by heavy siltation, causing a loss inecological diversity, and the hydraulics of floods changedsince the capacity of the river decreased. In the mid-1980s,the CNR started a program to restore the connectivity of thedead arms. This program was enlarged to include ecologicalaspects in 1994 when a 10-year master plan for hydraulic andecological restoration of the Rhone was launched and fi-nanced by the Water Agency, the Ministry of Environment,and the CNR.

7.4.2. Hydrology and Temperature

The Rhone has an annual mean discharge of 1720 m3/s at itsmouth, contributing #40% (with the Po River) of the fresh-water inputs into the Mediterranean Sea (Poussard and

Madrid 1999). The Rhone flow regime and its specific dis-charge of 17.8 L/s/km2 result from continuous mixing ofglacier-melt, snowmelt and rainfall (Vivian 1989). Thecatchment can be divided into four large hydrological water-sheds. The Swiss watershed from the source to the LakeL!eman is dominated by high mountains averaging1630 m asl and a glaciated area of 9.4%. Around 50% ofthe annual precipitation comes as snow. The flow regime ofthe Swiss Upper Rhone is characterized by low flows duringwinter from November to April and high flows in late springand summer due to snowmelt (Table 7.3, Figure 7.3). TheFrench Upper Rhone between Lake L!eman and the Saoneconfluence (length #200 km, mean slope #0.1%) has fourmajor tributaries: the Arve flows from the Mont-Blanc rangeand joins the Rhone in Geneva; the Fier and the Guiers flowfrom the foothills of the Alps; and the Ain flows from Jura.Lake L!eman mitigates flood discharges downstream of itsoutlet. For example, large floods from the Swiss UpperRhone can reach 1000 m3/s, whereas flow at the outlet wouldnot exceed 700 m3/s. Tributaries from the French Alps andJura have different flow regimes with high flows in spring(Fier and Guiers), summer (Arve River) and winter (AinRiver).

The middle Rhone from the Saone River confluence(163 m asl) to the Eyrieux confluence (92 m asl) (length#120 km, mean slope #0.05%) has two main tributaries:the Saone and the Is"ere. The Saone, the only typical ‘lowlandriver’ among the Rhone tributaries (mean slope 0.014%),flows from the north and has a typical oceanic pluvial regimewith a high winter discharge and low discharge in summer.

TABLE 7.3 Flow regime (in m3/s) of the Rhone River and its main tributaries

River Station Period Data origin Altitude A MQ SpQ NQ HQ Q2 Q10 Q50

Rhone Sion 1916–2003 1 484 3349 112 33.4 290 910 461 639 808Rhone Porte-de-Scex 1935–2003 1 377 5220 183 35.1 350 1370 629 867 1091Rhone Pougny 1925–2005 2 332 10 320 336 32.6 110 1440 990 1300 1600Rhone Lagnieu 1920–2005 2 192 15 400 457 29.9 160 2440 1300 1900 2300Rhone Perrache 1920–2005 2 162 20 300 598 29.5 190 4250 2100 3000 3800Rhone Ternay 1920–2006 2 158 50 560 1030 20.4 280 4780 3100 4500 5700Rhone Valence 1920–2005 2 102 66 450 1410 21.2 410 6700 3000 6000 8600Rhone Viviers 1920–2005 2 64 70 900 1490 21 450 7990 4300 6100 7600Rhone Beaucaire 1920–2005 2 6 96 500 1700 17.6 490 7800 5900 8300 10 000Ain Chazey Sur Ain 1959–2006 3 210 3630 123 33.8 11 1910 920 1500 2000Saone Monthureux-sur-Saone 1987–2006 3 250 228 3.25 14.2 0.3 155 77 140 110Saone Lechatelet 1965–2006 3 175 11 660 158 13.6 16 1650 830 1300 1700Saone Macon 1952–2006 3 168 25 690 403 15.7 40 2540 1700 2400 3000Doubs Neublans 1966–2006 3 181 7290 175 24 16 1760 1100 1500 1800Is"ere Moutiers 1903–2006 3 474 907 27.6 30.5 7.2 236 160 310 440Is"ere Beaumont-Monteux 1956–2002 2 116 11 800 337 28.6 23 2050 1200 1700 2200Drome Saillans 1910–2006 3 263 194 18.1 15.8 1.6 692 220 380 520Durance L’Argenti"ere 1910–2006 4 950 984 27.6 28.1 6.3 276 140 210 260Durance Meyrargues 1994–2006 4 186 12 500 26.1 2.2 6.2 2800 1000 2300 –

A: Catchment area upstream of gauging station. MQ: arithmetic mean annual discharge. SpQ: specific discharge (m3/s/km2). NQ: lowest measured discharge.HQ: highest measured discharge (instantaneous value). Q2: magnitude of a 2-year flood. Q10: magnitude of a 10-year flood. Q50: magnitude of a 50-year flood.Data origin: (1) Federal Office for Water and Geology (http://www.hydrodaten.admin.ch/), (2) Compagnie Nationale du Rhone, (3) Direction R!egionale del’Environnement Rhone-Alpes, (4) Direction R!egionale de l’Environnement Provence-Alpes-Cote d’Azur (French data available at http://www.hydro.eaufrance.fr/accueil.html) discharges are expressed in m3/s.

FIGURE 7.2 Longitudinal profile of the Rone River bed before and afterthe construction of a chain hydropower plants.


http://www.hydro.eaufrance.fr/accueil.html

http://www.hydrodaten.admin.ch/

http://www.hydro.eaufrance.fr/accueil.html

The Is"ere River has an alpine character with high spring andearly summer discharge from snowmelt. This watershed isaffected by water retention to meet peak energy demandsduring the cold season. The lower Rhone downstream fromValence has tributaries from the C!evennes Mountains thatdrain the eastern Massif Central (Eyrieux, Ard"eche, C"ezeand Gard) and from the Alps (Drome, Ouv"eze, Eygues andDurance). Watersheds of the southern tributaries have aMediterranean climate with high discharge during fall andspring, the Durance River in particular.Most of the dischargein these southern tributaries comes as floods that occur main-ly in September and October.

The area specific discharge of the Rhone decreases alongthe Swiss Upper Rhone, is relatively uniform from LakeL!eman to Lyon, decreases severely after the Saone conflu-ence, and is relatively stable to the delta (Figure 7.4).

At the mouth, the Rhone shows a high seasonal andannual regularity in flow (see Figure 7.3). The oceanic andMediterranean climate of the Rhone causes four main typesof floods: (1) oceanic, (2)Mediterranean, (3) extensiveMed-iterranean, and (4) general floods. The first type of floodresults from high oceanic rainfalls in Upper Rhone tributar-ies with relatively high probability of occurrence (40–60%).

The second type of floods are linked to Mediterranean rain-falls from the south–west (tributaries in the C!evennes area)or from Mediterranean tributaries. At times, Mediterraneanrainfalls reach the Saone, Is"ere and Ain watersheds causingthe third type of flood with low probability (#10%). Thefourth type of flood has intermediate probability (15–25%)and comes from oceanic and Mediterranean rainfall.

Downstream from Valence, tributaries of the lowerRhone often experience severe flash floods, the most de-structive natural hazard in thewesternMediterranean. Heavyrainfalls common in late summer and early fall are linked tothe proximity of the Mediterranean Sea, and meteorologicaland relief attributes. The C!evennes-Vivarais region in thesouthern Massif Central is especially affected by extremerainfalls that cause floods in the C"eze, Ard"eche and Gardrivers (Delrieu et al. 2005). These rains, called ‘c!evenoles’,can influence large regional areas as, for example, the floodin September 1890 in southern France on the Ard"eche at6500 m3/s at Vallon Pont d’Arc in the Ard"eche gorges (Parde1925). Mediterranean floods can extend far upstream as inNovember 1935, November 1951 and December 2003, andgenerate high discharges in the lower Rhone as in October1993 (9700 m3/s at Beaucaire, 25-year flood). Generalfloods result from different combinations of high rains, suchas those responsible for the floods at Beaucaire in October1840 (13 000 m3/s), May 1856 (12 500 m3/s) and January1994 (11 000 m3/s, 70-year flood).

Hydropower plants on the Rhone and high-head retentionschemes in the catchment have greatly modified the flowregime of the river (Vivian 1989). On the Swiss UpperRhone, peak electricity demand increased winter flows by20–150% and decreased summer flows by 15–30%. Flowsnow have a well-defined weekly cycle as well as sudden andfrequent daily flow oscillations (Meile et al. 2006) that alsoinfluence water temperature (Meier et al. 2004) and sus-pended sediments (Loizeau and Dominik 2000). The levelof Lake L!eman and the Rhone discharge at the outlet of thelake are regulated at Seujet dam in Geneva. Downstream ofthe Arve, discharge fluctuates according to the functioning(hydropeaking) of the Verbois, Chancy-Pougny andG!enissiat dams. Discharge from G!enissiat (gauging stationat Surjoux) shows severe daily variations ranging from 0 to700 m3/s, which are partly attenuated downstream by theSeyssel compensation dam.

Upstream of Lake L!eman, the annual mean temperatureis 6.9 !C at Sion (period 1974–2005, range $1.1 to 13.7 !C)and 7.2 !C at Porte-du-Scex (period 1971–2005, range 0.7–14.1 !C). The large volume and long residence time of LakeL!eman warms surface waters to an annual mean temperatureof 12.5 !C at Geneva (period 2003–2005, range 2.8–27.3 !C). Sudden and short decreases in water temperatureare recorded regularly at the outlet of Lake L!eman, especial-ly in summer with the renewal of water in the epilimnion inthe ‘Petit Lac’ by seiches induced by strong winds (Guilbaudet al. 2004). Temperature decreases can reach 9 !C and prop-agate downstream of Lyon.

FIGURE 7.4 Mean annual discharge (Q) and specific discharge (SpQ)along the Rhone River.

FIGURE 7.3 Mean monthly discharge of the Rhone River at sevengauging stations.


Twelve measuring stations are distributed along theFrench Rhone from the French-Swiss border at Pougny toAramon downstream of the Durance confluence. Three sta-tions are on the Ain, Saone and Is"ere. Long-term recordsshow that the Rhone temperature and air temperature are notbalanced. Water temperature depends on five main factors:(1) the upstream-downstream transfer of temperature of wa-ter flowing from the epilimnion of Lake L!eman, (2) meteo-rological conditions especially when significant gapsbetween air and water temperature occur, (3) main tributariesthat modify the water temperature below the confluence, (4)nuclear power stations that typically increase temperaturesdownstream by 1–2 !C, and (5) by-pass sections with lowregulated dischargewherewater temperature increases fasterthan in artificial channels.

From 1977 to 2005, the annual mean water temperatureof the Rhone was 10.9 !C at Pougny and 14.3 !C at Aramon.Temperature increases on average 2.4 !C in the Upper Rhonebetween the Arve and Saone and 1.2 !C from the Is"ere to theDurance River. The mean daily range in temperature in theRhone is#0.5 !C. The Ain is a cold tributary in winter but awarm tributary in summer (annual mean 11.4 !C, 1977–2005). The Saone is a warm tributary (annual mean13.4 !C, 1977–2005). The Is"ere is an alpine cold tributary(annual mean 10.5 !C, 1977–2005) that decreases the annualmean temperature in the Rhone by 0.2–1 !C during thewarmseason. Warmest temperatures occur in August and the low-est in January. From 1977 to 2005, water temperatures in theRhone and tributaries have increased, ranging from 0.7 (Is"ereRiver) to 1.8 !C (Rhone River at Aramon) The French UpperRhone now experiences temperatures that were found in thelower Rhone in the past, although the temperature increasewas higher in the lower Rhone. The temperature in LakeL!eman (data CIPEL) has increased by 1 !C at 5 m depth inthe last 30 years, and temperatures below 100 m depth arenow reaching 6 !C.

Climate change has increased temperatures duringspring, summer and winter, shortening the cold season andlengthening the warm season. For instance, the temperaturethreshold of 18 !C was reached 70 days earlier in 2003 thanin 1978 at Bugey 40 km upstream of Lyon.

7.4.3. Biogeochemistry

The geochemical characteristics of the Rhone were de-scribed by H.L. Golterman (1985a, 1985b, 1985c, 1985d).The Rhone has hard waters with increasing calcium con-centration and a small decrease in pH from the source tothe delta. It has high sulphate, chloride and phosphateconcentrations (Table 7.4). The Rhone loses its glacialsilt and some major changes occur in calcium-carbonatedynamics in Lake L!eman. Lake L!eman has a theoretical11.4-year water residence time that has a clear bufferingeffect on major elements. The lake outlet contributes non-reactive ions such as sulphates or magnesium and shows

strong seasonal patterns in silica, bicarbonates and calci-um. Downstream tributaries progressively modify theseconcentrations, depending on water origin and hydrolog-ical regimes. For instance, the Upper Rhone hydrologicalregime is defined by alpine components that shift to acomplex regime influenced by the pluvial-oceanic regimeof the Saone. The Saone increases the inputs of calcium,sodium, bicarbonates, sulphates, chlorides and nitrates inthe Rhone (Table 7.4), despite the inflow of the Is"ere withmore alpine characteristics.

The Alpine mountain range in the eastern catchmentdrained by the Rhone, Is"ere and Durance consists mainlyof sedimentary rocks with some siliceous crystalline andmetamorphic rocks in the inner Alps. The Jura and Vosgesmountains in the north in the Saone watershed are mainlycalcareous. Crystalline siliceous rocks dominate thesouthern Massif Central (C!evennes) in the south–westRhone basin (Gard, C"eze and Ard"eche). The Rhone alsois a large submontane alluvial river where upland/streaminteractions are important, exemplified by: (1) a develop-ment of a lateral aquifer >15 m deep upstream of Lyon,(2) large floodplains with numerous side arms althoughreduced by regulation, and (3) coarse sediments with highpermeability. This hydrogeological and geomorphologicalcontext contributes to the Rhone nutrient budget, espe-cially during low flows (Dahm et al. 1998). The naturalchemical composition of each watershed (Meybeck 1986),mixed origin of waters, hydrological features in the basin,and major anthropogenic inputs explain the high localvariability in water quality. For instance, the high chlorideconcentration in the Rhone at Geneva is directly linked tothe use of salt on roads during winter. Further, trace ele-ments (As, Sb, Ni and Ba) in the Rhone at Arles showincreased concentrations during floods in the Gard andC"eze rivers that drain silicate mountains and old minetailings (Ollivier et al. 2006).

7.4.4. Bedload

The main tributaries that provide suspended and bedloadsediments to the Rhone are upstream of the Ard"eche, exceptfor the Durance. A total of 800 000–900 000 m3/year ofsediments are poured into the Rhone. The Arve, Ain, Is"ereand Durance provide 75% of this sediment load, and thesediment transport velocity is estimated at 2 km/year inthe Rhone. The mean long-term annual sediment load inthe Rhone at Arles from 1967–1996 was 7.4 Mt, althoughrecent studies suggest that the annual mean value for the last40 years could have been between 9 and 10.1 Mt (Antonelli& Provansal 2002; Pont et al. 2002). Annual values werehighly variable, ranging from 1.2 Mt in a low flow year(Q = 1192 m3/s, 1973) to 19.7 Mt in a high flow year(Q = 2175 m3/s, 1994, 50 days of flooding). The highestestimated values during high flow years were 22 Mt (Surell1847) and 30 Mt (Parde 1925). Present values are mainly due


TABLE 7.4 Average main chemical characteristics of the Rhone River from upstream to downstream and some of its major tributaries (1985–2004)

River Station Altitude W (km2) (mS) pH Ca2+ Mg2+ Na+ K+ HCO$ SO42$ Cl$ NO3

$ NH4+ PO4

3$ BOD5 DOC

Rhone River Porte-du-Scex 377 5220 288 8.05 41.1 5.8 6.6 1.5 87.5 53.2 8.9 2.64 0.03 0.96Gen"eve 369 7987 287 7.98 42.8 6.1 4.9 1.4 46.0 6.6 1.72 0.03 0.05Chancy 347 10 294 308 8.06 47.4 6.5 5.7 1.6 119.3 45.6 7.9 2.77 0.11 1.53Pougny 342 10 300 302 8.10 47.3 6.0 5.4 1.6 113.8 45.8 7.6 2.29 0.21 0.11 1.77 1.58Murs-et-Geligneux 218 13 960 318 8.07 51.3 5.8 5.4 1.5 131.8 40.3 7.9 2.80 0.17 0.13 1.50 1.71St-Sorlin-en-Bugey 193 15 400 325 8.09 52.7 5.8 5.3 1.5 137.3 36.7 8.1 3.16 0.12 0.10 1.48 1.83Jons 180 20 300 337 8.09 58.3 5.2 5.1 1.4 158.8 32.0 7.5 3.92 0.11 0.12 1.30 1.80Chasse-sur-Rhone 150 51 080 400 8.03 66.3 4.9 11.0 2.0 175.1 33.3 18.7 6.11 0.25 0.21 1.76 2.43Saint-Vallier 120 54 650 415 7.98 68.9 5.0 12.1 2.0 174.1 38.9 20.2 6.88 0.31 0.39 1.56 2.38Charm es-sur-Rhone 103 66 450 428 7.98 71.8 6.3 11.9 1.8 167.9 53.9 20.7 6.41 0.25 0.31 1.45 2.06Donz"ere 58 70 900 422 7.99 69.8 6.1 11.4 1.8 166.4 49.7 18.5 6.60 0.18 0.24 1.38 2.22Aramon 16 88 600 418 8.03 69.3 5.9 11.2 1.9 163.8 48.2 18.1 6.45 0.15 0.24 1.58 2.29Arles 4 96 500 426 7.93 69.6 6.3 11.4 1.9 166.4 48.3 18.9 6.57 0.15 0.29 2.05 2.25

Ain St-Maurice-de-Gourdans 191 3650 394 8.13 74.0 2.8 2.8 1.0 227.0 4.0 6.4 4.17 0.02 0.05 1.40 2.00Doubs Sauni"eres 175 7500 432 8.12 75.2 3.3 7.5 1.9 216.2 19.2 16.4 7.47 0.03 0.14 2.86 2.18Saone Lyon 167 29 900 510 7.98 77.5 4.6 29.4 3.7 203.3 29.2 41.0 9.70 0.18 0.26 1.43 3.15Is"ere Chateauneuf-sur-lsere 128 11 800 469 7.96 79.4 9.7 11.7 1.1 152.2 100.7 18.5 3.83 0.21 0.15 1.30 1.12Durance Caumont-sur-Durance 39 14 400 502 8.14 83.2 12.8 15.1 2.0 204.7 86.4 24.1 3.25 0.02 0.03 2.32 1.14

The Doubs River is a tributary of the Saone River. Data from NADUF ‘Nationale Daueruntersuchung der schweizerischen Fliessgew#asser’ (Porte-du-Scex, Chancy), from SECOE ‘Service Cantonal de l’Ecologie de l’Eau &CIPEL ‘Commission Internationale pour la Protection des Eaux du L!eman (Gen"eve) and fromAgence de l’Eau Rhone-M!editerran!ee-Corse (other stations). Conductivity (mS/cm) at 25 !C. Concentrations of cations and anionsin mg/L. Biological Oxygen Demand (BOD5) in mg/L O2 and Dissolved Organic Carbon (DOC) in mg/L.

PART|I

Rivers

ofEu

rope

258

to reduced sediment loads in the Ain, Is"ere and Durance,partly reflecting lower flows since the middle 19th century(Table 7.5).

Sediment discharge of the Rhone has been alteredbecause of anthropogenic activities. The upper river be-tween Geneva and the Ain has seen a cessation in bedloadtransport due to gravel harvesting and retention in reser-voirs. CNR canals still contribute some sediment to the riverduring floods. Bedload transport was estimated at 1 Mm3/yearat the beginning of the 20th century but only 0.2 m3/yeartoday. Around 12 Mm3 of fine sediments are stored inG!enissiat reservoir. Presently, most suspended sedimentsnow pass through the downstream reservoirs because of anew hydraulic geometry. Excluding fine sediments stored inG!enissiat reservoir, the whole river has a sediment deficit of14 Mm3, the French Upper Rhone has a surplus of about 3–4 Mm3 and the lower Rhone has a deficit of#17 Mm3 (Dou-triaux 2006). The Ain River still provides a significant bed-load that has been extracted upstream of Lyon, but the Saoneprovides little or no bedload due to intensive gravel harvestingand the construction of a staircase of navigable reaches by lowdams.

Downstream of Lyon, a chain of diversion damsreduces velocity and prevents any erosion of the riverbed,and tributaries are usually managed to prevent the input ofbedload into reservoirs to decrease the risk of flooding.The Is"ere also has a chain of dams along its downstreamcourse. The Drome and Durance rivers still deliver bed-load, but gravel is retained upstream of the confluencewith the Rhone and extracted. Bedload entering by-passreaches from the Ard"eche River is extracted, and the riv-erbed is incising downstream from the last dam at Val-labr"egues. Mountain reforestation and land abandonmenthave further reduced erosion in the uplands, but not in thesame proportion as bedload.

7.4.5. Nutrients and Pollution

Over the past 30 years, marked changes have occurred in theproportion of the population connected to wastewater treat-

ment as well as in wastewater treatment technology. In gen-eral, organic pollution has decreased in many Europeancountries, as indicated in the improved water quality in theSaone, Doubs, Is"ere, Durance and Rhone. For instance, con-centrations of organic matter and ammonium have decreaseddramatically compared to the 1970s at Saint-Vallier, a heavilypolluted section in the Rhone. The status of smaller rivers ismore variable and can be critical when streams receive highpollution loads during low flow, especially wastewaters fromwine producing enterprises, cheese factories, and tourist areasas in Alps in winter and Mediterranean region in summer.

Concentrations of orthophosphate in the Rhone havebeen decreasing steadily over the past 20 years, followingmeasures introduced by national and European legislationto reduce eutrophication (Glennie et al. 2002). The twomain actions in the late 1980s and early 1990s were (1) areduction in the amount of sodium tripolyphosphate(STPP) to ‘alternative’ non-phosphate based detergentbuilders such as Zeolite A, and (2) improving wastewatertreatment through implementation of the Urban Wastewa-ter Treatment Directive (UWWTD). In Switzerland, ageneral decrease in phosphate resulted from the imple-mentation of phosphate removal in wastewater treatmentplants and from a phosphate-ban initiated in 1986 (Jakobet al. 2002). Although efforts were made in France toreduce polyphosphate compounds in detergents to<0.2 mg/L by 1996, a phosphate-ban decree has not beenadopted. In the lower Rhone, elimination in 1992 of alarge industrial effluent upstream of Saint-Vallier signifi-cantly lowered orthophosphate concentrations in the river(Poussard and Madrid 1999).

Pollution by nitrate is highly variable in the Rhonecatchment. Waters flowing from eastern mountains areof good to high quality while western tributaries showmoderate to bad quality. Nitrate is a non-point sourcepollution from agriculture. Among large rivers, only theSaone and its tributaries are particularly affected, espe-cially in Burgundy, and contribute >60% of the nitrateinputs to the Rhone. Nitrate values are low in the UpperRhone and increase progressively to a maximum at Jons

TABLE 7.5 Evolution of bedloads inputs of major tributaries of the Rhone River

Watershed area (km2) Natural regime Present regime

Bedload(m3/year)

Suspendedmatter (Mt/year)

Bedload(m3/year)

Suspendedmatter (Mt/year)

Ain 3713 100 000 0.15–0.3 60 000 0.1Saone 29 498 0 1.5–3 0 1.5–3Is"ere 11 865 100 000 4.5 0 3.5Drome 1642 40 000 0.2 30 000 0.2Ard"eche 2430 15 000–40 000 0.1–0.2 10 000 0.1–0.2Durance 14 322 300 000 6 0 1.8

Total 63 470 817–910 000 14.8–19.2 170–180 000 9.5–14

Etude globale pour une r!eduction des crues dues au Rhone – Hydratec, Sogreah, Minea. Etude du transport solide, synth"ese de premi"ere !etape, 2001.


due to inputs from intensive agriculture in the lower plainof the Ain River. There was a small decrease in nitrateconcentrations during the 1990s. From 1994 to 1995,nutrient inputs by the Rhone into the Gulf of Lions weremeasured at Arles (Moutin et al. 1998). The authors esti-mated the total input of nitrogen at 115–127 kt/year andmainly as nitrate (92.3–96.1 kt/year), which significantlyinfluences the nutrient balance and primary production ofthe western Mediterranean Sea.

7.4.6. Heavy Metals and OrganicMicropollutants

The most serious metals polluting the Rhone are mercuryand arsenic, and to a lesser extent nickel and zinc.

Small factories, sometimes concentrated in restrictedgeographical areas, are mainly responsible for metal pollu-tion. Metal pollution problems exist for the Arve, somevalleys in Alps, the Azergues (right bank tributary of theSaone) and the Guier River. In the Rhone, high poly-metalcontamination occurs between Lyon and the confluence ofIs"ere River along the ‘chemical corridor’ (Chasse to Saint-Vallier in the lower Rhone). Mono-metal inputs occur atPougny and downstream of the Ain River industrial plain(Santiago et al. 1994). Farm inputs clearly dominate most ofthe studied metals (animal farm breeding effluents – zinc –and soil amendments by chemical fertilizers). Copper is usedas fungicide in arboriculture and viticulture, especially thelower Rhone River, Provence, Beaujolais and Burgundy.

7.4.7. Pesticides

An environmental inventory of the Rhone (DIREN 2005)showed that pesticides are a real problem that may con-strain the ambitious aims of the Water Framework Direc-tive. Presently, the analytical assessment concerns >300active substances. In 2003, 60 surface water sites weresampled each month and 96 groundwater sites were sam-pled each season. In groundwaters, 40 pesticides wereidentified. There were 116 pesticides found in surfacewaters composed of herbicides (47%), insecticides(27%) and fungicides (22%). Over two-thirds of surfacewater sites had 10 or more pesticides detected, and highlycontaminated sites were distributed throughout the basin.For groundwater sites, contamination occurred mainly inagricultural areas such as the Burgundy foothills, theSaone River valley, the Upper Saone limestone plateau,and the Rhodanian corridor. In surface waters, glyphosate(N-phosphonomethyl glycine) and its metabolite amino-methylphosphonic acid (AMPA), amitrole (3-amino-1,2,4-triazole) and diuron were in >90% of the sites. Ingroundwaters, the most frequent products were atrazine,terbuthylazine, their metabolites and simazine (>30%sites), then diuron (#15%) and the fungicide oxadixyl

(10%). Total concentrations of active substances >5 mg/L(the drinking water limit) were observed in the Ardi"eres,Meuzin, Azergues, Seille, C"eze and Is"ere Rivers, alldownstream of vinyards and intensive agriculture areas.The Is"ere is affected by a pesticide factory on the DracRiver. Seasonally, inputs show a progressive increase dur-ing spring, a decrease in summer, and a peak again inOctober from herbicides used for winter cereals. The lastpublished survey indicated a general contamination ofrivers in the Rhone watershed by pesticides (Agence del’Eau RM&C 2004).

7.4.8. Priority Substances of the WaterFramework Directive

About 90% of 200 high-hazard substances were found inriver effluents, including mercury, nonylphenols, trichloro-methane, DEHP [di(2-ethylhexyl)phthalate]. Others danger-ous organic priority substances are found in effluents fromthe chemical industry, especially in the metropolitan areas ofLyon and Grenoble, and the heavily industrialized area alongthe lower Rhone. Urban effluents also contribute; 90% ofeffluents from wastewater plants had at least one prioritysubstance detected. Nonylphenols were found in >60% ofthe samples, and DEHP and pentachlorophenols in >30%.Polycyclic aromatic hydrocarbons (PAHs) are one of themost widespread organic pollutants in the Rhone catchment.Other micro pollutants (excluding PAHs, Polychlorinatedbiphenyls (PCBs) and pesticides) originate from two indus-trial sites on the Drac River near Grenoble and the DuranceRiver near Sisteron.

7.4.9. Artificial Radionuclides

In France, electricity is mainly produced by nuclear means:427 Twh, 78% of the electricity produced in 2004 (CEA2005). There are 15 nuclear power units and one spent fuelreprocessing plant on the river, and the Rhone valley has thehighest concentration of nuclear plants in Europe. A 40-yearsurvey of industrial radionuclide release in the Rhone (Lam-brechts & Foulquier 1987; Foulquier et al. 1991; Eyrolle etal. 2005) has been acquired by IRSN ‘Institut de Radiopro-tection et de Suret!e Nucl!eaire’ (see www.irsn.org). Radioac-tive isotopes originate from the atmospheric fallout fromweapon tests between 1945 and 1980. Radionuclide contam-ination of the atmosphere is still detectable today. The plumeof the Chernobyl meltdown on 24 April 1986 passed overeastern France in April and rainfalls in May deposited 1250–40 000 Bq/m2 of additional 137Cs to the Rhone catchment.After 20 years, the 137Cs terrestrial inputs are similar to the137Cs released byMarcoule reprocessing plant (Eyrolle et al.2005). The Marcoule plant has been reprocessing spent mil-itary and industrial fuel since 1961. A change in the treat-ment process in 1990 and the start of a dismantlement plan in1997 significantly decreased radioactive wastes from the


http://www.irsn.org/

plant. Today, 238Pu industrial inputs are still #10% higherthan those from global fallout, while 239+240Pu released byMarcoule equal the annual catchment contribution (Eyrolleet al. 2004). Recent research showed that floods causedcontamination pulses generated by the reworking of ante-cedent contaminated sediments (Eyrolle et al. 2006). Liquideffluents from nuclear installations are responsible for thelow-level presence of 60Co, 58Co, 110mAg, 54Mn, 137Cs and134Cs radionuclides. The 137Cs/134Cs activity ratio is a usefultool to describe the fate of radioactive sources reaching theRhone. Today, 137Cs terrestrial inputs are estimated at100 GBq/year, more than before the Chernobyl accidentand higher than recent industrial inputs (Eyrolle et al. 2005).

7.5. AQUATIC AND RIPARIANBIODIVERSITY

7.5.1. Algae

Information about algal composition of phytoplankton orperiphyton in the Rhone is scarce. The only available de-tailed data are those from a monitoring survey of the Saint-Alban/Saint-Maurice-l’Exil nuclear power plant 45 kmdownstream of Lyon with 395 species identified (320 phy-toplankton species, 361 periphyton species). Species of thegenus Diatoma and Chlorophyta are the most numerous(41% and 39% of the phytoplankton species, 86% and10% of the periphyton species, respectively). Cyanobacteriarepresent 9.7% of the phytoplankton species and 3.6% of theperiphytion species. Other periphyton species belong tothe Chrysophycea and Dinophycea (one species each). Themean species richness between 1985 and 2001 was #104species and ranged from 71 to 160 species. The number ofspecies has decreased continuously since 1992 (ARALEP2003).

In the by-pass section of the Pierre-B!enite water scheme,the most abundant taxon was Cladophora, and other taxawere rare (Spirogyra, Enteropmorpha intestinalis and Cya-nobacteria). In the by-pass sections of the Chautagne andBelley water schemes (data from 1994 to 1996), Cyanobac-teria (Oscillatoria limosa) were most abundant at the end ofwinter (Barbe & Barthelemy 1994; Barbe 1997). Later,substrates were colonised by Hydrurus foetidus and the fil-amentous diatoma Melosira varians, and green algae (Spi-rogyra sp. and Cladophora sp.) in summer. In the same area,rocky weirs were colonised by bryophytes typical of eutro-phic conditions: Fontinalis antipyretica, Amblystegiumriparium, Pellia epiphylla, Cinclidotus aquaticus, Cinclido-tus fontinaloides, Rhynchostegium riparioides, Brachythe-cium rivulare.

7.5.2. Macrophytes

Aquatic plant diversity in the Rhone and tributaries resultsmostly from the high number of abandoned channels. These

channels are shaped by river dynamics, and are consequentlyhighly diverse in terms of sinuosity, hydraulic capacity, anddistance from the river. This geomorphological complexitycombined with hydrology dictates (1) the frequency andduration of floods, (2) the net effect of floods (erosion versusdeposition), and (3) the discharge of groundwater exfiltratingin these channels (Bornette et al. 1998). The Upper Rhoneand several of its tributaries (e.g. Ain, Doubs, Ard"eche, Is"ere,Drome) are piedmont rivers, characterized by a coarse bed-load, and a relatively high slope. In such situations, floodduration is low (usually a few days), and floods causeincreases in flow velocity that damage plant communitiesand erode fine sediment, particularly cut-off channels withlow sinuosity and hydraulic capacity. In more sinuous chan-nels, floods have no or a silting effect, depending on thefrequency of connections between the river and the channels.Groundwater discharge is usually low in sinuous channelsthat are frequently clogged with fine sediment. Groundwaterdischarge can be quite high in others, depending on thechannel slope and substrate grain-size. This groundwatercomes either from nutrient-rich river seepage or from morenutrient-poor hillslope aquifers. Oligotrophic cut-off chan-nels are abundant along the Ain and in some places along theRhone. In most situations, the high human activity in thecatchment leads to fairly high (e.g. Upper Rhone, Is"ere) orvery high nutrient-content of the water (Saone, Doubs, lowerRhone). Highest species richness is observed in cut-off chan-nels with intermediate nutrient levels and the lowest speciesrichness occurs in nutrient-rich cut-off channels. Oligotro-phic communities have low richness but a high proportion ofrare species. Among the most abundant species that occur incut-off channels of the Rhone river and its tributaries areeutrophic species (Lemna minor, Ceratophyllum demersum,Spirodela polyrhiza, Myriophyllum spicatum) and speciesintolerant to flood scouring (Phragmites australis, Nupharlutea, Nymphea alba) (Bornette et al. 2001). Some relativelyrare species mainly occur along the Saone (Stratiotesaloides, Hydrocharis morsus-ranae, Nymphoides peltata).A few species including Callitriche platycarpa, Elodeacanadensis, Berula erecta, and Phalaris arundinacea occurin flood-disturbed cut-off channels (e.g. the Ain and Frenchupper-Rhone). Many species related to intermediate and lowtrophic levels occur along the Ain River (Potamogeton color-atus, Chara major, Luronium natans, Baldellia ranuncu-loides, Hydrocotyle vulgaris, Cladium mariscus,Schoenoplectus nigricans).

In an exhaustive study of aquatic vegetation in all cut-offchannels of the Rhone from Lake L!eman to the sea, Henryand Amoros (unpublished data, 1998) showed that speciesrichness is high (67 strictly aquatic species and 46 helophytespecies) but not uniformly distributed. Cut-off channelsalong the French Upper Rhone have a relatively low propor-tion of eutrophic species due to oligotrophic groundwaterfrom karstic origins and inputs from the Ain. From Lyon tothe confluence with the Is"ere, aquatic species that colonizecut-off channels are mainly eutrophic. Downstream from the


Is"ere confluence, the proportion of eutrophic speciesdecreases slightly, and some channels have oligotrophic spe-cies. Species richness increases significantly below the con-fluence with the Drome River with a high proportion ofoligotrophic species in cut-off channels. Further down-stream, cut-off channels of the Rhone again become highlyeutrophic. Some mesotrophic species occur exclusively incut-off channels upstream from Lyon, such as Hippuris vul-garis, Hottonia palustris, C. platycarpa, and Potamogetonnatans. Some species occur both in the upper river anddownstream of the Is"ere confluence (e.g.Groenlandia densa,Sparganium emersum) or the Drome River (e.g. P. coloratus,Sagittaria sagittifolia, Juncus articulatus). Finally, somespecies are found only in the eutrophic lower river, (Spiro-dela polyrhiza, Vallisneria spiralis, Lemna gibba). The mainnon-native aquatic plant species are Egeria densa, E. cana-densis, E. nuttallii, Lagarosiphon major, Ludwigia peploidesand L. grandiflora, Myriophyllum aquaticum.

7.5.3. Floodplain Forests

Alluvial forests have decreased by #50–80% during thepast 50 years, being replaced by agriculture and otherhuman activities. During the 18th century, forests werelimited to the riverbanks (riparian forest) and islands.Upper terraces were used for agriculture and lower ter-races as grassland. The ecological features of alluvialforests depend on geomorphology (braided, meandering,anastomosing). During the 18th century, allogenic succes-sions occurred in braided sections with a strong rejuve-nation process of fluvial landforms. Coarse sediments(sand, gravels and pebbles) were displaced by floodsand constituted unstable bars. Plant succession was usu-ally reset at a softwood stage (Salix daphnoides, Salixpurpurea, Salix eleagnos, Salix viminalis, Salix alba,Alnus incana), with the more mature species beingpoplars (Populus nigra). From the middle of the 19thcentury, anthropogenic pressure began to increase alongthe river corridor, and hard wood species such as Fraxinusexcelsior progressively colonized river margins (Marigoet al. 2000).

More recently, hydroelectric development of the Rhonevalley has led to severe changes in by-pass sections, causinga reduction in wetted areas and lowering of thewater table byup to 1 m in some areas. The main effects of the hydrologicalchangewere: (1) a regression in softwoods (S. daphnoides, S.purpurea, S. eleagnos, S. alba, S. viminalis, P. nigra and A.incana), (2) a progression of hardwoods (F. excelsior, Acerpseudoplatanus, A. platanoides, Tilia cordata, Coryllusavellana, Carpinus betulus, Quercus pubescens, Buxus sem-pervirens, F. sylvatica), (3) a progression of monopolisticspecies in open gaps (Impatiens glandulifolia, Solidagogigantea, Palaris arundinacea, Urtica dioica, Rubus fruti-cosus, Crataegus monogyna, Prunus spinosa, Berberis vul-garis, Humulus lupulus, Clematis vitalba, Parathenocissus

quinquefolia), (4) an arrival of new species usually associ-ated with Q. pubescens (sometimes Q. petraea) and C. betu-lus (Vinca minor, Mercurialis perennis, Phyteuma spicatum,Melica uniflora, Euphorbia sylvatica, Lamium galeobdo-lon), (5) the development of new types of plant communitiesas open poplar woods made up of P. nigra, Robinia pseuda-cacia (non-native species) and Q. pubescens (Klingemanet al. 1998). The creation of new areas by engineering workshas favoured the establishment of non-native species such asReynoutria sachalinense, Buddleia variabilis, Buddleiadavidii, Ambrosia artemiaefolia, Acer negudo, Amorpha fru-ticosa (in tributaries), Aster novi-belgii and Galega officina-lis (downstream from Lyon), Impatiens roylei (upstreamfrom Lyon on sandy beaches), Solidago canadensis andPhytolacca americana. Senecio inaequidens has been foundrecently in the Drac and Is"ere valleys. River embankmentand impoundment have reduced or eliminated ‘alpin’ pio-neer species as Myricaria germanica and Typha minima, aprotected species in the Swiss upper Rhone. Similarly, A.incana and somewillow species (S. daphnoides and S. trian-dra) have become rare. Along reservoirs, the increase inwater table and decrease in current velocity have led to thereplacement of species requiring well-oxygenated watersuch as A. incana by more tolerant species such as Salixcinerea and Alnus glutinosa.

Longitudinally, the following species are representa-tive: (1) limited populations of T. minima, Epilobiumfleischeri and E. rosmarinifolium are still present in theSwiss Upper Rhone, (2) Calamagrostis epigeios (sand,gravels, proximity of water table), T. minima (fine sand,proximity of water table), Salix eleagnos, Salix purpurea,M. germanica, Hippophae rhamnoides (gravels, proximityof water table, only upstream from Seyssel), Phalarisarundicea, Equisetum hiemale, Aegopodium podagraria,Salix triandra, S. daphnoides (sand, proximity of watertable), P. nigra (gravels, coarse sand, deep water table),A. incana (fine sand, mid-depth water table), F. excelsior,Ulmus montana (fine sand and silt, deep water table),Ulmus minor and Prunus avium are distributed in braidedsections upstream of Lyon, (3) in some meandering sec-tions upstream of Lyon, a decrease in agriculture in theearly 20th century allowed the development of oak forestsof Quercus robur; this forest was destroyed in the 1980s forthe Sault-Br!enaz dam, (4) between Lyon and Mont!elimarand in the Saone floodplain, Populus alba and Fraxinusangustifolia replace P. nigra and F. excelsior, typical alpinespecies are absent; in the former braided section (i.e. ‘LaPlati"ere’, nature reserve 50 km downstream of Lyon) P.alba and P. nigra co-occurred as well as F. excelsior andF. angustifolia with three maples (Acer platanoides, A.opalus, A. pseudoplatanus); in former meandering reaches(presently cut-off channels) where siltation was importantand water oxygenation lower, Salix alba, S. cinerea and A.glutinosa are dominant, and (5) in the Mediterranean partdownstream of Montelimar, large leaf trees progressivelydisappear: Tilia platyphyllos, A. glutinosa, F. angustifolia


and Q. pubescens replace Q. robur; Tamarix gallica andGlaucium flavum occur downstream of Arles. Long-termchanges in floodplain vegetation have been documented onmajor tributaries, especially the Ain (Marston et al. 1995)and Is"ere rivers (Pautou and Girel 1994; Girel et al. 2003).The main non-native plant species include Acer negundo,Ailanthus altissima, Ambrosia artemisiifolia, Amorphafruticosa, Aster lanceolatus, A. novi-belgii, A. squamatus,A. x-salignus, Bidens frondosa, Buddleja davidii,Helianthus tuberosus, H. x-laetiflorus, Heracleum mante-gazzianum, Impatiens glandulifera, Parthenocissusinserta, Reynoutria japonica, R. sachalinensis, R. x-bohe-mica, Robinia pseudo-acacia, Senecio inaequidens, Soli-dago canadensis and S. gigantea. In wetlands of theMediterranean zone, 6 non-native species occur: Acaciadealbata, Baccharis halimifolia, Cortaderia selloana, Pas-palum dilatatum, Paspalum distichum and Phyla filiformis.

7.5.4. Aquatic Invertebrates

Macrozoobenthic communities have been studied from theglacier’s outlet to the Rhone delta. Longitudinal, lateral andvertical patterns of aquatic invertebrate distribution havebeen examined as well as impacts of human activities oncommunity structure. Regardless, some compartments of theriver course have not been investigated as of this writing. Inthemost upstream section of the Swiss Rhone, the river flowsin an active floodplain (‘Gletschbode’) and is highly influ-enced by the glacier (low water temperature - rarely above4 !C - and mean conductivity values about 10 mS/cm, lowsubstrate stability and high concentrations in suspendedsediments) and receives its first tributary, the Mutt River.The Rhone upstream of the Mutt harbours a zoobenthiccommunity with low taxonomic richness (Knispel 2004).Among the 28 taxa present in the Rhone below the Muttconfluence and in the Mutt itself, Ecdyonurus picteti (Ephe-meroptera), Capnia spp. (Plecoptera), Dixidae and Thauma-lidae (Diptera) were absent in the Rhone upstream of theconfluence, and 12 taxa were more abundant in the Mutt anddownstream of the confluence the Rhone. The presence ofdifferent habitats in the floodplain (braided system after theconfluence) enhances the species diversity, allowing thepresence of Plecoptera (Perlodidae Perlodes intricatus,Nemouridae Protonemoura, Leuctridae), Trichoptera (Lim-nephilidae) and Ephemeroptera (Heptageniidae, e.g. Ecdyo-nurus picteti, Epeorus alpicola, Rhithrogena loyolaea). Inthe same upstream section, Ilg and Castella (2006) analysedthe longitudinal distribution of 5 groups of macroinverte-brates defined by 6 biological traits and demonstrated astrong upstream-downstream gradient (from 1830 m to1755 m asl). The upstream part (glacier snout) was dominat-ed by small deposit feeders or scrapers, feeding on detritus orperiphyton (Chironomidae, Diamesinae and Baetidae, e.g.Baetis alpinus). Small size, semivoltinism, absence of resis-tance forms and ability to live in the sediment intersticeswere important traits and considered adaptations to the harsh

conditions of glacial streams. These groups also occurreddownstream together with other taxa more heterogeneous insize, mainly crawlers, shredders that feed on various foodsources (Nemouridae and Limnephilidae).

A synthesis of data collected between 1991 and 1997 at142 stations in 20 rivers including the Upper Rhone andmajor tributaries highlighted the taxa distributed accordingto local habitat characteristics (substrate and vegetation),physico-chemical parameters, and hydrological regime, es-pecially hydropeaking (Baumann 2004). Hydropeaking as-sociated with the channelization of the river has led to animpoverishment of the macrozoobenthic community and adominance of Ancylidae (Mollusca), Gammaridae (Crusta-cea), Limnephilidae (Trichoptera) and Capniidae (Plecop-tera). For instance, Allogamus auricolis (Limnephilidae)seems quite resistant to hydropeaking. Some Plecopteraare almost extinct (Dol!edec 2000; Baumann 2004):Brachyp-tera trifasciata (Taeniopterygidae), Nemurella pictetii(Nemouridae), Isoperla obscura (Perlodidae) and Dinocrascephalotes (Perlidae). Perla grandis (Perlidae) is consideredrare. Siphonoperla montana, Chloroperla suzemicheli(Chloroperlidae), Perlodes intricate, Dictyogenus alpinum,D. fontium and Isoperla rivulum (Perlodidae) which werecommon alpine species in the 1980s are currently very rare.Substrate clogging by fine particles associated with algaeallowed the development of Oligochaeta (especially Naidi-dae) and Tipulidae (Diptera) associated with other pollution-resistant taxa such as Baetidae, Simuliidae, Psychodidae andChironomidae.

Bournaud et al. (1996) studied the longitudinal patternsof macroinvertebrate communities from the French-Swissborder to the sea. They identified 53 families (exceptedOligochaeta not identified to the family level) that repre-sented 85% of the entire sampled fauna.

Downstream from Lake L!eman, the longitudinal succes-sion of reservoirs, by-pass sections and short free-flowingsections associated with the original geomorphological fea-tures of the Rhone floodplain offer a large diversity of habi-tats for macroinvertebrates. Downstream from Lake L!emanto Verbois dam, three remarkable sectors were identifiedduring a 10-year survey of macrozoobenthic communities(Dethier and Castella 2002). From Lake L!eman to the Arvethe water is very clear, the river bottom is made of pebblesand cobbles, and the current velocity is about 1 m/s. In thissection, species diversity is relatively high and the dominanttaxa are Dugesia polychroa-lugubris, Dugesia tigrina, Pis-cicola geometra, Dreissena polymorpha, Hydroptila sparsa,Hydroptila teneoides, Agraylea multipunctata, Leptoceridaeand Lepidostomatidae. Other taxa, especially filtering col-lectors such as Hydropsyche spp., Neureclepsis bimaculata,were also present. Downstream of the Arve, turbidityincreases strongly, especially during spring and summer,and current velocity is reduced because the river enters theVerbois reservoir. This confluence represents a major dis-continuity in macrobenthic community with a decrease orloss of rheophilic and lithophilic taxa (Nemoura sp.,


Amphinemoura sp., Leuctra sp., Habrophlebia sp., Hydro-psyche sp., Simuliidae). These taxa appear again down-stream from Verbois dam (Perla sp., Leuctra sp.,Brachyptera sp., Heptagenia sp., Rhithrogena sp., Ecdyo-nurus sp., Rhyacophila sp., Baetis sp., Serratella ignita,Elmis sp. and Riolus sp.). Among the 61 taxa recorded inthe section between the lake and the Swiss-French border,several headwater species still occurred (i.e. Baetis alpinus,Odontocerum albicorne, Notidobia ciliaris, Hydropsychedinarica and H. instabilis).

Downstream, substantial data were collected from thelate 1970s after the first dams were built before 1950. Hy-droelectric development during the 1980s in the French up-per Rhone section (downstream from Seyssel) causedimportant changes in macroinvertebrate communities. Thecreation of artificial waterbodies such as reservoirs, head-races and tailraces, and the drastic reduction in discharge andalteration of the flow regime in by-pass sections, led to thecolonization by different communities. Rheophilic taxa suchas Hydropsychidae, Heptageniidae, Simuliidae and Elmidae(Elmis maugetii) were considered common in the Frenchupper Rhone prior to recent hydroelectric development.Most of the Trichoptera (18 taxa/24), Ephemeroptera(6 taxa/8) and Gammarus pulex were mainly or exclusivelyencountered in this section. Hydropsyche incognita wasfound exclusively at the border (Pougny), while H. contu-bernalis was dominant. Between Br!egnier-Cordon andLyon, the community had more potamophilic species suchas Heptagenia sulphurea, Polycentropus irroratus, Lypereducta and more thermophilic Trichoptera (Cheumatop-syche lepida and Hydropsyche exocellata). Within thedammed section, macrophytes favoured taxa such as Hydrasp., Chaetogaster sp., Potamopyrgus antipodarum, Physaacuta and Hydroptila sp. Planarians and other molluscs be-came very abundant. Three rheophilic and lithophilic speciesdisappeared in reservoirs, Theodoxus fluviatilis, Dendroce-lum lacteum andDugesia gonocephala (Dessaix et al. 1995).In by-pass sections, species typical of the Rhone beforeimpoundment such as Heptagenia sulphurea, Rhithrogenasp., Ecdyonurus sp., Baetis lutheri, Protonemura spp., Iso-perla spp., Perlodes intricata, Leuctra (gr. fusca) and Simu-liidae were still present but became rare (Dessaix et al. 1995;Cellot 1996). Moreover, seven species of Hydropsychidaewere found (H. angustipennis, H. contubernalis, H. exocel-lata, H. ornatula, H. modesta, H. incognita, H. siltalai).Bournaud et al. (1982) described the longitudinal distribu-tion pattern of these species: H. siltalai and H. incognitainhabited the upstream part of the French Upper Rhone; H.exocellata and H. contubernalis were common along theFrench Upper Rhone and, H. modesta was distributed allalong the river corridor. The hydraulic habitats are alsofavourable for potamic species such as Potamanthus luteusand Polycentropus flavomaculatus, and limnophilic taxasuch as Cloeon dipterum, Haliplus sp. and Platycnemis sp.A typology of side-arms, based on the distribution of macro-invertebrate species in the Rhone River upstream of Lyon

and in the Ain River was established by Castella (1987) andCastella et al. (1991). A total of 43 Mollusca species, 9Crustacea species, 29 Ephemeroptera species, 23 Odonataspecies, 112 Coleoptera species and 69 Trichoptera specieswere recorded. The lateral gradient in invertebrate commu-nities was similar to the well-known longitudinal gradients(Illies & Botosaneanu 1963; Giudicelli et al. 1980).

Downstream of Lyon, intense river regulation associatedwith strong chemical pollution and general habitat alterationled to a shift towards more lentic fauna. A total of 40 taxa wasfound both in the French upper and lower Rhone, includingOligochaeta, Chironomidae and the mollusc Ancylus fluviati-lis. Twenty seven taxa found in the French Upper Rhone werenot recorded in the lower Rhone, and 18 taxawere exclusive tothis part of the river (Berrahou 1993). Most taxa were partic-ulate detritus feeders (Mollusca, Asellidae and especiallyProasellus meridianus) and taxa resistant to pollution(Achaeta). Some taxa, such as Oligochaeta, Chironomidae,Gammarus fossarum, Asellus aquaticus (Crustacea), severalspecies of Mollusca (Bythinia tentaculata, P. antipodarum,Valvata piscinalis) were sampled all along the river but weremore abundant in the lower Rhone. Ecnomus tenellus, a lentictrichopteran, was highly abundant in the lower Rhone becauseof the reduction of hydraulic constraints after river regulation.However, macroinvertebrate communities in by-pass sectionsof the lower Rhonewere quite different from those of themainchannel, depending on the availability of habitats and hydrau-lic conditions (Fruget 1991; Fruget & Dessaix 2002). Forexample, in by-pass sections of Pierre-B!enite and P!eage-de-Roussillon, epipotamic species such as Ancylus fluviatilis,Baetis fuscatus, Heptagenia sulphurea, and Hydropsychespp. were found in some relict riffle areas (Fruget & Dessaix2002). Tributaries also may contribute to local increases ofbiological diversity (Berrahou 1993). Downstream from thelast dam (Vallabr"egues), water quality and progressivechanges of mesological conditions (salinity, granulometry,current velocity and water temperature) were responsible forthe decrease in taxonomic richness and presence of brackishand pollution-tolerant species. In a study of the invertebratefauna of the delta, Fruget et al. (1995) identified 60 taxa,mainly distributed according to their preferences in salinity(Echinogammarus sp., Theodoxus fluviatilis) and current ve-locity (e.g. E. tenellus, Hydropsyche modesta, Dugesia sp.).During the last decade, potamic and lentic species (Asellusaquaticus (Crustacea), Caenis luctuosa (Ephemeroptera), E.tenellus, Cyrnus trimaculatus, Ceraclea dissimilis (Trichop-tera)) have been favoured by the increase in water temperatureassociated with low water levels and low flow velocities inmost by-pass sections. However, rheophilic species similar tothose found upstream of Lyon such as H. exocellata andOulimnius tuberculatus (Coleoptera) are still present in theDonz"ere-Mondragon by-pass section (Pont-Saint-Esprit) lo-cated 195 km downstream of Lyon. Local hydraulic condi-tions and the nearness of tributaries such as the Ard"eche andEyrieux Rivers which enrich the Rhone with their driftingfauna can explain the presence of such taxa and other species


such as Serratella ignita and Procloeon bifidum (Ephemer-optera) (Berrahou 1993).

Long-term surveys allowed the study of the temporalvariability in zoobenthic communities. Temporal trendscombine the effects of dam building, the effects of climatechange, of invasive species spread and of more recent reha-bilitation measures. Daufresne et al. (2003) analysed thetemporal change of macroinvertebrates between 1980 and1999 around the nuclear power plant of Bugey just upstreamthe confluence with the Ain River. A gradual decrease ordisappearance of cryophilic and rheophilic taxa was ob-served (Chloroperla spp., Nemura spp., Protonemura spp.,Amphinemura spp., Brachyptera spp. (Plecoptera), Rhyaco-phila spp., Athripsodes spp. (Trichoptera), Ecdyonurus spp.,Caenis spp. (Ephemeroptera), Stratiomyidae (Diptera)) witha concomitant increase in lentic and thermophilic taxa (Cor-ixa sp. (Heteroptera), P. antipodarum, Theodoxus sp., Cor-bicula fluminea (Mollusca),Coenagrion sp., Platycnemis sp.(Odonata), Oecetis sp., Lepidostoma sp. (Trichoptera),Athricops sp. (Diptera)). The authors concluded that theincrease in water temperature under climatic warming to-gether with the development of hydroelectric schemes couldexplain the observed changes in invertebrate composition.Fruget and Bady (2006) analysed the temporal changes at thefamily level both in the French upper and lower Rhone. Theiranalysis was based on long-term surveys of the fauna near 4nuclear power plants (Bugey 50 km upstream of Lyon andSaint-Alban, Cruas-Meysse and Tricatin at a distance of50 km, 150 and 185 km downstream of Lyon, respectively).From a total of 106 macroinvertebrates families, the authorsfound a decrease in family number from upstream to down-stream that was linked to the alteration of physical habitat.They pointed out a gradual modification in community com-position between 1985 and 2004, a significant improvementin water quality, a general increase in water temperature andslight changes in hydrological regime were the most impor-tant factors explaining the observed changes. More recently,unusual summer temperatures observed since 2003 and twolarge floods in 2001 and 2002 caused an increase in thermo-philic species (e.g. E. tenellus) and exotic species (Dikero-gammarus villosus, Atyaephyra desmarestii, Hypaniainvalida, Hemimysis anomala) in the lower Rhone. The re-cent increase of minimum flows in the by-pass section ofPierre-Benite increased the proportion of rheophilic taxa.

7.5.5. Non-native Species

About30 invertebrate taxawere introduced in theRhoneRiver.Besides navigation, which allowed migrations and invasions,major changes in the river environment favoured colonizationby potamo-lentic species and expansion of others (Fruget et al.2001; Daufresne et al. 2007). The Saone River, connected bythe Freycinet canal network to rivers in northern Europe suchas the Rhine, is a primary source of invasive taxa, in particularduring floods. The main species include Cnidaria: Craspeda-custa sowerbyi (around 1900), Cordylophora caspia (down-

stream of Lyon), Bryozoa: Pectinatella magnifica, and veryrecently (2005)Urnatella gracilis (downstreamofLyon), Tur-bellaria:Dugesia tigrina (beginningof the 20th century), Poly-cheta:Hypania invalida (downstream of Lyon since 2002, theonly one freshwater Polycheta), Oligocheta: Branchiurasowerbyi (around 1950), Mollusca Gastropoda:Menetus dila-tatus, Physella heterostropha/acuta (beginning of the 20thcentury), P. antipodarum, Gyraulus parvus, Lythoglyphusnaticoides (downstream of Lyon, around 2000, thermophilousspecies) and Mollusca Bivalvia: D. polymorpha (1850), Cor-bicula fluminea (in1985 in theSaone, after 1993 in theRhone).Invasive species comprise a number of Crustacea.Gammarusroeseli, Gammarus tigrinus, Dikerogammarus villosus, Cran-gonyx pseudogracilis,Orchestia cavimana and A. desmarestiiwere founddownstreamofLyon in the 1990s.Chelicorophiumcurvispinum andHemimysis anomala appearedafter 2000alsodownstream of Lyon, Orconectes limosus was found around1960,whereasProcambarus clarkii (downstreamofLyon)andPacifastacus leniusculus (between Geneva and Lyon)appeared more recently. Dikerogammarus villosus was firstobserved in France in the Saone River in 1997, and then in1998 in the Rhone downstream of Lyon, in Lake L!eman in2002, and in the French Upper Rhone in spring 2006.

7.5.6. Protected Species

Few macroinvertebrates are protected, but Castella et al.(2005) mentioned the presence of Anisus vorticulus in sixside-arms in the French upper-Rhone (Br!egnier-Cordon).This species is listed in the annexes of the EU HabitatDirective and its conservation is clearly linked to theconservation of its habitat (lentic waterbodies with largedensities of macrophytes – Hydrocharis – and woody de-bris). Other species found in backwaters of the French UpperRhone are protected either at a national or European level:Pomatinus substriatus, Bidessus minutissimus, Stictotarsusduodecimpustulatus, Haliplus fluviatilis (Coleoptera),Caenis pseudorivulorum, Ephemera vulgate, P. luteus(Ephemeroptera), Nymphula stagnata (Lepidoptera), Ery-thromma lindeni, Erythromma viridulum, Gomphus vulga-tissimus, Libellula fulva (Odonata), Leptocerus interruptus(Trichoptera), Physa fontinalis, Theodoxus fluviatilis(Mollusca), Sialis nigripes (Megaloptera) (Paillex 2005).Lastly, four species present in the French part of the Rhonebelong to the Red List of threatened species in France(Maurin & Keith 1994): Coenagrion mercuriale, Leucorrhi-nia caudalis, Oxygastra curtisii, (Odonata), and Pisidiumpseudosphaerium (Mollusca Bivalvia).

7.5.7. The Rhone Groundwater System and itsObligate Fauna (Stygobionts)

Surface water-groundwater interactions are now recognizedto exert a major control on structural and functional attri-butes of river ecosystems. Nevertheless, the groundwater


realm flowing beneath alluvial floodplains was not obviousin the past, and its existence has only been recently incorpo-rated into the thinking of river ecologists (Gibert et al. 1994).From extensive studies carried out on the Upper Rhone River(Gibert et al. 1977; Seyed-Reihani et al. 1982; Dole 1983;Marmonier 1988), the four dimensional nature of streams,comprising their vertical extension, was documented, ex-plored and included in the ‘hydrosystem’ concept (Bravardet al. 1986b; Amoros et al. 1987b, 1988; Bravard et al. 1992).Such a precursory approach was later emphasized by Stan-ford and Ward (1988) and Ward (1989) and is now common-ly developed in many integrative studies (e.g. Malard et al.2003). The influence of exchange processes between theepigean river and its aquifer will not be detailed here, asseveral syntheses are available in the literature (e.g. Stanfordand Ward 1993; Brunke and Gonser 1997; Boulton et al.1998).

The alluvial aquifers of the Rhone host a large numberof invertebrate species, ecologically heterogeneous, espe-cially in the subsurface interactive zone (hyporheic zone)where epigean invertebrates (stygoxenes and stygophiles,Gibert et al. 1994) can seek refuge during unfavourablesurface conditions, or protect their eggs and young stagesfrom predation or other disturbances. The Rhone aquifersare also colonised by a number of obligate groundwaterspecies called stygobionts. A total of 64 stygobiotic spe-cies is currently reported from the French part of theRhone (tributaries excluded). Such a richness is high forthe groundwater environment (Culver & Sket 2000), but itis comparable to other European large rivers (Rhine,Danube) (Dole-Olivier et al. 1994). The taxonomic com-position of the stygobiotic fauna is unique compared tothat of the epigean fauna. The Rhone stygobiotic fauna ischaracterized by a quasi-absence of insects (one Coleop-tera species) and a strong dominance of crustaceans (36species, 56% of total richness) and molluscs (16 species,25% of total species richness). Three groups of crusta-ceans (Copepods, Amphipods and Ostracods) represent47% of total richness, often demonstrating higher speciesrichness in groundwater than in surface water (e.g. 10species belong to the stygobiotic genus Niphargus com-pared to 4 species in the epigean genus Gammarus). Nev-ertheless, this remarkable composition is a characteristicof stygobiotic assemblages worldwide (Ferreira et al.2003). The Hydrachnidia frequently colonise the hypor-heic zone of the Rhone, but little is known on its speciescomposition and distribution in the Rhone due to a lack oftaxonomic expertise. Despite unique features such as rar-ity, endemism, strong biological and ecological singular-ities, vulnerability, and high patrimonial value, thestygobiotic fauna still does not benefit from clear protec-tion status (Juberthie 1995; Danielopol et al. 2004).

Spatially, stygobiotic richness is non-uniformly distrib-uted along the Rhone valley corridor. Hot spots probablyresult from the co-occurrence of high amounts of Quater-nary alluvial deposits and lateral aquifers offering possible

connections between these entities. The thickness of allu-vial deposits and their lateral expansion vary along theriver continuum. Alluvial plains and constrictions succeedeach other along the river course like beads on a string(Stanford & Ward 1993). Each plain is shaped by its owngeologic settlement and its geomorphic and hydrologichistory with the active channel, thereby offering variableenvironmental conditions for the development of the sty-gofauna in terms of permeability, water circulation pat-terns within interstices and other parameters (temperature,oxygenation, food availability). From present knowledge,most of the obligate groundwater species are found in thevicinity of Lyon in the Miribel-Jonage section (Creuz!e desChatelliers 1991). This area (20 km % 5 km) is especiallyspecies rich (Culver and Sket 2000; Danielopol and Pos-pisil 2001), hosting half of the total richness reported inthe French Rhone valley (33 species). In this section, theRhone drains highly permeable glaciofluvial and fluvialdeposits (thickness &20–30 m, hydraulic conductivity&3.10$3 m/s). The water circulation pattern within thealluvia (drainage versus recharge areas) was stressed as amajor determinant of faunal composition at spatial scalesranging from a few meters to several kilometres. Hypor-heic flowpaths were described in relation with the geo-morphology of the river (lateral channel types,meandering, changes in slope, outcrops and knickpoints)(Dole & Chessel 1986; Marmonier & Dole 1986; Marmo-nier 1988; Creuz!e des Chatelliers & Reygrobellet 1990;Creuz!e des Chatelliers 1991; Dole-Olivier & Marmonier1992a; Marmonier et al. 1992). The composition of thestygobiotic fauna was shown to change in a similar wayalong the longitudinal, lateral and vertical dimensionsfrom the main channel to the floodplain margins, fromupstream to downstream reaches and from subsurface todeep phreatic zones (concept of repeated gradients, Dole-Olivier et al. 1993, 1994). Temporal changes in stygobioticfauna during flow disturbances matched those observedalong the three spatial dimensions of the river system(Marmonier & Dole 1986; Dole-Olivier & Marmonier1992b; Dole-Olivier et al. 1997). Less species-rich areaswere investigated by Creuz!e des Chatelliers (1991) in theBr!egnier-Cordon section (French Upper Rhone) andDonz"ere-Mondragon section (200 km south of Lyon), butthese investigations were restricted to the active channel ofthe river and did not extend into the floodplain. Stygobio-tic biodiversity of the Rhone is clearly under-studied.Promising and extensive explorations must be developedin poorly known areas to better assess species richness,especially in the southern part of the river corridor (e.g.Avignon-Arles section), focusing on the floodplain dimen-sion rather than the active channel submitted to intenseclogging downstream of the Is"ere. The concomitant influ-ence of past marine transgressions/regressions (i.e. sourceof marine colonizers), the absence of Quaternary glacia-tions and the presence of permeable aquifers suggest highlevels of biodiversity in these areas.


7.5.8. Fish

Geologically, the Rhone catchment emerged from a sea basinin front of the Alps orogenesis. The colonization by fresh-water organisms, especially fish, can be considered as ratherrecent. During the Pliocene, the alpine Rhone and Rhineformed the head of the Danube catchment and the fish faunaof these rivers was probably the same (Persat et al. 1995).Glaciations depleted Danubian species in the Rhone and theRhine (Persat 1988; Persat & Berrebi 1990): the most ther-mophilic and limnophilic species such as Silurus glaniswereeradicated from the Rhone and Rhine. On the other hand,glaciations allowed the colonization of the Rhone by cryo-philous species such as salmonids, bullhead (Cottus gobio),and burbot (Lota lota) (Persat & Keith 1997). The apron(Zingel asper) is the only endemic species in the Rhoneand, until recently, the only major endemic fish in France.It testifies to the ancient connection of the Rhone with theDanube that was inhabited by two species of the same genus:Z. zingel and Z. streber. The fish fauna of the Rhone alsoincludes some southern species: blageon (Telestes souffia),southern barbel (Barbus meridionalis), soiffe (Chondros-toma toxostoma) and river blenny (Salaria fluviatilis) (Persat1988). The southern part of the catchment served as refugeduring glaciation for several species. For instance, the north-ern limit in distribution of southern barbel is the Is"ere sub-basin. Colonization of the different sub-basins by speciesafter glaciation and before construction of major barriers inthe 19th century led to the current distribution of nativespecies.

The history of the Rhone corridor during the last twomillion years shows that several sections in the river can bedistinguished. The upper river, including the Swiss UpperRhone and Lake L!eman, was covered by the W#urmian gla-cier. After glacial retreat, the ‘Perte du Rhone’, a crevice likecanyon cut in the thick Urgonian limestone layer 42 kmdownstream of Geneva was a natural barrier for most fishesdownstream. Only some rheophilic species as brown trout(Salmo trutta), minnow (Phoxinus phoxinus) and grayling(Thymallus thymallus) might have colonized the UpperRhone by crossing this canyon (Persat & Keith 1997). Severaluncertainties exist about how other species colonized the riverupstream of ‘Perte du Rhone’, and the native status of manyspecies is still questionable. From ‘Perte du Rhone’ toAvignon, the Rhone was a large braided river with high flowsand sediment inputs provided by alpine tributaries. Thesehydro-morphological features allowed the persistence of arheophilic community typical of the barbel zone (Kreitmann1932; Persat et al. 1995). Presently, the Rhone extendingdown to the Saone confluence, usually called ‘French UpperRhone’, is considered as a separate entity, and the lowerRhone from the Saone to the Camargue delta is the last entity.

Excluding the Rhone, Changeux and Pont (1995) recog-nized three main ichthyogeographic regions in the Frenchcatchment: the Saone basin, the Durance basin and Mediter-ranean tributaries of the Rhone, and the Isere basin as a

typical alpine drainage. Each ichthyogeographic region hasdifferent climatic features: cool and humid in the Saonebasin (oceanic regime), dry and hot in the south (Mediterra-nean regime), and cold water in the alpine region (elevationeffect). Fish communities in lowland rivers (Saone basin)and moderate altitude rivers from the Jura mountains includ-ed bitterling (Rhodeus amarus), three-spined stickleback(Gasterosteus aculeatus), burbot, ruffe (Gymnocephalus cer-nuus), black bullhead (Ameiurus melas), and silver bream(Blicca bjoerkna). In southern rivers (southern Alps and theMediterranean) fish species include southern barbel, eel(Anguilla anguilla), soiffe, pike perch (Sander lucioperca),apron, blageon and carp (Cyprinus carpio) (Changeux1995). Alpine fish communities included brown trout, bull-head, grayling and brook lamprey (Lampetra planeri). In theinternal Alps, non-native species such as brook trout (Salve-linus fontinalis) and rainbow trout (Oncorhynchus mykiss)could constitute major fractions of the community. Fish rich-ness was positively correlated with the natural logarithm ofthe size of the sub-basins. At the intra-regions scale, the mainfactors influencing the presence or absence of different spe-cies were river slope, distance from the source, and hydro-logical features of the different sub-basins. In the Alps,altitude, which influences water temperature and flow re-gime, was an important factor explaining species presence.

Since 1860, 11 fish species have been introduced. In1860, the ruffe (Gymnocephalus cernuus) was found in theRhone. The species was native in the Meuse, Moselle andRhine catchments, and was probably native in the OgnonRiver, a tributary of the upper Saone. In 1880s, the nase(Chondrostoma nasus) colonized the French basins throughartificial waterways. It was first in the Saone (around 1840),then in the French Upper Rhone (around 1880) and in the AinRiver (in 1903). Rainbow trout (O. mykiss) were introducedin the Rhone catchment in the same decade than the nase. In1920, two other North American species were found in theRhone: the black bullhead (Ameiurus melas) and the pump-kinseed (Lepomis gibbosus). In 1930s, the pike perch (Sand-er lucioperca) colonized the Rhone and probably arrived inthe Saone and Doubs rivers from the Rhine-Rhone Canalaround 1910. The mosquitofish (Gambusia affinis) was in-troduced in south France to control mosquito populations. In1940s, largemouth bass (Micropterus salmoides) was intro-duced for angling and the species developed small popula-tions in the Rhone, mainly in the south. In 1987, the wels(Silurus glanis) was caught in the lower Rhone. It was notedin the Doubs River around 1930 and was introduced in theSaone in 1975 by anglers from where it colonized the lowerRhone and then the French Upper Rhone. The species wasalso introduced in several reservoirs (i.e. Vouglans in the AinRiver). This species was native in the Rhone River during theMiocene but was extinct during glaciations. In 1989, thePrussian carp (Carassius auratus gibelio) and topmouth gud-geon (Pseudorasbora parva) were recorded for the first timein the French lower Rhone and then in the Upper Rhone in2003, whereas goldfish (Carassius auratus) was probably


introduced earlier. Carps were introduced by Romans andcrucian carp (Carassius carassius) probably colonized theRhone during the 18th or early 19th century.

7.5.9. Fishes in the Swiss Upper Rhone

Originally, 17 native species belonging to 8 families occurredin the Swiss Upper Rhone and tributaries (Fatio 1882, 1890).These included cryophilic species (bullhead, brown trout –and Lake resident trout – , grayling), rheophilic species suchas schneider (Alburnoides bipunctatus), minnow, stone loach(Barbatula barbatula) and limnophilic species (roach – Ruti-lus rutilus – , rudd – Scardinius erythrophthalmus – , tench –Tinca tinca). The native status of several species, such asburbot, eel, perch (Perca fluviatilis) and some thermophilicones (roach, rudd, tench) is still doubtful because of the ‘Pertedu Rhone’ barrier. Of these species, only 9 are present todayand brown trout is the dominant species (K#uttel 2001; Peter &Weber 2004; Weber et al. 2007). The first (1863–1894) andsecond (1930–1960) river corrections for flood protection ledto a decrease in lateral connectivity, floodplain habitat het-erogeneity and fish diversity (Weber et al. 2007). Five specieswere introduced in the Swiss Upper Rhone (three-spinedstickleback, crucian carp, carp, rainbow trout and brooktrout), although carp is no longer present.

7.5.10. Fishes in Lake L!eman

Thirty species are present in Lake L!eman; 20 are consid-ered native and 10 were introduced (e.g. rainbow trout andbrook trout). Salvelinus alpinus and Coregonus lavaretusare native species that occurred naturally only in LakeL!eman and Bourget Lake. These species are sensitive toeutrophication and, presently, S. alpinus populations aresustained by stocking (http://www.thonon.inra.fr/poisson/pacagelacustre/pacagesalmonides/omblechevalier/omble-pacage.htm). The two initial forms of C. lavaretus presentin the lake disappeared at the beginning of the 20th cen-tury, being replaced by other forms of the same species(Gerdeaux 2001). These salmonids, perch, and to a lesserextent trout (S. trutta, lacustrine form) and pike are spe-cies of interest for anglers and professional fishermen(http://www.cipel.org/sp/). The native status of severalspecies (especially the thermophilic cyprinids) is stilldoubtful. The three-spined stickleback was introduced ina pond near Hermance (L!eman basin) in 1872 and wasprobably in Lake L!eman at the end of the 19th century(Fatio 1890). Burbot was introduced during the 15th cen-tury (Lunel 1874).

7.5.11. Fishes in the Rhone Downstream FromGeneva and French Upper Rhone

Gobin (1868) divided the French upper Rhone into five sec-tions. In the first, from the border (24 km downstream fromLake L!eman) to the ‘Parc’ (57 km from Lake L!eman, pres-

ently just downstream from G!enissiat Dam) were browntrout, chub (Leuciscus cephalus), burbot, gudgeon (Gobiogobio) and minnow. Barbel (Barbus barbus), carp, pike andperch were present only downstream of ‘Perte du Rhone’.Upstream of the ‘Perte du Rhone’ canyon, barbel was intro-duced in 1888 (Anon. 1938) and bream (Abramis brama) atthe beginning of the 20th century (after 1938). A few eelswere able to migrate above the ‘Perte du Rhone’ (Kreitmann1932). The cold turbid water of the Arve River near LakeL!eman outlet decreased the temperature in the Rhone duringsnow and ice melting periods, and species recorded in theArve (brown trout, grayling, chub and gudgeon) probablycolonized the Rhone downstream of the confluence (Kreit-mann 1932).

The second section between the ‘Parc’ and the outlet ofBourget Lake (‘Canal de Savi"ere’) had a diversity of habitatsfor fish as the Rhone flowed through a wide braided flood-plain with numerous side-arms and wetland areas in the‘Chautagne’ and ‘Marais de Lavours’, respectively. Lenticside-arms were used by phytophylic species for reproduction(pike, perch, tench and carp). The floodplain was consideredthe most productive fishery in the French upper Rhone. Thetributaries of ‘Les Usses’, ‘Dorche’ and ‘Fier’ were com-monly used by several species for breeding and as refugeagainst floods. Trout, grayling, barbel, pike, eel, perch, chub,dace (Leuciscus leuciscus) and gudgeon were the most abun-dant species (Gobin 1868; Anon. 1938). The grayling pop-ulation was very important. Among non-native species, thenase had a high population density in the braided section andnotable migrations into tributaries were reported, especiallyin the Usses and Fier (Anon. 1938).

The third section runs from the confluence with BourgetLake outlet 84 km downstream of Geneva to Sault-Br!enaz154 km from Geneva. It had large floodplains, three narrowcanyons and 13 tributaries. The Guiers River flowing fromthe Chartreuse Mountains is the main tributary with trout,grayling, burbot, barbel, chub, and dace. River blenny, anabundant species in Bourget Lake, was mainly present in theRhone downstream from the lake outlet (L!eger 1943). Theapron was also present (L!eger 1943). This third section wasthe upstream limit of twaite shad (Alosa fallax rhodanensis)and sea lamprey (Petromyzon marinus) in the Rhone. Burbotwas abundant because of the presence of small tributariesand numerous ditches in wetlands. Grayling abundance waslower than in the second section. In the lower part, fishproductivity decreased because side-arms and backwaterswere less numerous. Eel was abundant both in the Rhoneand tributaries.

The fourth section between Sault-Br!enaz and the Ainconfluence was considered relatively poor compared to theupper sections. In this stretch, the river flowed between steepbanks, islands were scarce and it had only one tributary: theBourbre River. The main fishes were pike, tench, barbel,chub, perch, carp and bream.

In the last section from the Ain confluence to Lyon, theRhone flowed through ameandering zone until Jons and, then,


http://www.thonon.inra.fr/poisson/pacagelacustre/pacagesalmonides/omblechevalier/omblepacage.htm



http://www.cipel.org/sp/

in a 15 km long braided section until Lyon. Engineeringworksconducted after 1850 to improve navigation led to the discon-nection of meanders which occur now as oxbow-lakes (Bra-vard 1987; Roux et al. 1989). The floodplain offered a largediversity of habitats and species such as trout, dace, barbel,pike, carp, tench, chub, perch, bream, roach, rudd, gudgeonand minnow. The presence of blageon was cited in the FrenchUpper Rhone by Kreitmann (1931) and L!eger (1943). Soiffewas not noted by Gobin (1868) but was probably present, atleast from the third section to Lyon (L!eger 1945). Its occur-rence was noted later in the last section (Anon. 1938). Bull-head was mentioned in the Rhone but was probably moreabundant in tributaries. Bitterling was recorded only in the5th section but should have been present upstream.

Today, the fish community of the Rhone between Genevaand Lyon is strongly affected by 10 hydroelectric schemes. Inthe Swiss Rhone, downstream from Lake L!eman, monitoringrevealed the presence of 28 species, including brown trout,grayling, pike, bream, bleak (Alburnus alburnus), barbel, gud-geon, chub, blageon, minnow, bitterling, roach, rudd, tench,stone loach, black bullhead, three-spined stickleback andperch. Some other species are present but rare: schneider,carp, pumpkinseed and bullhead. The very rare occurrenceof species such as S. alpinus, Coregonus spp. and burbot islinked to the proximity of the lake. Eel and goldfish werecaught occasionally, mostly from recent introductions (S.F.P.N.P. Gen"eve 2003). The G!enissiat damwas built from 1937 to1947 and represented the first and highest barrier to fishmigration. Daily water level fluctuations from hydroelectric-ity production are currently regulated by the Seyssel compen-sation dam built in 1951 and located 7.5 km downstream ofG!enissiat dam, and the fish community is quite poor. Five lowwaterfall hydroelectric schemes with by-pass sections be-tween Seyssel and Lyon fragment the river and create newartificial waterbodies as reservoirs and canals. Despite analtered discharge regime, by-pass sections provide suitablehabitat for most fish species, especially lithophilic fishes.

Presently, 44 species occur in the French Upper Rhone.Eels are stocked, soiffe are very rare, and natural trout andgrayling populations are found (Persat & Eppe 1997). Theintrogression of foreign genes in grayling populations showsthe impacts of stocking operations during the last 50 years.Trout and grayling are still present up to Lyon, at least in by-pass sections and connected flowing channels. The cyprinidcommunity in the French Upper Rhone is dominated bychub, barbel, schneider, gudgeon and minnow. Dace, naseand blageon are less common, probably because of habitatalteration. Lentic reservoirs and some side-arms and back-waters provide habitat for limnophilic (rudd, tench, cruciancarp, pike, pumpkinseed and, locally, pond loach – Misgur-nus fossilis) and potamophilic species (bream, bleak, pikeperch, carp, ruffe, wels, bitterling and perch) (Persat et al.1995; Klingeman et al. 1998). River blenny is regularlysampled near the Bourget Lake outlet, and burbot is rare.Coregonus sp. from Geneva Lake or Bourget Lake is sam-pled periodically. In the past 20 years, the increase in mean

annual water temperature is believed to be responsible for thedecrease in dace and increase of thermophilic species such aschub, schneider and barbel (Daufresne et al. 2003).

7.5.12. Fishes in the French Lower Rhone

Kreitmann (1932) noted that the same fish species (exceptAcipenser sturio and euryhaline fishes) were found along the248 km of the Lower Rhone, classified as a barbel zone. Theaverage slope was relatively high (50 cm/km), discharge was1500 m3/s at Viviers, and the slope reached 1.44 m/km sev-eral kilometres downstream of Valence. The substrate wasmainly coarse alluvium with silt and sand deposited alongthe banks and in backwaters. Because of the influence ofMediterranean rivers on the hydrology of the Rhone, thelower Rhone was divided into three section, a first one fromthe Saone confluence to the Eyrieux confluence (125 km), asecond one downstream from the Eyrieux confluence to theDurance confluence (123 km) and a third one close to theMediterranean Sea. The fish community in the first sectionhad at least 36 species, including large migratory fishes(twaite shad, sea lamprey, river lamprey Lampetra fluviatilisand eel), brown trout, grayling, burbot, bullhead, dace, bla-geon, soiffe, bleak, roach, barbel, bream, perch, ruffe, stoneloach, apron, river blenny and pike. The channel of thesecond section was considered as unstable with coarse sed-iment transport and high loads of suspended matter. Sidearms and backwaters were more numerous in this sectionand played an important role for fish reproduction and nurs-ery for young-of-the-year. The floodplain area was reducedby engineering works (Girardon’s embankments) in the 19thcentury for navigation, and in the 1930s a decline in commonsturgeon (A. sturio) was already noted and recorded only upto the Ard"eche confluence. The lowest part of the river isdivided in two arms: the ‘Petit Rhone’, right branch of thedelta, and the ‘Grand Rhone’, left branch of the delta(Photo 7.4). Both marine and freshwater species occurredin these arms because of the mixture of salt waters andfreshwater. Seven species were commonly fished in the ‘PetitRhone’ (carp, tench, common bream, silver bream, rudd,perch and chub), three were rare (blageon, dace, Rhonestreber) and common white fish and grayling were foundvery occasionally. Marine species were reported upstreamfrom the river mouth (brackish water): flounder (Platichthysflesus), brill (Scophthalmus rhombus), turbot (Psetta maxi-ma), common sole (Solea solea), bass (Dicentrarchus lab-rax), thick-lipped grey mullet (Chelon labrosus), goldengrey mullet (Liza aurata). Other species as sea lamprey,common sturgeon, eel, shad and flat-headed grey mullet(Mugil cephalus) were found more upstream (freshwater).In the ‘Grand Rhone’, the marine species were also recordedand some of them far upstream in freshwaters (i.e. bass).Freshwater species were found from Arles to Saint-Louis(upstream limit of brackish water): carp, common bream,silver bream, rudd, barbel (rare), perch, pike and trout (main-ly after large floods) (Gourret 1897).


Today, 43 fish species occur in the Lower Rhone, and thecommon sturgeon is extinct (Table 7.6). The French LowerRhone fish community differs from the French Upper Rhonecommunity because of the near absence of brown trout, dace,minnow, blageon, soiffe, burbot and bullhead. The graylingis absent, eel densities are higher than upstream of Lyon andtwaite shad is only found downstream ofMont!elimar. The 12hydroelectric schemes altered channel morphology, thermalregime, and substrate of the river. This river section also wasregularly impacted by pollution. From the Saone River to theIs"ere River, the fish community mainly comprises potamo-philic and limnophilic species that are well-adapted to slow-flowing regulated sections with warmer temperature andmoderate pollution (e.g. roach, chub, bleak, bream, tench,rudd, carp, pike perch, black bullhead and pumpkinseed).Barbel and nase still occur in bypass sections and riverblenny is found in low abundance. From the Is"ere River tothe Ard"eche River, the community structure changes becauseof the presence of rheophilic species such as schneider,blageon, gudgeon, barbel, nase, stone loach, bullhead, andto a lesser extent dace and soiffe. Here, water temperature islower, slope is higher, and bypass sections longer than up-stream, thereby providing suitable feeding and spawninghabitats for rheophilic species. Bitterling is present in bothsections, mainly in bypass reaches.

Downstream of the last hydroelectric power plant, theRhone flows to Arles and then forks in two arms that formits delta. Between Vallabr"egues power plant and the delta,several artificial backwaters (dikes built during the late 19thcentury) and dead arms are used as reproduction and nurseryareas (Poizat & Pont 1996; Pont & Nicolas 2001). The mainspecies occurring in this river section are chub, pumpkin-seed, silver bream, roach, nase and gudgeon. Other speciessuch as bleak, black bullhead, rudd, bream, tench and barbel

are present in lower numbers. The fish community comprises45 species, amongwhich 10 are euryhaline species (Atherinaboyeri, Gobius niger, D. labrax, C. labrosus, L. aurata, Lizaramada, Liza saliens, M. cephalus, P. flesus, Syngnathusabaster) and four are large migratory species. Physical bar-riers from river regulation have reduced longitudinal distri-butions of euryhaline species. Bass (D. labrax) and mullets(L. ramada, M. cephalus) migrated as far as the DuranceRiver confluence 85 km upstream from the sea (Kreitmann1932). Today, the bass distribution is limited to the 65 kmlower section of the river because of the Vallabr"egues power

PHOTO 7.4 Mouth of the Rhone Riv-er near Port-Saint-Louis (Photo: Com-pagnie Nationale du Rhone).

TABLE 7.6 Total number of fish species (including ex-tinct), number of remaining native, non-native species,and extinct fish species in various river sections alongthe Rhone River

River sections Total Native Non-native Extinct

Swiss Upper Rhone 22 9 5 8Lake L!eman 30 20 10 0French Upper Rhone 45 29 15 1French Lower Rhone 44 29 14 1Rhone delta 46 34 11 1Ain Upstream Vouglans 18 16 2 0Ain Vouglans Reservoir 25 16 9 0Lower Ain 32 24 7 1Upper Saone 38 28 10 0Lower Saone 38 23 12 3*

Upper Durance 5 4 1 0Serre-Poncon Reservoir 11 10 1 0Middle Durance 28 20 8 0Lower Durance 37 23 13 1

* the endangered speciesZingel asper is notofficially consideredas extinct but hasnot be observed for a long time.


plant and weir in the by-pass. The two Mullets benefitedfrom rehabilitation measures for twaite shad to pass up-stream of theVallabr"egues hydropower station and have beenfound upstream as far as Avignon. The thin-lipped greymullet (L. ramada) is most abundant.

7.5.13. The Apron

The apron, Zingel asper, is a small endemic fish in the Rhonebasin. In 1900, it was distributed throughout the Rhonecatchment but was absent upstream from ‘Perte du Rhone’.Today, it inhabits a few locations in the Ard"eche, Duranceand Doubs basins. The two most important populations arethose in the Durance and Beaume (Ard"eche) basins. A fewindividuals were still present in the Ain River in 1989 andmore recently in the Drome River. In the Rhone, a fewindividuals were recorded between 1950 and 1980, at Yenne93 km downstream of Geneva near Bourget Lake and, inMiribel Canal a few km upstream of Lyon. The last knowncapture in the Rhone was in May 1985 at Vernaison 12 kmdownstream of Lyon. The presumed reasons of the decline ofthe apron are (1) the general decrease of the natural flowregime, (2) the degradation in both quality and quantity ofwater in rivers and, (3) gravel extraction and other works inriverbeds. Recent studies improved the knowledge of thespecies regarding habitat use and reproductive behaviour(Labonne et al. 2003; Danancher 2005; Labonne & Gaudin2005), population genetics (Laroche & Durand 2004) anddiet (Cavalli et al. 2003). A Conservation Program has beeninitiated to protect the remaining individuals and restore thepopulation (http://www.cren-Rhonealpes.fr/part2/progs/life_apron.htm). Indeed, the apron is referenced in AppendixII of the Bern Convention on the Conservation of theWildlifeand Natural Environments in Europe, in Appendices II andIV of the European Fauna-Flora Habitats Directive, and inthe Red List of threatened species in France.

The program aims to (1) perform a demographic surveyof known populations, (2) develop suitable fishways for thespecies, (3) increase the connectivity between habitats (Du-rance, Ard"eche) to increase the number of individuals, thespatial distribution of the species, and the genetic mixing inthe population, (4) organize and develop amonitoring surveyof river stretches where the species is present, (5) developexperiments for artificial reproduction, (6) experimentalrestocking operations, and (7) inform people about theresults and the biology of the apron.

7.5.14. Migratory Fishes in the Rhone

Five migratory species occurred in the Rhone catchment inthe early 20th century: sea lamprey (P. marinus), Europeanriver lamprey (L. fluviatilis), sturgeon (A. sturio), twaite shad(Alosa fallax rhodanensis) and European eel (Anguilla angu-illa). The biology and ecology of the two lampreys are poorlydocumented. They occur in Camargue and in the ‘Petit

Rhone’. Their upstream distributions are limited to Avignonand the downstream part of the Durance. The sturgeon wasreported as highly abundant during the Middle Age. Over-fishing reduced the population as early as the 13th and 14thcenturies, and the decline was intensified during the 19th and20th centuries because of river embankment and regulation.The last known catch occurred in 1969–70 at the river mouthand in 1954–55 near the Ard"eche confluence, although an-other catch was reported downstream of Arles in 1989. Thisspecies is now protected and a project to restore populationsin the Rhone is under development.

Two species of shad were mentioned by L!eger (1945/48), the Rhone twaite shad (Alosa fallax rhodanensis) andthe allis shad (Alosa alosa). Today, only the Rhone twaiteshad, considered as a discrete group within the speciesAlosa fallax, is present in the lower Rhone (Le Corre et al.1997, 1998, 2000, 2005). Alosa alosa was stocked in the1950s in the Rhone and a few hybrids (A. fallax % A.alosa) were found in the Aude River, a small FrenchMediterranean river. In the Saone, before 1882, the up-stream migration of the twaite shad reached Auxonne at220 km upstream of Lyon and was reported from thedownstream end of the Doubs River. Until 1937, in theFrench Upper Rhone, shad migrated up to Bourget Lake.In the Isere River, the migration was reported as far asGrenoble. The lower parts of southern tributaries(Ard"eche, C"eze, Durance, Gardon) were also used forreproduction. The migration of the twaite shad along thelower Rhone and the main tributaries has been constrainedby 27 dams or weirs. The first barrier on the Saone wasthe ‘La Mulati"ere’ dam built in 1882 just upstream of theconfluence with the Rhone. In 1921, access to the IsereRiver was closed by the construction of ‘Beaumont-Mon-teux’ dam (#7 km from the confluence with the Rhone).The twaite shad was stopped at Lyon in 1937 after thecompletion of the ‘Jons’ dam only few kms upstream ofLyon, and the use of the reproduction sites along thelower Rhone was limited by the ‘Donz"ere-Mondragon’(1952) and ‘Vallabr"egues’ (1974) hydroelectric schemes.Until recently, the reproduction of the twaite shad waslimited to the lower Rhone downstream of the last dam.Since 1994, a large program that aimed towards restoringmigration ways for the twaite shad in the Rhone catch-ment allowed spawning migration in the Rhone in theDonz"ere bypass section and in 3 tributaries (Ard"eche,C"eze and Gardon). In 2006, young-of-the-year shad weresampled near Mont!elimar at 165 km from the sea, provid-ing evidence of reproduction in this area. Morphologicaland biological data on upstream migrant twaite shads arenow available on the Rhone (Le Corre et al. 2000). Thespawning migration occurs between March and June, andthe age of spawners ranges from 2 to 8 years for malesand 3–8 years for females. Sizes range from 255 to490 mm for males and 335–520 mm for females. Thegrowth rate and longevity of the Mediterranean twaiteshad are greater than those of Atlantic allis shad.


http://www.cren-rhnealpes.fr/part2/progs/life_apron.htm



The eel population is poorly described and few relevantdata are available to estimate the stock in the Rhone, exceptcatches by professional and amateur fishermen. The meanannual eel catch by professional fishermen is 9239 kg/yearin the delta and 7275 kg/year in the lower Rhone, and38 kg/year (delta) and 179 kg/year (lower Rhone) by am-ateur fishermen (data 1999–2002, Conseil Sup!erieur de laPeche 2005). Crivelli (1998) indicated biomasses rangingfrom 3.0 to 2269 kg/ha in canals in Camargue and from 0.2to 40 kg/ha in ponds. Data collected from 1979 to 2005 bydiverse groups monitoring fishes downstream of Lyon wereused to assess spatial and temporal changes in eel popula-tion densities and size-class structure. Over 400 electro-fishing operations were used and 4 size classes wereconsidered (Figure 7.5). The smallest eels (first and secondsize-classes) were mostly present downstream of Val-labr"egues dam, highlighting this structure as a migrationbarrier. Large eels (>440 mm, >4 years) dominated up-stream samples. The largest individuals were still presentas far as 300 km from the sea while the CPUEs (catch perunit effort) remained very low. The long-term survey of theCPUE (data from 1980 to 2005 collected from Arles to thethird dam downstream of Lyon) showed a regular decreasein eel densities as the number of dams from the sea in-creased. In each section, CPUEs decreased after 1995, andthis fact agrees with the general decline in European eelpopulations reported in the literature. Moreover, the pres-ence of dams near the delta affects strongly the upstreammigration and, the availability of suitable habitats for eeldownstream of Lyon is very low. Eel abundance in theRhone is 10–100% lower than in the Loire River, France.Further, a large number of young eels are probably re-moved and sold locally or exported to Italy and Japan.To restore eel populations, resource managers shouldimprove migration pathways, better manage flows, connectbackwaters, and improve water quality. Several studies are

examining the potential effects of Anguillicola crassus, anematod parasite affecting yellow and silver eel physiolo-gy and swimming capacity. Infection rates can reach 80%of silver eels in Camargue (Lefebvre et al. 2006).

Eels are currently restocked as well as in the FrenchUpper and Lower Rhone.

Between 1994 and 2004, 3 243 000 EU have been spentto improve the upstreammigration of fish in the lower Rhoneup to Avignon and several tributaries (Ard"eche, Gardon,C"eze).

7.5.15. Amphibians

Of the 35 amphibian species recorded in France and Switzer-land, 23 occur in the Rhone floodplain (7 species of Urodelaand 15 species of Anoura) and one in Lake L!eman (Ranalatastei). Among the Anoura, Rana kleton esculenta is ahybrid between Rana ridibunda and Rana lessonae, andRana kleton grafi was from the natural crossing betweenRana ridibunda and Rana perezi. Loss of wetlands and smallponds, water pollution, habitat fragmentation and use ofinsecticides are responsible for the decline in most amphib-ian populations, except for the marsh frog (R. ridibunda).This species disperses from central Europe and competeswith native species such as Perez’s frog (R. perezi), westernspadefoot (Pelobates cultripes) and yellow-bellied toad(Bombina variegata). The common toad (Bufo bufo), firesalamander (Salamandra salamandra) and palmate newt(Triturus helveticus) occur locally all along the Rhone. Thecommon frog (Rana temporaria), pool frog (R. lessonae) andedible frog (Rana kleton esculenta) are absent in the Rhonedelta. Perez’s frog (Rana perezi) and Graf’s frog (Ranakleton grafi) are typical Mediterranean species and occurfrom the delta marshes to Mont!elimar. Common frog popu-lations have decreased since the 1950s. The midwife toad(Alytes obstreticans), well-distributed along the Rhone Riv-er, has disappeared in the Swiss Rhone downstream of LakeL!eman since 1980, despite multiple re-introductions (Gene-va Herpetological Society).

Some amphibians are mainly found in the northern partof the Rhone. Smooth newt (Triturus vulgaris) and alpinenewt (Triturus alpestris) are rare in the French UpperRhone and Lower Rhone floodplains upstream of Mont!eli-mar but are absent downstream. South great crested newts(Triturus cristatus) are scarce because of the lack of re-production areas. Backwaters around Arles have beenidentified as the last southern reproduction areas for thisspecies. Around Geneva, South great crested newt popula-tions have been declining since 1987 because of the non-native Italian great crested newt (Triturus carniflex).Among the Anoura, the common tree frog (Hyla arborea)occurs mainly in the Upper Rhone floodplain, while theagile frog (Rana dalmatina) is found as far as Mont!elimar.Other amphibians are mainly present in the south part ofthe Rhone River. The Mediterranean tree frog (Hyla

FIGURE 7.5 Mean abundance of eel (four size classes, in mm, and for allsize classes combined, total) between the Mediterranean Sea and Lyon(average values; 1979–2005). Catch per unit effort (CPUE) corresponds tothe number of eel caught at a given station by electrofishing during 20 min.The location of the hydropower stations is indicated.


meridionalis) is commonly found in Camargue and in thelower Rhone floodplain. Until recently, the Mediterraneantree frog and common tree frog co-occurred between Lyonand Valence but river regulation and the loss of wetlandsled to their quasi-extinction. The Western spadefoot ispresent from the delta to Valence, and the Parsley frog(Pelodytes punctatus) and natterjack toad (Bufo calamita)are found in the south with abundances decreasing fromsouth to north. The natterjack toad occurs regularly alongthe river. The yellow-bellied toad has disappeared in theMediterranean part of the Rhone since the early 20th cen-tury. Most amphibian species are protected.

7.5.16. Reptiles

Three snake species and two turtles are linked to aquaticbiotopes along the Rhone River. The dark green snake (Colu-ber viridiflavus), European grass snake (Natrix natrix), vi-perine water snake (Natrix maura) are protected bothnationally and internationally (Bern Convention), and thedark green snake is listed in annex IV of the directive 97/62/EEC on the conservation of natural habitats and of wildfauna and flora. Two populations of European pond terrapin(Emys orbicularis) are known, one at ‘l0Ile Cr!emieu’ on theFrench Upper Rhone and the other in Camargue. Europeanpond terrapin populations have declined because of compe-tition with the non-native Trachemys scripta (red-eared slid-er). The European pond terrapin is listed in annexes II and IVof the directive 97/62/EEC on the conservation of naturalhabitats and of wild fauna and flora.

7.5.17. Birds

Some common bird species occur regularly along the RhoneRiver, including the grey heron (Ardea cinera), commonkingfisher (Alcedo atthis), common black-head (Larus ridi-bundus), black kite (Milvus migrans), Eurasian coot(Fulicula atra), Eurasian reed-warbler (Acrocephalus scir-paceus), mallard (Anas platyrhynchos), mute swan (Cygnusolor), and common moorhen (Gallinula chloropus). Theblack kite and Eurasian reed-warbler are common alongthe Rhone and have stable populations. Population sizes ofgrey heron, Eurasian coot, mallard and common moorhenhave increased because of their ability to colonize artificialwetlands and waterbodies, especially around towns. Thegrey heron was classified as harmful (it has almost disap-peared during the 19th century) and as a game bird in 1967,but today this species is protected. The mute swan wasconsidered very rare in 1936, but the number of individualshas increased considerably since the 1970s as the speciesbecame sedentary. In contrast, habitat alteration caused adecrease in populations of common kingfisher populationsduring the 20th century. The same is true for the little bittern(Ixobrychus minutus), which occurs in the L!eman basin anddownstream in the lower Rhone. The Eurasian curlew

(Numenius arquata) and bluethroat (Luscinia svecica) havestable populations, although these species are uncommonand nest only in the French Upper Rhone. The grasshopperwarbler (Locustella naevia) and marsh warbler (Acrocepha-lus palustris) occur in the French and Swiss Upper Rhoneand their distributions are increasing, especially the marshwarbler since 1980–1990. This species seems to tolerateurbanization and dry habitats.

The Camargue is famous for birds because it has thehighest species richness in France with 398 species (of which132 nest there) out of 512 recorded. The Carmargue also is awintering area (many duck species), an important nestingground (little bittern, squacco heron – Ardeola ralloides),and a stopping place for migratory species (i.e. red knot –Calidris canutus). Long-term trends were reported for severalspecies. The grey heron, buff-backed heron (Bulbucus ibis)and little egret (Egretta garzetta) populations have increased,while black-crowned night-heron (Nycticorax nycticorax)and purple heron (Ardea purpurea) populations have de-clined. The Camargue is the only French site where ninespecies of heron nest. It is also the main wintering area forducks in France (average of 150 000 individuals), especiallyfor common teal (Anas crecca), mallard, Eurasian wigeon(Anas penelope), gadwall (Anas strepera), northern shoveler(Anas clypeata) and common pochard (Aythya farina). Mal-lard, gadwall and red-crested pochard (Netta rufina) repro-duce in Camargue. Several species of waders have relativelystable populations such as the common ringed plover (Char-adrius hiaticula), kentish plover (Charadrius alexandrinus)and pied avocet (Recurvirostra avosetta). In France, collaredpratincole (Glareola pratincola) nests only in Camargue; thisspecies is mainly associated with rice fallow lands and tem-porary ponds and is only found in a few places in the delta.Among Laridae, some populations are increasing such asCaspina gull (Larus cachinnans) or Mediterranean gull(Larus melanocephalus), while others are decreasing suchas little gull (Larus minutus) or common tern (Sterna hir-undo). Several other species such as the common black-headhave relatively stable populations. The Camargue is the mostimportant site for greater flamingo (Phoenicopterus roseus)nesting in the west Mediterranean region and the only one inFrance. Between 1983 and 2002, the number of nesting pairsvaried from 8600 to 22 200 pairs. The number of individualshas increased during the past 50 years, especially after thecreation of a special area at ‘Salins de Giraud’.

7.5.18. Mammals

Among mammal species that occur along the Rhone, at-tention will be on species with useful and accurate data. Inthe past, European otter (Lutra lutra) and European beaver(Castor fiber) were common along the Rhone, but werehunted for their fur and almost disappeared during the 19thcentury. In 1909, the protection of European beaver insouthern France allowed the recolonization of the Rhonebasin and the species reached Lyon around 1960. The


species is now found along the whole river from Switzer-land to Camargue.

The European otter is protected at national (France andSwitzerland), European and international levels. This spe-cies is classified as ‘near threatened’ in the IUCN red list ofthreatened species. The European otter is sensitive to habitatalteration but is still present in low numbers in the Frenchpart of the river. The last observations in the Swiss UpperRhone were in 1970.

The Southwestern water vole (Arvicola sapidus) is alsolisted in the IUCN red list as ‘near threatened’. Indeed, thepopulations are critically threatened and the species is nei-ther protected at the national or European level. This specieswas very common, but wetland draining, river embankmentsand competition with the non-native coypu (Myocastor coy-pus) and muskrat (Ondatra zibethicus) were probably re-sponsible for its decline. The situation is the same forEurasian water shrew (Neomys fodiens) but it is protectedby French law and the Berne Convention.

In 1940, the North-American muskrat was absent in theRhone valley. It was recorded in 1960 in the Ain and in theFrench Upper Rhone floodplain in 1970. In Switzerland, oneobservation was reported around 1990 upstream of LakeL!eman. Today, the main goal is to limit its distribution be-cause it is considered as a vermin in France. Coypu wasintroduced in France at the end of the 19th century and itspread along the Rhone river valley during the 20th centuryand will probably reach Switzerland (Geneva) soon. Thecoypu is considered as a vermin because they cause damagesby digging holes and usually feed on agricultural plants.Among the 19 Chiroptera species identified along the Rhone,Daubenton’s bat (Myotis daubentonii) and Geoffroy’s bat(Myotis emarginatus) are protected. They occur in alluvialforests and their presence seems to be strongly related towaterbodies.

7.6. MANAGEMENT AND CONSERVATION

7.6.1. Economic Importance

Minor hydroelectrical production occurred in the Valaisfrom 1902 to 1950. An exponential increase in high-headhydropower schemes in the Swiss Upper Rhone occurredfrom 1951 to 1975. From 1976 to 2003, small hydropowerschemes, adaptation of powerhouses, and a dam heighteningwere implemented with a cumulative effective capacity of allreservoirs being 1195 Mm3, corresponding to 21% of theannual mean flow of the Rhone at Porte-du-Scex (Meileet al. 2006). Hydroelectric production in the Valais is#10 billion KWh/year of which 1.5 billion is produced inthe Swiss Upper Rhone. French production of electricity was#542.3 TWh in 2003, 78% (420 TWh) of which were pro-duced by nuclear power, 12% (64 TWh) by hydropower, and10% (54 TWh) by thermal means. The Rhone-Mediterra-nean watershed produced #25% of this electricity, and>50% of hydropower potential occurs in the Rhone catch-

ment. In the French Rhone (excluding the littoral area), themean annual production of electricity is 128 412 GWh,16 TWh (3%) of which is from hydropower. The four nuclearpower plants on the Rhone produce 22% of the French nu-clear electricity (#93 TWh/year).

The Rhone-Saone corridor is a main axis for navigation(517 km long). Since the Gallo-Roman period, the Rhonehas been used to transport various goods and materials (met-al, wood, fabrics, and cereal). This large way is connected bysmaller canals to the Rhine River (‘Canal du Rhone auRhin’), the Moselle River (‘Petite Saone’), the Seine River(‘Canal du Centre’), the Yonne River (‘Canal deBourgogne’), and the Marne River (‘Canal de la Marne ala Saone’), and the Garonne basin via the ‘Canal du Rhone aS"ete’. The transport capacity of the Rhone downstream ofLyon is 22 Mt/year, most of it being local and made of bulkgoods such as oil and gravels. Container transport is increas-ing rapidly. In 2005, 58 807 TEU (20-feet Equivalent Unit)were transported on the Rhone River, #20% greater than in2004. Rough minerals, agricultural products, oil productsand fuel represented 41%, 14%, 13% and 7%, respectively,of transported goods. One of the most important transportcentres is Port Edouard Herriot at Lyon with interconnectedrailroads, roads and navigation. The amount of transport is#10 Mt/year, 935 000 tons transported by ships.

The Rhone River catchment is an important reserve forgroundwater. Over 200 Mm3/year taken in the alluvial aqui-fer is used as drinking water for three million inhabitants.

7.6.2. Flood Control

Large floods in the Valais were relatively frequent in the past(1640, 1740, 1778, 1846, 1860), most often between Augustand October. Damages were often extreme, and led to thefirst correction of the Swiss Upper Rhone (1863–1894). Itssuccess created more industrial and economic developmentsin the Valais. After two large floods in 1930 and 1960, asecond correction was completed using large amounts of bedmaterial to build dikes. In October 2000, the Rhone flooded1000 ha (#980 m3/s at Branson), causing 306 Me in dam-age and initiating a third correction. The project will run for30 years from Gletsch to Lake L!eman and will cost 612 Me.The project aims include environmental needs, flood securi-ty, and socio-economical development such as agriculture,hydroelectricity production and tourism. The main goal is towiden the river, deepen the bed, and reinforce dikeswherevernecessary.

After World War II, economic development along thelower Rhone valley was intense: hydroelectricity production,navigation and irrigation were promoted by the ‘CompagnieNationale du Rhone’ and lead to new farms, industries andtowns along the river. Today, over 556 000 inhabitants in 310towns live in the Rhone floodplain. Three floods occurred in2002 and 2003, with damages of 846 Me from the 2003flood. In January 2004, the government introduced a ‘globalstrategy’ to prevent flooding of the Rhone and its main


tributaries. The main objectives are to limit extreme floods,to control future development in flood areas and to changeland use to reduce flood damage. To mitigate flood peaks, 12large flood expansion areas have been identified from Chau-tagne to Vallabr"egues, and the delta. Flood risk in the FrenchUpper Rhone is already low because of several large over-flowing areas. To reduce flood duration, 50% of existingdikes will be reinforced before 2015, especially betweenBeaucaire and Arles. In the delta, priority efforts will bemade to improve water drainage into the sea to better protectthe Camargue. Management policies will focus on a‘protection plan for inundation risk’ for the Rhone Riverand the main tributaries, on rehabilitation of flood expansionareas, on dike improvement, and better information on riskprevention and management.

7.6.3. Fishery

Statistics on professional and amateur fisheries on theRhone and Saone are available since 1988 (http://195.167.226.100/peche/carnet.html) (Table 7.7). Fisher-men are more numerous and fish catches higher in thelower Saone than in other sections (29% of all amateursand 38% of all professionals). For amateur fishermen, themean annual fish biomass caught is #80 kg/year per li-cence. For professional fishermen, fish biomass caught is1 ton/year per licence and each fisherman usually has sev-eral licences. The most important fishery (48%) is largecyprinids (breams, carp, roach, barbel, nase and chub).Small cyprinids (especially bleak) are an important fish-ery of professional fishermen in the Saone and Doubs.Large cyprinids represent 44% of the fishery biomass inthe French lower Rhone and 81% in the French UpperRhone. Pike-perch and pike are carnivorous fishes withcommercial interest. Pike-perch catches are important in

the Saone and French lower Rhone. Pike populations arerelatively abundant in the French Upper Rhone and upperSaone but they are less common in the lower Saone. Themean individual weight of caught fish (pikes) ranges be-tween 2.1 and 2.5 kg. Trouts are caught by amateur fish-ermen in the French Upper Rhone (average fish biomassis #1.1 kg). The Wels fishery increased significantly after1994, especially in the lower Saone. This fish is alsoimportant in the lower Doubs and to a lesser extent inthe Lower Rhone. The Wels population has increasedsince 1990. Burbot is mainly caught by amateur fishermenin the French Upper Rhone (#73 kg/year, period 1988–2001). In the delta and French lower Rhone, shads, mul-lets and bass can constitute a large part of the fishery(14% in the lower Rhone). The eel fishery also is impor-tant in the lower Rhone and delta, but is low in the Saonedespite stocking efforts.

7.6.4. Conservation and Restoration

Along the French Rhone and its main tributaries, 86 areashave protection status at the regional, national or Europe-an level. Sites of community importance (Directive 92/43/EEC) and Special Protection Areas (European Union di-rective on the Conservation of Wild Birds, 79/409/CEE)represent 22% of these protected areas (http://natura2000.environnement.gouv.fr/regions/idxreg.html). The areawithin the Natura 2000 network is 98 590 ha (1614 alongthe Ain River, 680 at the Rhone-Ain confluence, 19 658along the Saone River, 8455 along the Doubs River, 575along the Is"ere River, 6047 along the Durance River, and61 561 along the Rhone River). Nine sites in the Natura2000 network are on the French Rhone River(FR8201771, FR8212004 Lake Bourget – Chautagne –Rhone; FR8210058, Upper Rhone islands; FR8201785,

TABLE7.7Mean estimated biomasses (kg/year) caught by amateur and professional fishermen in the Rhone River (FrenchUpper Rhone (FURh), the French Lower Rhone (FLRh) and delta (DELT)), the Saone (Upper Saone (USao) and LowerSaone (LSao)) (1988–2001) and the Doubs River (1997–2001)

Fish categories FURh FLRh DELT USao LSao Doubs

Pr. F Am. F. Pr. F Am. F. Pr. F Am. F. Pr. F Am. F. Pr. F Am. F. Pr. F Am. F.

Eel 5 170 5308 170 6528 145 5 65 49 115 34 98Other amphihyalins – – 3255 566 18 509 7981 – – – – – –Large cyprinids 9368 9652 3668 8694 2385 882 2221 4875 25 232 7264 7695 1906Small cyprinids 407 133 264 1114 0 14 1047 216 13 107 905 6150 222Carnivorous fishes 623 1556 1279 1620 477 503 1055 1340 5710 4163 1640 765Salmonids 78 178 1 9 0 9 0 2 7 5 4 1Catfish 74 224 278 581 112 121 374 206 3925 2880 660 1168Others 247 826 316 992 114 232 662 2333 9300 6273 820 1531

Total 10 802 12 739 14 369 13 746 28 125 9887 5364 9037 57 330 21 605 17 003 5691

Estimated values are calculated by using the declared biomasses and estimated biomasses caught by fishermen who did not return their capture forms (oneconsiders that their fish efficiency was the same as that of fishermen declaring their captures). Eight groups of fish species have been selected. The ‘otheramphihyalins’ species group includes shads, bass, mullets, lampreys; the ‘carnivorous fishes’ species contains largemouth bass, pike, perch and pikeperch; the‘others’ group contains, among others, burbot, black bullhead, stone loach and pumpkinseed.


http://natura2000.environnement.gouv.fr/regions/idxreg.html

http://195.167.226.100/peche/carnet.html



floodplain and waterbodies of Miribel-Jonage island;FR8201749, FR8212012 Plati"ere island; FR8212010,Printegrade; FR8201677, alluvial floodplain of the lowerRhone; FR9310019, Camargue). Except for the Is"ere andAin Rivers, the main tributaries and the Rhone have areasof community importance. Four main tributaries have na-tional natural reserves, the Saone (‘La Truch"ere Rate-nelle’), the Doubs (‘l0ile du Girard’, ‘Lac du Remorey’,http://www.maisondelareserve.fr/), the Drome (‘LesRami"eres du Val de Drome, http://ramieres.val.drome.reserves-naturelles.org/), and the Ard"eche (‘R!eserve Nat-urelle Nationale des Gorges de l0Ard"eche’, http://www.gorgesdelardeche.fr/reserve-naturelle.php).

The Camargue is a Special Protection Area with 358 birdspecies being recorded and 132 nesting regularly.

There are 200 migratory species, including purple heron(Ardea purpurea), white stork (Ciconia ciconia), black stork(Ciconia nigra), and squacco heron (Ardeola ralloides). AR!egional Natural Park was created in 1970, allowing tourism,environmental protection, water resource management andthe agricultural and economical development of local popula-tions. Three main areas are usually distinguished: (1) the‘Upper Camargue’ north of the ‘Etang deVaccar"es’ is a fluvialarea with freshwater ponds, (2) the ‘Middle Camargue’around the ‘Etang de Vaccar"es’ where soft salt soils are usedfor agriculture (rice, corn, maize) and livestock grazing, and(3) the ‘Lower Camargue’ of marine – lagoon origin with azone of salt ponds used for salt production (Salin de Giraud).

Strictly protected areas in Camargue (private and public)cover 21 700 ha, the National Natural Reserve (13 117 ha,http://www.reserve-camargue.org/) is one of the most impor-tant. Its biodiversity includes 513 plant species, 50 Molluscaspecies, eight amphibian species (Mediterranean tree frog(H. meridionalis), western spadefoot (P. cultripes)), 14 rep-tile species (European pond turtle (E. orbicularis), southernsmooth snake (Coronella girondica)), and 34 mammalianspecies (Kuhl’s pipistrelle (Pipistrellus kuhli). The reserveis a Biogenetic Reserve for Europe. Two other NationalNatural Reserves exist along the Rhone, the ‘Marais deLavours’ (http://www.reserve-lavours.com/) and ‘l0Ile de laPlati"ere’ (http://www.ile.platiere.reserves-naturelles.org/).The ‘Marais de Lavours’, created in 1984 with 474 ha, isan ancient peat swamp (alkaline peat of Carex lasiocarpa)connected during floods to the Rhone before regulation(Chautagne – Belley section). The arachno and entomofaunahave been well studied: 185 spider species, 436 butterflyspecies, 120 Coleoptera species, 257 Diptera species and41 dragonfly species were recorded. Among the 215 verte-brate species, 12 amphibian species (natterjack toad (Bufocalamita), agile frog (R. dalmatina), pool frog (R. lessonae),common tree frog (H. arborea), and yellow-bellied toad (B.variegata)), 4 reptiles species (Aesculapian snake (Elaphelongissima)), and 207 bird species (Eurasian curlew (Nume-nius arquata) and bluethroat (Luscinia svecica)) were found.

‘L’Ile de la Plati"ere’ reserve is located #50 km down-stream of Lyon in the ‘P!eage-de-Roussillon’ by-pass sec-

tion and covers 484 ha. About 64 Mollusca species, 669Coleoptera species (Lucanus cervus), 43 dragonfly spe-cies (Coenagrion mercuriale), 165 butterfly species (63diurnal species, 102 nocturnal species), and 280 verte-brate species were recorded. Among the vertebrates are4 amphibians, 6 reptiles (dark green snake (C. viridifla-vus)), 215 bird species (70 nesting) and 36 mammals.Over 700 plant species were identified, including trees(83 species), aquatic plants (35 species) and herbaceousplants. These two national reserves are also Special Pro-tection Areas for birds.

A Regional Natural Reserve comprising an alder (A.glutinosa) ash (F. excelsior) forest (Alno-Padion, Alnionincanae, Salicion albae) was created in 1988 on the FrenchUpper Rhone (‘Iles du Haut-Rhone’, 226 ha).

7.6.5. Restoration Activities and Potential

The initial corrections of the Swiss upper Rhone havegreatly altered the ecological value of the floodplain,whereas the last correction, although questionable becauseof hydroelectrical production, aims to rehabilitate alluvialhabitats and biodiversity. From 1999 to 2001, a large allu-vial area (100 ha) was rehabilitated just downstream ofGeneva: ‘les Teppes de Verbois’ (http://etat.geneve.ch/dt/site/protection-nature/master-content.jsp?componentId=kmelia274&nodeId=2007) and is now protected at localand international scales (Ramsar Convention).

In 1998, a large restoration program was developed forthe French Rhone to (1) increase minimum flow rates in 8by-pass sections (Chautagne, Belley, Br!egnier-Cordon, Mir-ibel-Jonage, Pierre-B!enite, P!eage-de-Roussillon, Mont!eli-mar and Donz"ere-Mondragon), (2) rehabilitate connectionswith secondary channels and backwaters, and (3) restoremigratory pathways for largemigratory fishes (eel and twaiteshad). Presently, the minimum flow has been increased atChautagne (2004), Belley (2005), Br!egnier-Cordon (2006)and Pierre-B!enite (2000), and 23 side-arms being restored inthe French upper Rhone (Photo 7.3) and 3 at Pierre-B!enite.Cooperation between water managers, scientists and stake-holders led to the development of scientific surveys to eval-uate the success of restoration. It involves collection andmanagement of data, the development of new protocols forpre and post-restoration assessments, the development ofpredictive models of the effects of restoration, and the anal-ysis of pre and post-restoration data, and evaluating post-restoration responses (Lamouroux et al. 1999, 2007). Forfish, expected effects of the increase in minimum flow arean increase in ‘midstream’ species that prefer fast-flowingand deep microhabitats (barbel, nase, bleak, schneider, daceand grayling). In the Pierre-B!enite restored section in 2000,the proportion of these species already doubled five yearsafter minimum flow increase.

River side-channels follow ecological succession thatover time (decades) leads to terrestrialization (Bravardet al. 1986a). Under natural conditions, fluvial dynamics


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compensate for this succession to terrestrialization by cre-ating new side-channels (Amoros et al. 1987a). The RhoneRiver corrections reduced the fluvial dynamics of the riverand increased the sedimentation in backwaters that in-creased the rate of succession (Bravard et al. 1986a). TheRhone was also impacted during the 20th century by hy-droelectric projects. Consequently, the restoration of side-channels concerns both natural processes (succession andalluvial deposition) and human impacts (embankments andhydroelectric production). Regardless, rehabilitation tar-gets must be defined according to the present environmen-tal conditions and incorporate realistic strategies. Forinstance, some environmental alterations such as changesin water quality and flow regime caused by human activ-ities appear irreversible, especially in large rivers, limitingpotential restoration options. The guiding principle fordefining rehabilitation targets was to improve habitat di-versity, thereby increasing biodiversity to previous succes-sional stages (Amoros 2001). Two distinct approacheswere used in the Rhone system (1) flood pulsing as adisturbance to increase habitat diversity, reduce competi-tive exclusion, and reset ecological successions, and (2)use of groundwater to enhance habitat diversity and biodi-versity. For example, the bed of three side-channels atPierre-B!enite was excavated, sediments were removed in1999, and the minimum discharge was increased in 2000.Further, two side-channels were designed to be floodscoured, one having an additional supply of groundwaterand the other being connected to the river at both ends. Thethird side-channel was designed to have river backflowthrough a downstream connection. A 5-year monitoringprogram showed that the number of aquatic plant speciesremained 8–12 in the control side-channel, whereas it in-creased in the three rehabilitated channels, being highest(21–23 species) in the flood-scoured and groundwater-sup-plied channel (Amoros et al. 2005). The number of speciesunique to one side-channel (downstream connexion) was35% of the total species number in the bi-connected chan-nel (downstream and upstream connexion), 29% in thegroundwater-supplied channel, and 9% in the back-flowchannel.

7.6.6. EU Water Framework Directive

Seven groundwater aquifers and 26 surface waterbodieswere defined along the French Rhone (http://195.167.226.100/DCE/RM/RM_etat-des-lieux.htm). Among the sur-face waterbodies, 19 were classified as heavily modified(85% of the river) and one was unclassified (the Chautagneby-passed section). Among the remaining waterbodies, 2were between Br!egnier-Cordon and Jons (Cusset hydroelec-tric scheme), 2 were bypass sections on the upper Rhone(Belley and Br!egnier-Cordon), and the others were bypasssections on the lower Rhone (P!eage-de-Roussillon andDonz"ere-Mondragon). Two waterbodies were identified ashaving a risk of failing to meet the WFD objectives (i.e. to

reach a ‘good surface water status’ by 2015): the sectionbetween the Swiss border and Seyssel dam and the riverstretch between the Saone and Is"ere confluences. For theother waterbodies, there was doubt concerning the risk offailing to meet the WFD objectives. At the watershed scale,552 ‘running water’ waterbodies were described (61% of allwaterbodies); 161 waterbodies were identified as heavilymodified, 116 had a great risk of failing to meet the WFDobjectives, and 204 had a chance to reach the objectives. Forother waterbodies, available information was not sufficientto evaluate their ability to reach WFD status by 2015. In2003, 20 surface waterbodies reached good status for clas-sical physico-chemical parameters and all waterbodies willprobably reach this status by 2015. For micropollutants suchas pesticides and metals, only 30–50% of the surface water-bodies reached good status in 2003, and no waterbodyreached good status for organic pollutants, especially poly-cyclic aromatic hydrocarbons (DIREN 2005). The biologicalstatus is difficult to evaluate for each waterbody because ofthe paucity of information. As the Water Framework Direc-tive objectives are very ambitious, success of its implemen-tation will depend on the efficiency to solve critical issuessuch as the management of water pollution (nitrates, pesti-cides), river restoration, hydroelectricity production, andnavigation. For large rivers such as the Saone and Rhone,pristine biological references are lacking and objectivesmust be defined for each waterbody. Camargue comprises10 transitional waterbodies, including the two arms of theRhone, the littoral and 7 lagoons. Only onewaterbody wouldpotentially reach a good ecological status by 2015. TheVaccar"es and ‘Salins d’Aigues Mortes’ ponds have a highrisk of failing to meetWFD objectives by 2015. For the otherwaterbodies, there is doubt whether they will reach a goodecological status by 2015, because their hydromorphologicalfunctioning is highly disturbed.

7.7. THE AIN RIVER

7.7.1. Geomorphology

The Ain is 200 km long and drains 3713 km2 in the JuraMountains (Photo 7.5). Its main tributary is the Bienne River.The Jura Mountains are composed of eastern sedimentaryfolds (1600 m asl) and western low plateaus (500–1200 m asl) sloping to the west. This well-watered and for-ested range has karstic features due to the predominance ofJurassic limestone. It was affected by Quaternary glaciationsthat layed thick till deposits. Local erosion of till providesmost of the sediment from clay to boulders entering into theAin network. The coupling of slope and fluvial processesgenerates a steep gravel-bed river. The channel of the Ainis confined inside gorges along most of its length, and achain of four hydroelectric dams were built from 1930 to1970. Downstream of the gorges, the Ain wanders almostfreely for 40 km, except where bridges constrict the chan-nel. Erosion of fluvio-glacial deposits from the W#urm Ice


http://195.167.226.100/DCE/RM/RM_etat-des-lieux.htm


Age provides most coarse sediments to the river. Side-channels from the ancient braided belt and more recentmeanders, as well as a recently developed alluvial forest,contribute to the rich biodiversity of the river corridor.However, the river can no longer cut off meanders andcreate new palaeo-channels. Further, bed incision (2 m inthe last 30 years) has lowered the groundwater level caus-ing a loss of riverine wetlands.


The Ain catchment has an oceanic hydrologic regime,despite altitude, because it is highly influenced by rainfall(Table 7.3). The isotherm of 0 !C is at 1000–1200 m a.s.l.The annual mean discharge is 132 m3/s. Strong dailyfluctuations (>100 m3/s) usually occur downstream ofthe reservoirs due to hydroelectricity production. Evenin March, the month of maximum discharge, snowmeltonly contributes #30% of discharge. Most floods occurbetween October and March, when rainfalls and snowmeltare combined. The Ain discharge during floods can reach2400 m3/s and exceeds that the Rhone River at its conflu-ence with the Ain River.


Water of the Ain is quite hard due to the predominance oflimestone, whereas sulphate concentration is low. The sedi-ment load of the Ain has never been monitored. In the 1950s,most of the bedload in the Rhone at Lyon (#30 000 m3/year)

originated from the Ain,. The chain of hydro-dams built alongthe Ain trapped sediments and most of the suspended load.Today, most sediment transfer to the Rhone is from bank andbed erosion in the 40 km long downstream reach. The Aincatchment is dominated by extensive agricultural practices(breeding, meadows and forests) with some local viticultureand cheese production in the JuraMountains. The lower valleyis characterized by intensive farming. The main industrialpollutants are from mechanical factories, surface treatments,cheese processing, plastics and sawmills. Most industries areconcentrated along the lower part of the Ain River includingthe ‘Plainede l’Ain’ (confluencewith theRhone), the ‘PlasticsValley’ (city of Oyonnax), the city of Morez with its eyeglassindustry, and the city of Saint-Claude well-known for tradi-tional hand-crafted pipes.Water quality problems are linked totoxic pollutants including pesticides in the Ain and sometributaries and metal and organic micro-pollutants in tributar-ies such as the Ange River and Oignin River (Oyonnax).

7.7.4. Aquatic and Riparian Biodiversity

The cut-off channels of the Ain have high species richnessand many rare macrophytes. Aquatic plant communities incut-off channels are adapted to flood disturbances (mainlyruderal, small-sized species, for example C. platycarpa,Potamogeton pusillus, P. natans, S. emersum), and oligotro-phic (C. major, P. coloratus) or mesotrophic waters. The Ainis highly dynamic and surface water and groundwater nutri-ent content is intermediate and low, respectively.

A large alluvial forest develops along the lower 40 km ofthe river where the floodplain widens. The riparian forest is

PHOTO 7.5 The Lower Ain River(Photo: J.M. Olivier).


mainly composed of hard wood species such as F. excelsior,F. angustifolia, U. minor, T. cordata and Ailanthus glandu-losa. Coarse and permeable sediments favour the establish-ment of communities dominated by P. nigra, S. eleagnos,Acer platanoides, Viburnum lantana, Cornus sanguinea,Ligustrum vulgare and mesophilous species such as Q. pub-escens and Robinia pseudacacia. The alteration of the riverhydrology by anthropogenic activities (lateral embank-ments, dam construction) associated with natural processes(incision) have lowered thewater table by#1 m, and pioneercommunities have been progressively replaced by more ter-restrial-like species (Marston et al. 1995).

Data for aquatic invertebrates are mainly available for thealluvial floodplain of the lower Ain where habitat diversity ishigh (Berahou 1993; Cog!erino 1989). The most representa-tive limnophilic taxa are Helobdella stagnalis, Erpobdellaoctoculata (Hirudinea), Galba corvus (Mollusca), Leptoph-lebia marginata, Habroleptoides modesta (Ephemeroptera),Taeniopteryx schoenemundi (Plecoptera), Aeschnidae (Odo-nata), Callicorixa praeusta, Micronecta sp. (Heteroptera),Athripsodes sp., Mystacides azurea, Setodes argentipunctel-lus, and Oxyethira distinctella (Trichoptera). The most abun-dant rheophilic taxa are B. lutheri, B. rhodani, Ecdyonurusvenosus (Ephemeroptera), Chloroperlidae, Siphonoperla tor-rentium (Plecoptera), Esolus parallelepipedus, Elmis aeana,E. maugetii (Coleoptera), Rhithrogena semicolorata, Rhyaco-phila dorsalis, Hydropsyche incognita, C. lepida (Trichop-tera), and Wilhelmia equina (Diptera). Among the maintributaries of the French Rhone River (Arve, Guiers, Ain,Saone, Is"ere, Eyrieux, Drome, Ard"eche, Durance, Gard),Berahou (1993) considered the lower Ain as one of the mostlotic. Among the macroinvertebrates collected in thesetributaries, five rheophilic taxa are exclusive to the Ain (Orec-tochilus villosus, Riolus subviolaceus (Coleoptera), Agrayleasp.,Hydropsyche contubernalis and Setodes argentipunctellus(Trichoptera)). The side-arms of the lower Ain harbour 36species of Trichoptera, 94 species of Coleoptera, 10 speciesof Odonata, 12 species of Ephemeroptera, and 6 species ofCrustacea (including the stygobiotic amphipods Nipharguskochianus and Niphargopsis casparyi). The composition ofColeoptera reflected successions within side-arms (Castella1987). Elmidae were dominant in lotic side-arms but wereprogressively replaced by Dysticidae and Haliplidae in paleo-channels with strong groundwater input.

L!eger (1926) recorded 18 fish species from the Bienneconfluence (the major tributary of the Ain) to the Rhoneconfluence. The dominant species were nase, brown trout,grayling, chub and bream. Brook lamprey, eel, twaite shad,barbel, gudgeon, blageon, schneider, carp, stone loach, pike,and bullhead, perch and ruffe were present in lower densities.Minnow inhabited small tributaries and was probably alsopresent in the Ain. Apron was not recorded by L!eger (1926)but it occurred in the Ain and Bienne. The upper course of theAin offered impressive lotic sections suitable for brown trout.

The main alteration of the river by large dams occurredfrom 1901 (Sault-Mortier) to 1968 (Vouglans). The canyon

was transformed into a succession of several reservoirs withfavorable conditions for limnophilic species. Presently, theAin can be divided in three parts: the upper section from theheadwaters to the large dam of Vouglans, including the firstlarge dam at Blye (48 km from the spring); the intermediatesection fragmented by a succession of 5 large dams; and thelower Ain downstream from the last dam (Allement) to itsconfluence with the Rhone.

In the upper section, the river has been fragmented bywatermills built to produce energy for small factories andfive low dams. The impact of these dams on fish populationshas not been studied. Eighteen species occur in this section.They are mainly rheophilic species (stone loach, bullhead,schneider, barbel, soiffe, blageon, gudgeon, dace, minnow,trout and grayling) although limnophilic species such astench, rudd, and black bullhead also occur. Other native(bleak, roach, pike, perch, and chub) and non-native species(carp and pumpkinseed) are also present. Vouglans is thelargest reservoir along the river. Among the 25 species pres-ent in the reservoir, 9 are non-native and several of themwereintroduced for angling (pike-perch, brook trout, lake char –Salvelinus namaycush – and wels).

The following reservoirs downstream hold a limnophiliccommunity.

The lower Ain is a large floodplain with meanders andseveral side-arms and backwaters often fed by groundwater.Thirty-one species belonging to 12 families occur in thelower floodplain. Rheophilic species (trout, dace, nase, bar-bel, schneider, minnow, stone loach, and blageon) dominate,and the grayling population is the most important one in theRhone catchment. Apron, burbot and eel are very rare. Pota-mophilic (bream, pike, and carp) and limnophilic species(tench and rudd) maintain small populations because of thepresence of lentic side-arms and backwaters (Mallet 1999).Soiffe is absent although a large natural population is presentupstream in the Suran River, a tributary flowing from Jura.Six of the 31 species are non-native. Hydrology and watertemperature depend on hydropeaking. Minimum instreamflows increases water temperature during summer, stressingthe most sensitive species such as grayling.

Among the three species of Urodela present along theAin, the fire salamander and palmate newt are the mostcommon species whereas the alpine newt is present only inthe lower river. Eight species of Anoura occur along the river.The common toad and midwife toad are well distributedalong the river, whereas the natterjack toad occurs in a fewspots within the valley. The southern species, Pelodytespunctatus (Parsley frog), inhabits mainly the lower Ain,and could be present upstreamwhereas agile andmarsh frogsoccur only downstream of Vouglans dam. Only the Europeangrass snake inhabits the Upper Ain valley. This species co-occurs with dark green snake in the lower Ain. The Europeanpond terrapin is probably present.

The bird community of the Ain is composed of commonspecies with stable populations such as mallard or Eurasiancoot. Other populations such as grey heron or little ringed


plover (Charadrius dubius) are increasing as the number ofsand and gravel pits increase. Hen harrier populations (Cir-cus cyaneus) have been declining in the whole Rhone-Alpes} region since the 1970s, but this species is still nestingin a few places in the lower Ain floodplain. Gadwal nested inthree places in 1977 and is still nesting today in a few places.Four species of falcon inhabit side-arms and backwaters ofthe floodplain: Eurasian hobby (Falco subbuteo), rock kes-trel (Falco tinnunculus), merlin (Falco columbarius) andperegrine falcon (Falco peregrinus).

Most records of mammals are in the lower Ain includingEuropean otter, European beaver, Southwestern water voleand Eurasian water shrew populations. Muskrat and coypupopulations are probably limited to the lower Ain.

7.7.5. Management and Conservation

Mean annual hydroelectricity production along the Ain isabout 750 GWh. Tourism is well developed along the riverand Vouglans reservoir (canoe-kayak, angling, swimming).About 27 000 and 9000 persons/year practice nautic sports inthe Ain and Vouglans reservoir, respectively. The UpperRiver and lower valley are famous for trout and graylingangling, respectively. Because of the good water quality,recreation activities are very intense in the lower Ain duringsummer and recent attempts aim to regulate tourism activi-ties. Groundwater resource management is an important is-sue in the lower Ain floodplain because agriculture mainlydepends on groundwater supply. The ‘Lower Ain floodplainmanagement program’ aims to stabilize the amount ofpumped groundwater in the future.

The lower Ain is considered as a highly important flood-plain. The water quality is very good and this running part ofthe river offers suitable habitats for sensitive species such asgrayling. Further, although the discharge regime is influ-enced by hydroelectricity production, the lower valleyremains dynamic. Unfortunately, recent bed incision reducedthe meandering potential of the river and caused drying ofseveral backwaters. River bed armouring is progressing at#500 m/year. In 2002, a Life program (Conservation ofhabitats created by the Ain River dynamic, http://www.bas-sevalleedelain.com/life/fr/index.php) was approved by theEuropean Community and two contiguous sites (2294 hec-tares) were included in the Natura 2000 network(FR8201653 and FR8201645). Five backwaters were reha-bilitated (2.6 km) to stop the drying process and increasewetted areas, resulting in 20 new macrophyte species suchas L. natans. In order to favour the development of alluvialforests and especially pioneer species, 1500 ha will be pro-tected against human activities. A special management pro-gram including 40 communes and covering 602 km2, 53 kmof the lower Ain and 18 tributaries, was set up in 2006. Themain objectives of the program are: to restore the fluvialdynamics of the Ain; to improve discharge management; toimprove flood risk management; to protect groundwaterresources; to increase water quality; to preserve river and

backwater biodiversity; to restore the fish community; tomanage tourism; and to evaluate the efficiency of the pro-gram.

Among the 10 waterbodies identified along the Ain, only3 are not classified as heavily modified: the most upstreamsection from the first large dam (Blye), the river part betweenBlye dam and Vouglans reservoir, and the lower part down-stream of the last power station. Four waterbodies are reser-voirs. The first upstream waterbody is the only one having agreat probability to meet the WFD objectives (good status)and a reference site was selected in this section (Champag-nole, Jura). For all other waterbodies, the probability ofmeeting the WFD objectives is low.

7.8. THE SAONE RIVER


The Saone River drains an Eocene-Oligocene, north–southgraben filled in the early Tertiary. Between 1.8 and 3 Ma BP,the Bresse Marl Complex was made of fluvial gravels fromthe Aar and Doubs rivers and marl deposits in lacustrineareas. Between 1.2 and 1.7 Ma BP, overlaying sedimentscame from the surrounding plateaus. During the Quaternary,flowing southward and shifting to thewest, the Saone incisedits valley through uplifting sedimentary fill. The tectonicwarping was complex since the middle of the river flowedacross subsiding areas, while the lower river evolved to theuplift of the Lyon area and incised a narrow valley throughmetamorphic rocks of the eastern Massif Central. For in-stance, 40 km from its confluence with the Rhone, the SaoneWurmian gravel deposits are 10 m lower than those of theRhone in Lyon (relative subsidence: 0.2–0.4 mm/year). Theactive tectonic explains the low gradient of the Saone, whichis #0.068 m/km between the Doubs and Azergues rivers(132 km). The steepest gradient downstream of this reach(0.17 m/km) is probably due to bedload inputs from theAzergues River (about 30 km upstream from Lyon) duringthe late Holocene. The floodplain of the Saone, prone tolong-lasting floods, was protected by low levees constructedbetween 1843 and 1895. About 17 000 ha of agriculturallands are now protected except from 1 to 6 year summerfloods which sustain floodplain meadows.


The Saone has an annual mean discharge of 445 m3/s at Lyon(period: 1920–2001) (Table 7.3). The specific discharge is14.8 L/km2/s, and the river has a typical rain regime (oceanictype) with high winter flows and floods (rainfall) and lowsummer flows (evapotranspiration). The spring flow fromthe Doubs tributary is increased by snowmelt from theVosges and Jura Mountains. The 100-year flood (3180 m3/s at Lyon) is modest but its duration can be >30 days in themiddle river. Discharge of historic floods was much higher,reaching 4300 m3/s for the 1840s flood.


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Except for restricted northern headwaters on granitic sand-stone of the Vosges mountains, the river drains natural min-eralized waters from limestone bedrock and received waterinputs from the Doubs River that flows over large karsticareas. Chemical industrial inputs from Solvay (Tavaux, Jura)increase mineralization (chlorides) and a strong agriculturalpressure contributes to high levels of nitrates (Figure 7.6).

The Saone is a large slow-flowing floodplain river thatcontrasts with the alpine character of the Rhone. The riverflow is slowed even more because of navigation control.Most tributaries have a lentic character as well and waterin the catchment is highly eutrophic. The catchment mainlycomprises agricultural landscapes with strong pressuresfrom farming, and intensive culture and viticulture activitiescausing diffuse pollution by nutrients and pesticides. Severalcities located along tributaries such as Dijon (Ouche River),Besancon (Doubs River) and Belfort and Montb!eliard(Doubs River basin) are sources of urban and industrialpollutants. The Doubs catchment contains traditional milkand cheese production on the plateaus, and also automanufacturing and surface treatment facilities that releasevarious micropollutants. Tributaries from the Burgundy andBeaujolais wine producing areas add organic pollutants andpesticides, as well as urban and industrial pollutants.Tributaries in the eastern catchment have cumulativeimpacts from nutrients and pesticides linked to viticultureand intensive agriculture (corn). The upper Saone has highloads of nickel and arsenic.

The sediment load in the Saone is impacted by gravel andsand removal as well as by flow reduction from dams thatalso trap sediments from tributaries. As such, the Saoneessentially carries suspended sediments that are depositedon the alluvial plain during floods. Astrade (2005) reported asuspended load of 50 mg/L over 10 days during the January1995 flood (1585 m3/s). A concentration peak (70 mg/L) insuspended sediment occurred at bankfull discharge. In com-parison, the Rhone at Arles can transport >2500 mg/L dur-ing floods in a similar time period.


Phytoplankton are sparse in the Saone with only 24 speciesbeing recorded (e.g. 66 species are found in the Loire River).The most common taxa were Diatomophyceae, Cyanophy-ceae and Chlorophytae. In 1998, the average cell abundancewas #13 200 cells/L and chlorophyll-a concentration was4 mg/L. In summer, algae concentration can reach#567 000 cells/L and chlorophyll-a concentration 69.5 mg/L. These are relatively low values compared to the LoireRiver where average algae concentration is 40 % 106 cells/L and chlorophyll-a concentration ranges between 100 and200 mg/L. Zooplankton numbers are also low with a defi-ciency in Rotifera (14 species). Planktonic Crustacea aredominated by Copepoda (80% of the community), and Clao-docera are mainly represented by Daphnia, Diaphanosomasand Bosmina (Fruget and Persat 2000).

In the main channel, 67 macrophyte taxa were recorded(22 submerged species, 28 species with floating leaves, 16emergent species, and 15 filamentous algal species). Born-ette et al. (2001) found 66 species of which 12% were absentin the Doubs, Rhone and Ain rivers. Eutrophic species suchas C. demersum, S. polyrhiza and L. minor are highly abun-dant as well as species intolerant to floods (N. lutea and N.alba). Other species such as S. aloides, Valisneria spiralis,Potamogeton nodosus andM. spicatum are also common. E.Canadensis, Potamogeton lucens, P. perfoliatus and S. sagit-tifolia occur in a few backwaters. The most common helo-phytes are tolerant to high nutrient levels and preferundisturbed wetlands (Glyceria maxima, Rumex hydrola-pathum, Acorus calamus and P. australis). Rare species suchas Trapa natans, Hydrocharis morsus ranae, and Butomusumbellatus occur in the river. Cut-off channels along theriver are mostly eutrophic with little influence by floods.They are consequently dominated by the same tall, compet-itive eutrophic species (Godreau et al. 1999).

Because of its hydrology and fertile soils, the floodplainhas been used for agriculture since the Middle Age. Present-ly, there are many poplar plantations. Two natural sectionsinclude: (1) fragments of alluvial forests upstream fromChalon-sur-Saone where F. angustifolia reaches its northernlimit and occurs with A. glutinosa and Salix alba; and (2)small and fragmented hard wood forests downstream fromChalon-sur-Saone, composed of Q. robur, F. angustifolia, F.excelsior, Ulmus laevis, U. minor, A. glutinosa, and Salixalba. These residual forests are endangered by developmentof poplar plantations and agriculture. U. laevis is mainlypresent along the Saone in the Rhone catchment.

The Saone is characterized by typical limnophilic macro-invertebrate species. From the spring to the confluence withthe Doubs River, communities are dominated by burrowingand pollution intolerant species such as Pisidium supinum(Mollusca), Ephemera lineata, Ephoron virgo, C. luctuosa(Ephemeroptera), whereas epibenthic and pollution tolerantspecies are abundant downstream (Helobdella stagnalis,Her-pobdella octoculata, Glossiphonia complanata (Hirudinea),

FIGURE 7.6 Long-term change of the nitrate concentration (mg/L) in theSaone at Auxonne (Cote d0Or).Source: Data from Agence de l0Eau Rhone-M!editerran!ee-Corse.


Asellus aquaticus (Crustacea)) (Tachet et al. 1988). A recentstudy of the lower Saone upstream from Lyon showed thathalf of recorded taxa were alien species: Chelicorophiumcurvispinum,Dikerogammarus villosus, A. desmarestii (Crus-tacea), D. polymorpha and Corbicula fluminea (Mollusca),Pectinatella magnifica (Bryozoa), Hypania invalida (Poly-chaeta), and Dugesia tigrina (Planaria) (Persat and Fruget2007). Except for Oligochaeta and Chironomidae, commonnative species are potamolentic species such as E. tenellus(Trichoptera) and C. luctuosa (Ephemeroptera). The abun-dance of non-native species probably explains the rarity ofburrowing species such as E. lineata and E. virgo (Ephemer-optera) orG. fossarum and Asellus aquaticus (Crustacea), andthe recent decline of Gastropoda. Over 40 Odonata specieswere recorded in cut-off channels of the Saone River (God-reau et al. 1999), some quite common (Ischnura elegans,Platycnemis pennipes and Coenagrion puella) and othersspecific to particular habitats (C. mercuriale and Gomphuspulchellus). Both C. mercuriale and O. curtisii have threat-ened status in Europe and Cordulegaster bidentata is restrict-ed to France. Mollusca received a particular attention becauseof their high abundance in the Saone. Annual surveys (1996–2004) at five sites along the Saone (116, 194, 225, 332 and472 km downstream from the source) and two sites on twotributaries (Ognon and Doubs) identified 24 species (14 Gas-tropoda, 10 Bivalvia). Nearly 60% of the total density wasrepresented by three species: Valvata piscinalis, Pisidiumsubtruncatum and Corbicula fluminea. Other important spe-cies were the gastropods P. antipodarum, Gyraulus albus andBithynia tentaculata, and the bivalve SphaeriidaeMusculiumlacustre, Pisidium nitidum, P. casertanum, P. amnicum and P.moitessierianum.

The longitudinal gradient in aquatic insects along theDoubs and 11 tributaries was described by Verneaux et al.(2003). Species in upstream springs, brooks and karstic out-lets includedRhadicoleptus spinifer,Drusus annulatus, Aga-petus fuscipes (Trichoptera), N. pictetii, Nemoura cinerea,and N. cambrica (Plecoptera). In the middle and lower sec-tions (Doubs, Loue) were E. lineata, E. virgo, P. luteus, andCaenis horaria (Ephemeroptera), and E. tenellus, A. multi-punctata, and Leptocerus tineiformis (Trichoptera). Fruget etal. (1996) examined macroinvertebrate communities in themain channel and backwaters of the Doubs River and Rhine-Rhone Freycinet canal that connects the Rhine to the Saone.High richness (149 taxa) was related to the habitat diversity.Dominant taxa were Chironomidae and Oligochaeta. Othercommon taxa included Trichoptera (27 species), Mollusca(22 species), Coleoptera (18 species) and Ephemeroptera (17species). Plecoptera (Leuctra sp.), Trichoptera (Hydropsychesiltalai, H. incognita, H. exocellata, H. contubernalis, C.lepida, Psychomia pusilla, and Rhyacophila spp.), Ephemer-optera (B. fuscatus, S. ignita, and Potamanthus lutheus),Coleoptera (E. parallelepipedus and Elmis maugetti) andDiptera (Simulium spp.) were common in lotic stretches.Lentic species (L. tineiformis, C. dipterum, Caenis sp., Cae-nis robusta, the Odonata Coenagrionidae,Ancylus fluviatilis,

Physa acuta) were found in backwaters and canals. Leutri-dae, Rhyacophila spp., and Baetis spp. were associated withthe bryophyte Fontinalis upstream of the confluencewith theAllan River.

Originally, at the beginning of the 20th century, the fishfauna was composed of 30 native species, including twaiteshad and sea lamprey (mainly in May during migration) thatare no longer able to migrate. River lamprey was notrecorded by L!eger (1945), but it probably migrated in theSaone in the 19th century. Apron was present in the Loue andDoubs rivers. Nine-spined stickleback (Pungitus pungitus)and probably ruffe were native in the Saone basin. The upper13 km of the river was the trout zone, the following 93 kmthe barbel zone, and the last 366 km the bream zone (L!eger1945). In the last 70 km before Lyon, L!eger (1945) found 14abundant species (bleak, roach, bream, nase, chub, carp,tench, gudgeon, perch, pike, eel, burbot, black bullheadand pumpkinseed), 4 common species (dace, soiffe, barbeland rudd) and 10 less common species (brown trout, ruffe,pike-perch, large-mouth bass, minnow, bullhead, three-spined stickleback, stone loach, bitterling and brook lam-prey).

The Saone is usually divided into the upper Saone fromthe source to the Doubs confluence (315 km) and the down-stream lower Saone (157 km). Thirty-eight species occur inthe upper Saone and 10 are non-native. Apron is present inthe Lanterne and Ognon rivers. Cyprinids are well repre-sented in the Saone. Rheophilic fishes such as bullheadand trout are rare and grayling is absent. In the Doubs, 35fish species are known, and nine-spined stickleback is ab-sent. The brown trout is widely distributed in the Doubs, andgrayling occur in the upper Doubs near the French-Swissborder and in the tributaries Dessoubre and upper Loue. Thelower Doubs has a typical potamic ichthyofauna such asroach, bleak, bream and silver bream, chub, wels, perch,but also rheophilic species such as dace, schneider, soiffe,nase, barbel, minnow and gudgeon (Fruget et al. 1998).Regularly, 38 species occur in the lower Saone and 12 arenon-native. Brown trout, bullhead, and burbot are rare orabsent. Dominant fishes are potamic and limnophilic spe-cies, including roach, chub, bleak, pumpkinseed, silverbream, rudd, gudgeon, perch, tench, topmouth gudgeon, bit-terling, bream, carp and pike-perch. Large carp occur in theSaone. Pike are not abundant in the lower Saone because ofthe disappearance of submerged meadows used for repro-duction. Wels is very abundant. Cut-off channels and side-arms provide suitable reproductive and nursery areas forphytophylic and litho-phytophilic fishes (Grenouillet et al.2000, 2001).

Five species of Urodela and eight species of Anouraoccur in the Saone catchment. The common toad, midwifetoad, common frog, fire salamander, palmate newt and alpinenewt are common. The smooth newt is less abundant than thealpine newt and the great crested newt is relatively rarebecause of fish predation. The Saone floodplain providessuitable habitat for the common tree frog. The common toad


and agile frog, which are abundant in southern France, arerare along the Saone. The yellow-belled toad is only found inthe upper Saone valley; its absence downstream is probablydue to the loss of flooded forests in the floodplain (Craney &Piston 1994). The dark green snake and European grasssnake are present along the river.

Several common birds occur in the Saone valley, includinggrey heron, Eurasian reed-warbler, meadow pipit (Anthuspratensis), reed bunting (Emberiza schoeniclus), yellow wag-tail (Motacilla flava) and northern lapwing (Vanellus vanel-lus). Other species such as the grasshopper warbler have stablepopulations. Eurasian curlew populations have increased overthe last 10 years with 200–300 breeding pairs today. Corn-crake (Crex crex) occurred everywhere in France except thesouth in the early 20th century, but the number decreased by>35%between 1984 and 1997 (Mayaud 1936). Presently, thisspecies is found only in the Saonevalley (30–50 singingmaleswere recorded in 2000). The situation was the same for thesedge warbler (Acrocephalus schoenobaenus). Whinchat(Saxicola rubetra) is rare in France but present in the Saonefloodplain and is increasing in numbers. The marsh warblershows the same pattern with an increase since 1990. In con-trast, the little bittern has almost disappeared in the Saonefloodplain. Several species of Ardeida are found on the |Ilede laMotte}, a Natura 2000 site in the lower Saone, includingblack-crowned night heron, little egret and buff-blacked her-on. A colony of 30–50 pairs of black-crowned night-heronestablished in this Natura 2000 site is of special importancebecause the species is decreasing in the Rhone-Alps region.The Saone floodplain provides suitable habitats (feeding andresting areas) for overwintering migratory birds such as thegreat white egret (Egretta alba), white stork (Ciconia cico-

nia), osprey (Pandion haliaetus), and several waders such asthe wood sandpiper (Tringa glareola).

European beaver have been present in the Saone flood-plain since 1991, although few data exist. European otter arefound in the lower Saone, and Eurasian water shrew, muskratand coypu have been recorded all along the Saone.


Hydroelectricity is mainly produced on the Doubs and Louerivers (#300 GWh/year). The Saone valley is an old axis ofcivilisation, being a natural link between the Mediterraneanand lands in the north (Bravard et al. 2002a) (Photo 7.6).The Saone has been regulated for navigation since the 19thcentury, using low navigation dams upstream after 1841 andnarrowing the channel downstream after 1845 (Astrade2005). Although the navigation system was completed by1882, transport was modest (400 000 tons/year) due to rail-way competition. Around 400 km of the Saone are navigablefromCorre to Lyon. Only small ships (<400 tons) are able tonavigate the ‘Petite Saone’ where 22 dams were built.Between 1958 and 1991, five navigation dams were builton the ‘Grande Saone’ and the riverbed was dredged to allowlarge barges up to 4400 tons. Four large commercialports are present along the Saone: Pagny-Seurre (mean an-nual traffic: 244 000 tons), Chalon-sur-Saone (mean annualtraffic: 903 000 tons), Macon (mean annual traffic:418 000 tons) and Villefranche-sur-Saone (mean annualtraffic: 696 000 tons).

The alluvial aquifer of the Saone is a large reserve fordrinking water with 700 000 inhabitants currently using thisresource.

PHOTO 7.6 Saone River, 25 km up-stream from Lyon (Photo: J.M. Olivier).


Fishery is an important activity on the Saone, mainlycyprinids, wels and pike-perch. Spinycheek crayfish (Orco-nestes limosus) is also harvested (367 kg/year in the upperand 4217 kg/year in the lower Saone).

The Saone floodplain has been colonized by typicalplants and animals with large areas covered by meadowsor alluvial forests. From the source to the confluence withthe Rhone River, four sites belong to the Natura 2000 net-work (FR4301342, Saone Valley; FR 2612006, alluvial mea-dows and associated environments of Saone-et-Loire; FR2600976, floodable meadows and forests of the Saone valleybetween Chalon/Saone and Tournus and the lower GrosneRiver; FR 8201632, wetted meadows and alluvial forest ofthe Saone valley). These alluvial habitats are especially im-portant for birds, for instance, as a nesting place for corncrake and a biotope for Eurasian curlew. The lower Doubsconstitutes another Natura 2000 site (FR 4301323), provid-ing several habitats for aquatic and terrestrial vegetation andanimals (little ringed plover, stone curlew (Burhinus oedic-nemus), European bee-eater, purple heron, and little bittern).

The Saone valley was heavily impacted in the past (toxicand agricultural pollution, sand and gravel extraction, naviga-tion). Rehabilitation success of the Saone River is dependentupon the improvement of water quality and ecological statusof polluted tributaries. A large management program wasdeveloped for the catchment. A Life program (1997–2001,1.32 Me) was launched in a floodplain area (72 000 ha) withthe goal to harmonize agricultural practices, flood manage-ment and biodiversity conservation. Specific objectives wereto: (1) increase compatibility between navigation and agricul-ture by improvement of navigation equipment; (2) restorefloodplain functioning; (3) to protect groundwater resourcesby limiting N and P inputs; and (4) to set up a monitoringprogram to inform and educate local people. An experimentalprogram (1998–2001) was developed on 6500 ha (six sites) inthe floodplain, and included restoration and maintenance offloodgates, ditches and canals, access of spawning sites forphytophylic fish species (especially pike), management of thealluvial forest for native species, limiting bank erosion byusing eco-engineering technology, and protection of ground-water quality and quantity. Shore protection against wavescreated by ships favoured vegetation development (V. spiralis,Najas spp.), microphytic and microbenthic species, andprovided nursery places for fishes.

Among nine waterbodies along the Saone, only the last40 km between Villefranche/Saone and the Rhone conflu-ence were classified as heavily modified. Indeed, water pol-lution in the lower river is high (mainly pesticides andmetals, but also nutrients), and backwaters often were al-tered. The status between the Doubs confluence and Ville-franche/Saone (126 km) is still doubtful. Two waterbodieshave a high risk of failing to meet the WFD objectives. Therisk is low for only one waterbody and there is doubt con-cerning the other waterbodies. The upper three waterbodiesare morphologically unaltered but water pollution by pesti-cides, metals and organic pollutants is high and diffuse. The

high human use of the valley and river gives little hope thatthe river will reach good ecological status by 2015.

7.9. THE DURANCE RIVER


The Durance is an alpine Mediterranean river that drains14 322 km2. The tributary Clar!ee is considered as the mainupstream origin of the Durance, although the river lies at#2400 m a.s.l. on the slopes of Montgen"evre Pass on theFrench-Italian border. The Durance merges 320 km down-stream with the Rhone at 12 m a.s.l. The upper catchment(3688 km2 or 25% of the total area) from the source to theUbaye River confluence is alpine. The middle part(8152 km2 or 57%) from the Ubaye to Mirabeau Gorge issubalpine as are the tributaries Bu#ech (1485 km2) and Ver-don (2302 km2). The lower catchment (2565 km2 or 18%)fromMirabeau to Avignon is a Mediterranean river (SDAGERM&C, 1996). The river crosses different geological forma-tions including metamorphic rocks, alternated marls andlimestone. The latter were deposited during the SecondaryEra and folded during the Tertiary, thereby inducing thedeposition of flysch and conglomerates. Limestones domi-nate in the catchment. Pleistocene glaciers covered the upperpart of the catchment and created U-shaped valleys, but theydid not reach the lower catchment. In the middle Durance,Pleistocene terraces still constitute a large part of the valley-floor, while such terraces were eroded in the lower part in theHolocene. The Durance before regulation was a good exam-ple of sedimentary continuity with a regular and high slopeprofile that was linked to flow increases and sediment loadinputs from tributaries. The homogenous character of thesetributary alluvial inputs gave the Durance a more or lessuniform coarse granulometry (SGFH 1916a; Belleudy &Lefort 2001). Excluding some areas (Briancon, Argenti"ere,Saint-Cl!ement, Serre-Poncon, Sisteron and Mirabeau), theriver essentially flows through a 1–6 km wide valley. In thelate 19th century, channel widths ranged from <100 to>1600 m. Water widths in 1890 ranged between 200 and700 m in the middle and lower Durance and <100 m in theupper river (Warner 2000).

Deforestation combined with geomorphic characteristicsand climate was responsible for severe erosion and the braid-ed pattern of the river before 1860. The channel width wasreduced due to a natural decrease in runoff at the end of theLittle Ice Age. Later, the channel width was reduced furtherby channel protection works, gravel extraction, afforestationof side-valleys and mid-20th century regulation schemes(Miramont et al. 1998), resulting in a channel width 30%and wetted width 25% less than that before 1890 and anincrease in agricultural land area of #62 km2 (Warner2000). The river flows through fertile basins which havebeen intensively used for irrigated agriculture for centuries.As early as the late 12th century, canals were used to deliver


water to mills and for irrigation. In 1554, the engineer Adamde Craponne built a canal to irrigate the plain of La Crau, theancient Durance delta (Balland et al. 2002). In the late 19thcentury, diverted flows in the lower Durance exceeded80 m3/s at times, above the lowest summer discharge of42 m3/s in 1870 (Garot 1903). Conflicts caused the Ministryof Agriculture to pass a law in 1907 to control water use.Various projects were initiated (Wilhelm 1913), and the 5January 1955 law was passed to satisfy three objectives:flood control, irrigation, and hydroelectricity (Clebert andRouyer 1991). Large regulation schemes on the Durance andVerdon rivers started afterWorldWar II and finished in 1974.Currently, the hydroelectric network includes 18 power-houses, operated by two major high-head storage schemes:Serre-Poncon dam (1955–1959, 1270 Mm3) on the Duranceand Sainte-Croix dam (1971–1974, 767 Mm3) on the Ver-don. The middle Durance, regulated since 1962, has sevenpower plants supplied by a side-channel carrying 250 m3/s(Reverchon et al. 2006). For flood control, waters from theDurance and tributaries flow through an 185 km by-passsystem from 780 to 0 m asl in the Etang de Berre lagoon.Thewealth and well being of the Provence region has been tothe detriment of the river and alluvial landscape. About250 km of the Durance downstream of Serre-Poncon damcarry 1/40th of the initial annual flow. This low flow isinterspersed by short and large floods, partly controlled bywater retention in reservoirs but which still are geomorpho-logically active (Photo 7.7). The silting of new channels isfast and encroachment of riparian forest regularly requiresmaintenance to hold high flows. Large volumes of freshwaterand silt enter the Etang de Berre, a lagoon of 155 km2 af-fected by urban and industrial wastes, and contribute to itsdegradation.


A Mediterranean climate prevails in the Durance catchment,which receives less annual precipitation than other alpinetributaries of the Rhone. Parde (1925) suggested that modestrainfall was caused by the surrounding mountains that open tosoutheast. Rainfall is minimum in July and maximum in Oc-tober and November, and can be intense. The upper Durancehas a nival flow regime, little influenced by glacial inputs(#1% of the catchment area) but with high snowmelt flowsin spring (maximum in June) and low flows in winter (Table7.3). In the middle and lower Durance, the influence of Med-iterranean rainfalls increases with decreasing altitude. Here,the flows become highly irregular, especially in autumn withlarge floods. Parde (1925) reported that the 19th century highfloods in the Durance at Mirabeau were 5000 m3/s in Novem-ber 1886, 5100 m3/s in October 1882 and 5200 m3/s in No-vember 1843 (436 L/s/km2). Along with afforestation, theDurance-Verdon regulation schemes reduced the probabilityof high floods. However, the risks and socio-economic costs oflarge floods increased in the valleys because of extensivefloodplain developments (Balland et al. 2002). The Durancehydrology changedmostly because of the high-head reservoirsand theirmanagement.Most floods in the upper catchment arestored in Serre-Poncon and Saint-Croix reservoirs.

Short-term temperature data were recorded for the mid-dle Durance near Serre-Poncon and near the Asse tributary.The upper section has cold water released from Serre-Poncondam, temperature then increases rapidly in the first 10 kmdue to the Mediterranean climate (range is 6 !C in summerand 1–2 !C in winter). Maximum summer temperature is#21.5 !C, but 11 !C below the dam (Gras and Gilbert1987). Further downstream, groundwater inputs from

PHOTO 7.7 The regulated middleDurance near Manosque, Alpes-de-Haute-Provence (Photo: Georges Carrel).


alluvial and hillslope aquifers increase spatial variability intemperature. In 1990–1991, water temperature ranged from0 !C in February to 27 !C in July, and the daily range was7–8 !C in spring, 5–6 !C in late summer and autumn, and3–4 !C in winter (Dumont et al. 1993).

7.9.3. Biogeochemistry, Bedload and Sediment

The Durance has hard waters due to sulphates from Triassicgypsum located in the upper catchment. The sediment inputsfrom the Durance are the highest in the Rhone catchment at abedload of 300 000 m3/year and suspended load of600 000 m3/year. The specific erosion exceeds 1000 tons/km2/year. Jurassic black marls or ‘Terres Noires’ cover2200 km2 of the Durance catchment. These areas are oftenunvegetated and subject to weathering and erosion, resultingin a badlands topography with suspended solids in the rivers(Mathys et al. 2003). Results obtained on small experimentalbasins (see Web site: DRAIX ORE) located in the Bl!eoneRiver watershed, a left-hand tributary of the middle Durance,show usual sediment concentration of 100–300 g/L, up to800 g/L on the Laval basin (86 ha). The annual erosion ratereaches 190 tons/year on totally unvegetated area (Mathys etal. 2003). The high sediment loads cause technical, socio-economic and ecological problems on the regulated Duranceby silting by-pass sections, intermediate reservoirs of theDurance-Verdon hydroelectric complex and the Etang deBerre lagoon. The intermediate reservoirs were>50% filledby silt since their setting up from 1959 to 1991, requiringspecial management flushing rules (Reverchon et al. 2006).

7.9.4. Nutrients and Pollution

Upstream to Serre-Poncon reservoir is a remarkable naturalmilieu (Durance valley, Clar!ee River valley, Queyras NaturalRegional Park, and Ecrins National Park) that is relativelypristine but with high tourism pressure of skiing in winterand whitewater sports, boating and bathing in Serre-Ponconin summer. Sewage inputs associated with seasonal peakscause organic pollution in the river near densely populatedresorts. Flow regulation by 29 hydroelectric power plants inthe catchment, and weirs in the mountains for irrigation,increase pollution problems during peak tourism. Down-stream from Serre-Poncon dam, irrigation developmentand agricultural practices caused increases in pollution bypesticides and organics, augmented by minimum flow poli-cies in the river. Pollution by chlorinated solvents in themiddle Durance remains a major problem for the river healthand supply of drinking water.


The algal community of the middle Durance consists of 99species belonging to the Chromophytae (69 species), Chloro-phytae (20), Cyanophytae (7), Euglenophytae (1) and Rhodo-

phytae (2). Diatoms are typical species of mineralized water(Cocconeis pediculus,Cymtopleura solea) andmeso- or poly-saprobic waters (Gomphonema parvulum, Nitzschia palea).Green algae such as Cladophora, Pediastrum and Scenedes-mus are also present. Macroalgae such as Characeae (Charavulgaris var. foetida) and Vaucheriacae (Vaucheria sp.) arecommon. Flow regulation led to the proliferation of filamen-tous Chlorophyceae and the disappearance of stenothermicspecies likeH. foetidus (Chlorophyceae) and Fragilaria arcus(Diatomophyceae) (Cazaubon & Giudicelli 1999).

The upper Durance is considered as a dynamic systemwith pioneer communities and softwood species (T. minima,Calamagrostis littorea, M. germanica, S. eleagnos, andS. purpurea). Downstream, dikes, dams, and water intakesreduce the fluvial dynamics. Downstream from Sisteron, thefloodplain covers a large area that contain shrubs ofS. triandra and S. purpurea, and large trees like Salix alba,A. and dominated by F. angustifolia. They also have somefoothill species such as Q. pubescens and Acer campestre.Mediterranean species found in the lower Durance flood-plain include T. gallica, Acer monspessulanum, Erianthusravennae, Imperata cylindrical, and Spartium junceum.

No data are available on macroinvertebrates before im-poundment and regulation of the river (1950s). The earliestecological studies of the Upper Durance were done between1977 and 1979 (Dumont 1980; Dumont & Rivier 1980).Upstream to Serre-Poncon dam, fauna is typical of alpinerivers with species such as Amphinemura sulcicollis, A. tri-angularis, Capnia vidua, Dictyogenus alpinus, D. cepha-lotes, Nemoura fulviceps, N. mortoni, N. sinuate, P.grandis, Perlodes microcephala, Protonemura brevistyla,P. intricate, P. lateralis, P. nimborum, P. praecox, Rhabdiop-teryx neglecta, Taeniopteryx kuehtreiberi (Plecoptera) andAllogamus auricollis, Drusus discolour, Rhyacophila simu-latrix, and R. torrentium (Trichoptera). First studies of themiddle Durance and lower Asse tributary in 1984 (Pr!evot1984) recorded 207 taxa and showed the simultaneous pres-ence of southern species, Sericostoma galeatum, Agapetuscravensis (Trichoptera),Dryops algiricus (Coleoptera), Teti-simulium bezzii, Wilhelmia mediterranea (Diptera Simulii-dae), and alpine species, Ecdyonurus ruffii (Ephemeroptera),Chloroperla tripunctata, Perla bipunctata, P. marginata(Plecoptera). Simuliidae, Plecoptera (D. cephalotes, Dictyo-genus ventralis, C. tripunctata), Ephemeroptera Heptagen-iidae (Ecdyonurus ruffii, Ecdyonurus lateralis, Rhithrogenasp.) and Oligoneuriidae (Oligoneuriella rhenana) werefound in the Asse, an unregulated gravel-bed river. Macro-invertebrates in the regulated Durance were eurythermic andpotamic. The main species inhabiting the river are Caenismacrura, C. luctuosa, B. lutheri, P. antipodarum, H. exocel-lata, Eusimulium rubzovianum, Hydropsyche modesta,Hydroptila vectis, B. fuscatus, Lymnea ovata peregra, Poly-centropus flavomaculatus, Esolus pygmaeus, Lebertia pul-chella, Psychomyia pusilla, Micronecta meridionalis, andLimnius intermedius. In flowing side-arms, the communitywas dominated by Hirudinea (Hemiclepsis marginata, E.


octoculata), Trichoptera (Hydropsyche incognita), Gastro-poda (P. antipodarum, Ancylus fluviatilis) and Odonata(Coenagrion mercuriale).

In the early 1990s, Cazaubon and Giudicelli (1999)collected 77 taxa in the middle Durance. Densities rangedfrom 1060 to 1900 individuals/m2. Thirty taxa were con-sidered common and represented #90% of the sampledindividuals. The community was composed of potamalspecies such as H. exocellata (the dominant species con-sidered as the ‘fundamental species in the macroinverte-brate community of the regulated Durance’), H. modesta,B. fuscatus, and B. pavidus, and by rhithral species of H.incognita, B. lutheri and Ecdyonurus helveticus (sporadic)and Caenis pusilla, a southern species along with Leuctrageniculata (Plecoptera), Hydroptila vectis, Polycentropusflavomaculatus and Setodes sp. (Trichoptera). Two spe-cies of Simuliidae occurred: Simulium pseudequinum wasabundant during spring and summer while S. bezzii wasabundant in winter. Orthocladiinae (Diptera, Chironomi-dae) were present throughout the year. Because of highsummer temperatures, stenothermic species such as D.cephalotes (Plecoptera) present upstream and just down-stream of Serre-Poncon dam are not found in the middleDurance. As is the case for Dictyogenus ventralis, Iso-perla grammatica, Perla bipunctata, P. marginata (Ple-coptera), Acentrella sinaica, Rhithrogena sp. andEpeorus sp. (Ephemeroptera) that are present in the Asseand Buech rivers. Univoltine species such as the Ephe-meroptera Serattella ignita and O. rhenana are probablyfavoured by the warmer thermal regime in the middleDurance. Other species such as the polyvoltine speciesSimulium bezzi maintain populations by having a singlegeneration in winter. The reduction in discharge and nar-rowing of the riverbed decreased habitat availability andeliminated backwater habitats, leading to the extinction ofspecies such as E. octoculata, Dina lineata, and H. mar-ginata (Hirudinea). Near the confluence with the Rhone,the macroinvertebrate community has several Mediterra-nean taxa like Crustacea Gammarus pulex gallicus. Thetwo exotic species of Crustacea, A. desmarestii (also aMediterranean species) and O. limosus were recorded.

Data of fishes in the Durance come mainly from the greyliterature. Based on Rabotin (2002), Holley and Guibert(2005) and Irz (2006), 43 species from 14 families wererecorded along the Durance river, including the introducedlacustrine species S. alpinus and C. lavaretus. The Durancebasin can be considered a special ichthyoregion in the Rhonecatchment (Changeux & Pont 1995) with high occurrencesof southern cyprinids, soiffe, southern barbel and apron.Longitudinally, the fish assemblages include introducedbrook trout in the headwaters, followed successively bybrown trout, then cyprinids along with blageon and barbel,to migratory species of eel, twaite shad and mugilids M.cephalus, L. ramada near the Rhone confluence. Of the 42species sampled in the Durance, 16 are non-native and in-clude carp and crucian carp. Most non-native species were

introduced in the late 20th century for angling, especiallysalmonids (brook trout, rainbow trout, grayling) and in reser-voirs (C. lavaretus and S. alpinus). Other non-native speciescolonised the Durance from the Rhone, including cyprinidsChondrostoma nasus, Carassius gibelio, Carassius auratus,percids Sander lucioperca, centrarchids Lepomis gibbosus,M. salmoides, the ictalurid Ameiurus melas, and the poeciliidGambusia affinis. The last exotic species were the topmouthgudgeon (1999) and the cobitid Cobitis bilineatawhich sud-denly occurred after 1994. River regulation altered the lon-gitudinal distribution of fishes, partially known from anhistorical fish map (L!eger 1934). Flow regulation causedother ecological changes including an increase in fish growth(Bouchard et al. 1998) and hybridization between two cypri-nidsC. toxostoma andC. nasus (Costedoat et al. 2005, 2007).

Three species of Urodela and seven species of Anoura arefound along the Durance; the common toad, marsh frog andfire salamandra are the most common. The alpine newt ispresent in the upper Durance valley, and this constitutes itssouthern distribution limit. The palmate newt occurs only atone location in the lowerDurance.TheMediterranean tree frogis present far upstream in theDurancevalley up toGap,where-as the Parsley frog occurs in the middle and lower river in lownumbers. Perez’s frog is inhabits the lowerDurance, but popu-lations are decreasing with the expansion of marsh frog.

The dark green snake was recorded only in the middleDurance, and viperine water snake occurs in the middle andlower river. The European pond terrapin colonizes in thelower Durance.

The Durance holds one of the highest species richness inFrancewith over 260 bird species.Most of the French species,except those occurring in mountains and those linked to ma-rine shores, are present. Over 60 species having a EuropeanCommunity status were recorded, including little bittern (20–30 pairs), black kite (100–150 pairs), calandra lark (Melano-corypha calandra) (6–10 pairs, 20%of the French population)and little bustard (Tetrax tetrax) (#15 individuals). Alluvialforests are colonized by several species of herons (little egret,black-crowded night heron, and buff-backed heron). Severalpaludicole (marsh-inhabiting) species occur in reedbeds suchas purple heron, Eurasian bittern (Botaurus stellaris), littlebittern, spotted crake (Porzana porzana), moustached warbler(Acrocephalus melanopogon), and penduline-tit (Remiz pen-dulinus). Gravels bars and movable shores offer habitats forcommon stern, little ringed plover, European bee-eater (Mer-ops apiaster) and common kingfisher.

The European beaver is found in the Durance valley, evenin the upper river (upstream of Sisteron). The southwesternwater vole, muskrat and coypu were recorded only in thelower Durance.


The national and regional socio-economic importance of theDurance basin is considerable. Hydropower generationwhich in <60 years has increased from 95 MW (SGFH


1916b) to 2000 MW provides 10% of the hydroelectric pro-duction in France. The reservoirs guarantee water for irriga-tion and drinking through a complex network of canals.Southwest users benefit from a water supply of 114 m3/sfrom the Durance by-channel and southeast users receive awater supply of 40 m3/s from the Verdon scheme. About100 000 ha in Provence are irrigated through the Durance-Verdon schemes. Water demand linked to tourism in largereservoirs and rivers in the Durance basin has been under-estimated. The high tourism in winter for skiing is causing agrowing demand of water for artificial snow (average3000 m3/ha) and tourism in summer relies on rivers forwhitewater sports and Serre-Poncon reservoir (3000 ha) forbathing and boating. Lake Sainte-Croix and the famous Ver-don canyon have also become a touristic attraction. Tourismnow represents 11% of the gross regional product in theProvence-Alpes-Cote d’Azur.

A large part of the Durance (from 678 to 12 m a.s.l., area:15 954 ha, FR9312003) belongs to the Natura 2000 networkwith many natural habitats in the mountains and Mediterra-nean zone. Durance hydrosystem requires complex mea-sures for rehabilitation that consider all socio-economicand environmental stakeholders. A large governmental mis-sion (Balland et al. 2002) was recently entrusted with thedecision-making processes and coordination between envi-ronmental stakeholders. The main concern is the change offlow regime in the regulated river, the transfer of fine sedi-ment load, and the rehabilitation of Berre lagoon. The trans-port of suspended matter was partly restored up to1.8 million tons (SOGREAH 2000), and flow of the Durancecanal was modified in 1995 to restore the threatened Etangde Berre. Here, freshwater inputs decreased from 3.6 Gm3

per year (1966–1995) to 2.1 Gm3 and floods carrying high-suspended sediment loads were maintained in the formerchannel to reduce sediment deposition in the Etang de Berre.

The upper Durance has three waterbodies that are notconsidered as heavily modified. The most two upstreamwaterbodies have high probability to reach the good statusby 2015, the third one, located on a tributary affected byhydropower and gravel abstraction, has a risk of failing tomeet the good ecological status by 2015. The Serre-Ponconreservoir is classified as heavily modified. Downstream, twowaterbodies are also identified as heavily modified and thelower part upstream of the confluence with the Rhone Riverhas a high risk of failing to meet the WFD objectives by2015. For other waterbobies, there is doubt about their abilityto reach good status by 2015.

7.10. CONCLUSIONS AND PERSPECTIVES

Before regulation, the Rhone basin had a large array of land-scapes and high biological diversity. Tributaries flowingfrom Alps, Jura, Vosges, Massif Central and Cevennes haddifferent morphological, hydrological and ecological char-acteristics that differentially influenced the physical, chem-

ical and biological features of the Rhone. In the Swiss Valais,the first and second river training works and the constructionof high-head storage reservoirs considerably modified thehydrological regime, suspended sediment load, surface wa-ter temperature, ground water level and temperature andoxygen content in Lake L!eman. Present ecological featuresand biodiversity in the Rhone valley are far from those pre-vailing at the end of the 19th century. Today, natural habitatsof the Swiss Rhone floodplain cover only 6% of the wholefloodplain area and analysis of historic maps indicate a lossof 90–100% of alluvial habitats areas. The objective of thethird Rhone correction is to (1) insure effective protectionagainst floods as water levels during flooding are 3–4 mabove the floodplain and dikes can break because most areold and easily damaged, (2) improve the ecological potentialof the floodplain by promoting alluvial areas and restoringnatural processes, and (3) promote economic developmentand tourism in the Valais.

The Arve River flowing from Mont Blanc supplied theRhone with large amounts of coarse sediments up tothe 20th century, whereas today only fine sediments reachthe Rhone because of the reservoirs just downstream ofLake L!eman. From an ecological point of view, the Arvecontributes cold water to the Rhone and modifies its phy-sico-chemistry. The presence of ‘Perte du Rhone’ a fewkilometres downstream from the lake played an importantrole for the colonization of the upper river by fishes. Thefirst French hydroelectric power station was built in 1884on the Valserine River near its confluence with the Rhoneand another was constructed in 1899 on the Rhone up-stream of ‘Perte du Rhone’; they contributed greatly tolocal industrial development. The gorge was used to buildG!enissiat dam, the head of a series of six hydropowerplants that greatly modified the physical and biologicalcharacteristics of the river. The confluence of Rhone withthe Saone River at Lyon is a major source of hydrological,physico-chemical and biological discontinuity along theFrench Rhone River. The urbanization and industrial de-velopment of Lyon, in peculiar downstream, has a majorinfluence on water quality in the Rhone. Downstream,arrival of cold water from the Is"ere allows the recoveryof several fishes present in the French upper Rhone. Thebeginning of the Mediterranean zone and the presence ofseveral tributaries downstream from the Is"ere lead to aprogressive change in riverine features, especially theincrease in water temperature. Downstream from Arles,the delta of Camargue constitutes an exceptional land-scape that, despite human interventions to limit damagescaused by large floods, remains an area of great ecologi-cal, cultural and sociological importance. Special atten-tion has been paid recently to improve flood managementdownstream of Vallabr"egues by creating overflow areasupstream and by increasing embankment security. Sedi-ment deficit in the delta led to a progressive advance ofthe sea, and studies are being conducted to find ways torestore sediment transport to the Rhone delta.


The natural functioning of the main tributaries Arve, Ain,Saone, Is"ere, Drome, Ard"eche, Durance, and Gard has beengreatly altered for the past 200 years. The main environmen-tal problems on the Ain are linked to flow management forhydroelectrical production and groundwater extraction forirrigation in the lower valley. The hydroelectric facility in themiddle valley led to the loss of areas with high morpholog-ical and ecological interest. Protection of the lower river andassociated wetlands and biodiversity is presently a greatchallenge. The Saone valley has been strongly altered byhuman activities (channel regulation and dredging for navi-gation, industrialization, agriculture) and, despite the will torestore and improve the morphology and ecology of the riverfloodplain, management goals to maintain existing econom-ic activities and diminish pollution levels (toxic pollution,pesticides), a protection program against floods, and en-hancement of ecological functions of the river remains adifficult task. The Durance catchment is extensively usedfor hydroelectricity production, irrigation, and drinking wa-ter production. Geomorphological and ecological processesare heavily altered by water abstraction and managementobjectives in the WFD context are quite difficult to attain.

At the watershed scale, much effort has been made tomove from a traditional strategy of flood defence to one offlood riskmanagement. Local populations and water manage-ment committees joined efforts to analyse the main causes offlooding and elaborate policies and action plans to reduce riskstochasticity, diminish vulnerability, and manage residualrisk. All these measures must be integrated in a more generalprogram that includes the set up of theWFD, the protection ofsurface and groundwater quality and quantity, and the preser-vation of biodiversity. The French government, WatershedManagement authorities, ‘Agence de l’Environnement et dela Maitrise de l’Energie’, ‘Voies Navigables de France’, localRegional Councils (Languedoc-Roussillon, Provence-Alpes-Cote-d’Azur, Rhone-Alpes, Bourgogne et Franche-Comt!e,)and the ‘Compagnie Nationale du Rhone’ signed a globalprogram towards sustainable development of the Rhone andits valley named ‘Plan Rhone’. This ambitious program hassix themes: (1) a patrimonial and cultural section devoted tothe improvement of the knowledge of cultural developmentlinked to the Rhone River along the valley, and a new appro-priation of the river by the public at large, (2) a global strategyto prevent flood risks, (3) a program ofwater quality improve-ment (WFD) and achievement of the inventory of Sites ofCommunity importance (Natura 2000 sites, Saone floodplainand Camargue), (4) promotion of hydroelectricity and eolianenergy production, (5) promotion of fluvial transport along theRhone-Saone axis especially by harbour infrastructure im-provement, and (6) development of tourism (river boating,‘v!eloroute’). The cost of this program (2006–2013) will be>800 Me.

After the period of river regulation and deteriorationduring the 20th century, the will to insure good water qualityand quantity has progressively led to the awareness of thenecessity to protect and restore, as much as possible, the

ecological functioning of rivers. Recent large floods, bothin the Swiss (Valais) and French part of the river (especiallythe lower Rhone and Camargue), have forced authorities toreconsider sediment and water transfer along the river.Knowledge about the impact of pesticides and toxins onthe biology is still insufficient and research is urgently need-ed. New problems linked to global climate change are oc-curring, especially regarding water temperature. To meetcurrent objectives (i.e. river ecological restoration and objec-tives of the Water Framework Directive) is a great challengeas well as for the small and large rivers of the Rhone catch-ment because of the necessity to integrate socio-economicand ecological objectives.

Acknowledgements

We thank the following people for their contributions tothe preparation of this chapter: Corinne Mass!e for help indata collection and preparation, Alain Poirel for watertemperature data on the French part of the Rhone Riverand tributaries, Fr!ederique Eyrolles (Laboratoire d’!etudesRadio!ecologiques en milieu Continental et Marin) fordata about radionuclides, Vincent Passani for hydrologicaldata of the Durance watershed, Jacky Girel, Henri Persat,Jean-Claude Rostan, Gudrun Bornette, Jean-Francois Fru-get, Sylvain Dol!edec, Bernard Dumont, Pierre Joly, San-drine Plenet and Eric Feunteun for their valuable help anddiscussions about biodiversity of the Rhone River water-shed. The Mus!eum National d’Histoire Naturelle providedhelpful data on biodiversity along river corridors ([Ed].2003–2006; Inventaire national du Patrimoine naturel(Web site: http://inpn.mnhn.fr/)). We thank Electricit!e deFrance (EDF) and the Compagnie Nationale du Rhone(CNR) for providing details about discharge, dam featuresand hydroelectricity production, the Commission interna-tionale pour la protection des eaux du L!eman for provid-ing physico-chemical data of the Rhone RiverDownstream from L!eman Lake and the Federal Officefor the Environment (FOEN) for hydrological and tem-perature data in the Swiss Upper Rhone valley.

REFERENCES

Agence de l’Eau Rhone-M!editerran!ee-Corse. 2004. Pesticides dans les eauxsuperficielles et souterraines des bassins Rhone-M!editerran!ee et deCorse – Donn!ees 2002 et 2003. Agence de l’Eau Rhone M!editerran!eeCorse, Lyon.

Alliot, E., Younes, W.A.N., Romano, J.C., Rebouillon, P., and Masse, H.2003. Biogeochemical impact of a dilution plume (Rhone River) oncoastal sediments: comparison between a surface water survey (1996–

2000) and sediment composition. Estuarine, Coastal and Shelf Science57: 357–367.

Amoros, C. 2001. The concept of habitat diversity between and within

ecosystems applied to river side-arm restoration. Environmental Man-

agement 28: 805–817.


http://inpn.mnhn.fr/

Amoros, C., Rostan, J.C., Pautou, G., and Bravard, J.P. 1987a. The reversible

process concept applied to the environmental management of large riversystems. Environmental Management 11: 607–617.

Amoros, C., Roux, A.L., Reygrobellet, J.L., Bravard, J.P., and Pautou, G.

1987b. Amethod for applied ecological studies of fluvial hydrosystems.Regulated Rivers: Research & Management 1: 17–36.

Amoros, C., Bravard, J.P., Reygrobellet, J.L., Pautou, G., and Roux, A.L.1988. Les concepts d’hydrosyst"eme et de secteur fonctionnel dans

l’analyse des syst"emes fluviaux au niveau des !ecocomplexes. Bulletind’Ecologie 19(4): 531–546.

Amoros, C., Piegay, H., Dufour, S., Grospretre, L., Piegay, H., and Henry, C.

2005. Flood scouring and groundwater supply in rehabilitated side-channels of the Rhone River, France: sedimentation and aquatic vege-tation responses. Archiv f#ur Hydrobiologie 15, Supplementband 155:

147–168.Anon. 1938. Le Rhone. Administrative Report.Antonelli, C., and Provansal, M. 2002. Characterisation and assessment of

sand fluxes in the lower Rhone river, France. Proceedings of River Flow

2002, Louvain–la-Neuve, Belgique,Balkema, Rotterdam.ARALEP. 2003. Suivi hydrobiologique du Rhone au niveau de la centrale

nucl!eaire de Saint-Alban – Saint-Maurice l’Exil. Synth"ese 1985–2001.

Report to Electricit!e de France. ARALEP, Villeurbanne.Arnaud-Fassetta, G., and Provansal, M. 1999. High frequency variations of

water flux and sediment discharge during the Little Ice Age (1586–1725

AD) in the Rhone Delta (Mediterranean France), Relationship to thecatchment basin. Hydrobiologia 410: 241–250.

Astrade, L. 2005. La Saone en crue. Dynamique d’un hydrosyst"eme anthro-

pis!e, Presses Universitaires de Lyon, Lyon.Balland, P., Huet, P., Lafont, E., and Leteurtois, J.P. 2002. Propositions de

simplification et de modernisation du dispositif d’intervention de l’Etatsur la gestion des eaux et du lit de la Durance. Contribution "a un Plan

Durance. Conseil G!en!eral des Mines, Minist"ere de l’Environnement etdu D!eveloppement Durable – Minist"ere de l’Agriculture, del’Alimentation, de la Peche et des Affaires Rurales –Minist"ere D!el!egu!e"a l’Industrie – Minist"ere de l’Equipement, des Transports, du Loge-ment, du Tourisme et de la Mer, Paris.

Barbe, J. 1997. Chasses franco suisses (1997). Le Vieux Rhone de Chau-

tagne: d!etermination des conditions physiques du milieu et des pro-

lif!erations v!eg!etales, Cemagref, Lyon.Barbe, J., and Barthelemy, R. 1994. Conditions de d!eveloppement et gestion

des prolif!erations de cyanobact!eries sur le vieux Rhone de Belley:

rapport final. Cemagref, Lyon.Baumann, P. 2004. Revitalisierung und Benthos der Rhone. Schlussbericht

SP I-6. Rhone-Thur Projekt, EAWAG, WSL, Limnex, AG.

Belleudy, P., and Lefort, P. 2001. La continuit!e s!edimentaire et sa rupture. Lagestion du transport solide: de la source "a la mer,Soci!et!e Hydrauliquede France, Lyon.

Berrahou, A. 1993. R!epartition longitudinale des macroinvert!ebr!es benthi-ques du Rhone francais et sa relation avec les principaux affluents.Doctoral thesis, University Lyon, p. 1.

Bornette, G., Amoros, C., Pi!egay, H., Tachet, J., and Hein, T. 1998. Eco-logical complexity of wetlands within a river landscape. BiologicalConservation 85: 35–45.

Bornette, G., Piegay, H., Citterio, A., Amoros, C., and Godreau, V. 2001.

Aquatic plant diversity in four river flood plains: a comparison attwo hierarchical levels. Biodiversity and Conservation 10:1683–1701.

Bouchard, P., Chappaz, R., Cavalli, L., and Brun, G. 1998. Influence ofenvironmental variables on the growth of Leuciscus cephalus (Linnaeus1766), in the River Durance, South-East France. Annales de Limnologie

34: 193–200.

Boulton, A.J., Findlay, S., Marmonier, P., Stanley, E.H., and Valett, M.H.

1998. The functional significance of hyporheic zone in streams andrivers. Annual Review of Ecology and Systematics 29: 59–81.

Bournaud, M., Tachet, H., and Perrin J.F. 1982. Les Hydropsychid!es (Tri-

choptera) du Haut-Rhone entre Gen"eve et Lyon. Annles de Limnologie18(1): 61–80.

Bournaud, M., Cellot, B., Richoux, P., and Berrahou, A. 1996. Macroinver-tebrate community structure and environmental characteristics along a

large river: congruity of patterns for identification to species or family.Journal of the North American Benthological Society 15(2): 232–253.

Braudel, F. 1986. L’ identit!e de la France. Espace et histoire. 3 tomes,

Arthaud, Paris.Bravard, J.P. 1987. Le Rhone du L!eman "a Lyon. La Manufacture, Lyon.Bravard, J.P., Amoros, C., and Pautou, G. 1986a. Impact of civil engineering

works on the successions of communities in a fluvial system. A Meth-

odological and Predictive Approach Applied to a Section of the Upper

Rhone River Vol. 47: Oikos, France, , 92–111.Bravard, J.P. Combier, J. and Commercon, N. (eds). 2002a. La Saone, axe de

civilisation, Presses Universitaires de Lyon, Lyon.Bravard, J.P., Peiry, J.L., and Salvador, P.G. 2002b. La diversit!e des enregis-

trements morpho-s!edimentaires tardiglaciaires et holoc"enes dans quel-

ques vall!ees du pi!emont des Alpes du Nord (Haut-Rhone, Arve, Is"ere).In: Les Fleuves ont une histoire: pal!eo-environnement des rivi"eres et deslacs francais depuis 15000 ans, Editions Errance, Bravard, J.P., Magny,

M. (eds). The River Handbook Hydrological and Ecological principlesVol. 1: Blackwell Scientific Publications, Oxford, , 426–447.

Bravard, J.-P., Roux, A.L., Amoros, C., Richardot-Coulet, M., Reygrobellet,

J.L., Bournaud, M., and Pautou, G. 1986b. Evolution spatio-temporelledes syst"emes fluviaux am!enag!es: recherches m!ethodologiques sur leHaut Rhone francais. La Houille blanche 4(18): 1–8.

Brunke, M., and Gonser, T. 1997. The ecological significance of exchange

processes between rivers and groundwater. Freshwater Biology 37:1–33.

Castella, E. 1987. Apport des macroinvert!ebr!es aquatiques au diagnostic

!ecologique des !ecosyst"emes abandonn!es par les fleuves. Recherches

m!ethodologiques sur le Haut-Rhone francais. Vols. 1 and 2, Doctoralthesis, University Lyon, p. 1.

Castella, E., Richardot-Coulet, M., Roux, C., and Richoux, P. 1991. Aquaticmacroinvertebrate assemblages of two contrasting floodplains: theRhone and Ain Rivers, France. Regulated Rivers: Research and Man-

agement 6: 289–300.Castella, E., Terrier, A., Pellaud, M., and Paillex, A. 2005. Distribution

d’Anisus vorticulus (Troschel 1834) dans la plaine alluviale du Haut-Rhone francais. Un Gast!eropode Planorbidae list!e en annexe de la

|Directive Habitats}. Bulletin de la Soci!et!e Linn!eenne de Lyon,

pp. 1–11.Cavalli, L., Pech, N., and Chappaz, R. 2003. Diet and growth of the endan-

gered Zingel asper the Durance River. Journal of Fish Biology 63:460–471.

Cazaubon, A., and Giudicelli, J. 1999. Impact of the residual flow on the

physical characteristics and benthic community (algae, invertebrates) ofa regulated Mediterranean river: the Durance, France. Regulated Riv-

ers: Research and Management 15: 441–461.CEA. 2005. Informations sur l’!energie 2005. Commissariat "a l’Energie

Atomique, Paris.Cellot, B. 1996. Influence of side-arms on aquatic macroinvertebrate drift in

the main channel of a large river. Freshwater Biology 35: 149–164.

Changeux, T. 1995. Structure du peuplement piscicole"a l’!echelle d’un grandbassin europ!een: organisation longitudinale, influence de la pente ettendances r!egionales.Bulletin Francais de la Peche et de la Pisciculture337/338/339: 63–74.


Changeux, T., and Pont, D. 1995. Ichthyogeographic regions and watershed

size in the French river Rhone network. Hydrobiologia 300/301:355–363.

Clebert, J.P., and Rouyer, J.P. 1991. La Durance. Rivi"eres et vall!ees de

France. Privat, Toulouse.Cog!erino, L. 1989. Les rives aquatiques de grands cours d’eau : car-

act!erisation m!esologique et faunistique. Doctoral thesis, UniversityLyon, p. 1.

Conseil Sup!erieur de la Peche – Protection des Milieux Aquatiques. 2005.Synth"ese nationale du suivi national de la peche aux engins pour lap!eriode 1999 "a 2002. Report.

Costedoat, C., Pech, N., Salducci, M.D., Chappaz, R., and Gilles, A. 2007.Novelties in hybrid zones: crossroads between population genomic andecological approaches. PLoS ONE 2: e357. doi:10.1371/journal.

pone.0000357.Costedoat, C., Pech, N., Salducci, M.D., Chappaz, R., and Gilles, A. 2005.

Evolution of mosaic hybrid zone between invasive and endemic speciesof Cyprinidae through space and time. Biological Journal of the Lin-

nean Society 85: 135–155.Craney, E., and Piston, H. 1994. Les amphibiens du lit majeur de la Saone –

Inventaire et !ecologie – Cartographie des biotopes. Report.

Creuz!e des Chatelliers, M. 1991. Dynamique de r!epartition des bioc!enosesinterstitielles du Rhone en relation avec des caract!eristiques

g!eomorphologiques (secteurs de Br!egnier-Cordon, Miribel-Jonage,

Donz"ere-Mondragon). Doctoral thesis, University of Lyon.Creuz!e des Chatelliers, M., and Reygrobellet, J.L. 1990. Interactions be-

tween geomorphological processes, benthic and hyporheic communi-

ties: first results on a by-passed canal of the French Upper-Rhone River.Regulated Rivers: Research & Management 5: 139–158.

Crivelli, A. 1998. L’anguille dans le bassin Rhone-M!editerran!ee-Corse :

une synth"ese bibliographique. Report COGEPOMI Rhone-

M!editerran!ee-Corse.Culver, D.C., and Sket, B. 2000. Hotspots of subterranean biodiversity in

caves and wells. Journal of Cave and Karst Studies 62(1): 11–17.

Curt, T., Prevosto, B., and Bergonzini, J.C. 2004. Boisements naturels desterres agricoles en d!eprise. Cemagref Editions, Antony.

Dahm, C.N., Grimm, N.B., Marmonier, P., Valett, H.M., and Vervier, P.

1998. Nutrient dynamics at the interface between surface waters andgroundwaters. Freshwater Biology 40: 427–451.

Danancher, D. 2005. Apport de l’!ecologie comportementale "a la conserva-

tion d’un poisson en voie de disparition: l’Apron du Rhone (Zingel

asper). Doctoral thesis, University Lyon, p. 1.Dangreaux, B. 2002. Le Rhone et le vin: du vin des cimes au vin des sables.

Gl!enat, Grenoble.

Danielopol, D.L., and Pospisil, P. 2001. Hidden biodiversity in the ground-water of the Danube floodplain National Park (Austria). Biodiversityand Conservation 10: 1711–1721.

Danielopol, D.L., Gibert, J., Griebler, C., Gunatilaka, A., Hahn, H.J., Mes-sana, J., Notenboom, J., and Sket, B. 2004. Incorporating ecologicalperspectives in European groundwater management policy. Environ-

mental Conservation 31(3): 185–189.Daufresne, M., Bady, P., and Fruget, J.F. 2007. Impacts of global changes

and extreme hydroclimatic events on macroinvertebrate communitystructures in the French Rhone River. Oecologia 151: 544–559.

Daufresne, M., Roger, M.C., Capra, H., and Lamouroux, N. 2003. Long-term changes within the invertebrate and fish communities of the UpperRhone River: effects of climatic factors. Global Change Biology 10:

124–140.Delrieu, G., Ducrocq, V., Gaume, E., Nicol, J., Payrastre, O., Yates, E.,

Kirstetter, P.E., Andrieu, H., Ayral, P.A., Bouvier, C., Creutin, J.D.,

Livet, M., Anquetin, S., Lang, M., Neppel, L., Obled, C., Parent-Du-

Chatelet, J., Saulnier, G.M., Walpersdorf, A., and Wobrock, W. 2005.

The catastrophic flash-flood event of 8–9 September 2002 in the Gardregion, France: A first case study for the Cevennes-Vivarais Mediterra-nean Hydrometeorological Observatory. Journal of Hydrometeorology

6: 34–52.Dessaix, J., Fruget, J.F., Olivier, J.M., and Beffy, J.L. 1995. Changes of the

macroinvertebrate communities in the dammed and by-passed sectionsof the FrenchUpper Rhone after regulation.Regulated Rivers: Research

and Management 10: 265–279.Dethier, M., and Castella, E. 2002. A ten years survey of longitudinal

zonation and temporal changes of macrobenthic communities in the

Rhone River, downstream from Lake Geneva (Switzerland). Annales deLimnologie 38(2): 151–162.

DIREN. 2005. Etat des lieux du bassin du Rhone et des cours d’eau cotiers

m!editerran!eens. Caract!erisation du district et registre des zonesprot!eg!ees. Document adopt!e par le Comit!e de Bassin du 4 mars 2005.Secr!etariat Technique SDAGE - DCE, Direction R!egionale del’Environnement Rhone-Alpes, Bassin Rhone-M!editerran!ee, Agence

de l’Eau - Rhone, M!editerran!ee & Corse, Lyon.Dole, M.-J. 1983. Le domaine aquatique souterra in de la plaine alluviale du

Rhone "a l’Est de Lyon, 1. Diversit!e hydrologique et bioc!enotique de

trois stations repr!esentatives de la dynamique fluviale. Vie et Milieu 33(3–4): 219–229.

Dole, M.-J., and Chessel, D. 1986. Stabilit!e physique et biologique des

milieux interstitiels, Cas de deux stations du Haut Rhone. Annales deLimnologie 22: 69–81.

Dole-Olivier, M.-J., and Marmonier, P. 1992a. Patch distribution of

interstitial communities: prevailing factors. Freshwater Biology

27: 177–191.Dole-Olivier, M.-J., and Marmonier, P. 1992b. Effects of spates on intersti-

tial assemblages structure. Disturbance-perturbation relationship, rate

of recovery. Hydrobiologia 230: 49–61.Dole-Olivier, M.-J., Creuz!e des Chatelliers, M., and Marmonier, P. 1993.

Repeated gradients in subterranean landscape. Example of the stygo-

fauna in the alluvial floodplain of the Rhone River (France). Archiv f#urHydrobiologie 127(4): 451–471.

Dole-Olivier, M.-J., Marmonier, P., and Beffy, J.L. 1997. Response of inver-

tebrates to lotic disturbance: is the hyporheic zone a patchy refugium?Freshwater Biology 37(2): 257–276.

Dole-Olivier, M.-J., Marmonier, P., Creuz!e des Chatelliers, M., and Martin,

D. 1994. Interstitial fauna associated with the alluvial floodplains of theRhone River (France). In: Gibert, J., Danielopol, D.L., Stanford, J.A.(eds).Groundwater Ecology,Academic Press, San Diego, pp. 313–346.

Dol!edec, S. (ETEC). 2000. Traitement statistique des donn!ees hydrobiolo-

giques du Canton du Valais. Rapport d’expertise. Service de la protec-tion de l’Environnement, Canton du Valais.

Doutriaux, E. 2006. Am!enagements sur le Rhone francais et bilan

s!edimentaire. Congr"es du Rhone, Gen"eve.Dumont, B. 1980. Les types d’am!enagement hydro-!electrique en zone

montagnarde et leurs effets sur la vie aquatique. In: Grover, J.H.

(ed). Proceedings of the Technical Consultation on Allocation of

Fishery Resources held in Vichy, France, 20–23 April 1980,Pub-lished by arrangement with the Food and Agriculture Organizationof the United Nations in cooperation with the American Fisheries

Society, pp. 93–96.Dumont, B., and Rivier, B. 1980. Equipements hydro-!electriques et vie

aquatique en milieu Alpin. In: Grover, J.H. (ed). Proceedings of the

Technical Consultation on Allocation of Fishery Resources held in

Vichy, France, 20–23 April 1980,Published by arrangement with theFood andAgriculture Organization of theUnitedNations in cooperation

with the American Fisheries Society, pp. 97–108.


doi:10.1371/journal.pone.0000357

doi:10.1371/journal.pone.0000357

Dumont, B., Le Coarer, Y., and Carrel, G. 1993.Moyenne Durance. Site du

Largue. Etude du d!ebit r!eserv!e. Morphodynamique et ichtyologie. U.R.Hydrobiologie, Cemagref Aix-en-Provence, Le Tholonet.

Eyrolle, F., Charmasson, S., and Louvat, D. 2004. Plutonium isotopes in the

lower reaches of the River Rhone over the period 1945–2000: fluxestowards the Mediterranean Sea and sedimentary inventories. Journal ofEnvironmental Radioactivity 74: 127–138.

Eyrolle, F., Louvat, D., Metivier, J.M., and Rolland, B. 2005. Origins and

levels of artificial radionuclides within the Rhone river waters (France)for the last forty years: Towards an evaluation of the radioecologicalsensitivity of river systems. Radioprotection 40: 435–446.

Eyrolle, F., Duffa, C., Antonelli, C., Rolland, B., and Leprieur, F. 2006.Radiological consequences of the extreme flooding on the lower courseof the Rhone valley (December 2003 South East France). Science of The

Total Environment 366: 427–438.Fatio, V. 1882. Histoire naturelle des poissons – premi"ere partie. H. Georg,

Geneva, Basel.Fatio, V. 1890.Histoire naturelle des poissons – deuxi"eme partie. H. Georg,

Geneva, Basel.Ferreira, D., Dole-Olivier, M.J., Malard, F., Deharveng, L., and Gibert, J.

2003. Faune aquatique souterraine de France: base de donn!ees et!el!ements de biog!eographie. Karstologia 42: 15–22.

Foulquier, L., Garnier-Laplace, J., Descamps, B., Lambrechts, A., and Pally,M. 1991. Exemples d’impact radio!ecologique de centrales nucl!eaires

sur des cours d’eau francais. Hydro!ecologie Appliqu!ee 3: 149–208.Fruget, J.F. 1991. The impact of river regulation on the lotic macroinverte-

brate communities of the lower Rhone, France. Regulated Rivers: Re-

search and Management 6: 241–255.Fruget, J.F., and Persat, H. 2000. Changement de l’!equilibre hydrobiologi-

que de la Basse Saone. Impact de l’eutrophisation et de la contaminationtoxique. Unpublished report, Grand Lyon.

Fruget, J.F., and Dessaix, J. 2002. Biodiversit!e structurelle et fonctionnelledes peuplements de macroinvert!ebr!es en tant que descripteur de lavariabilit!e hydraulique: exemple de deux parties court-circuit!ees du

Rhone Moyen. Revue des Sciences de l’Eau 15(1): 209–221.Fruget, J.F., and Bady, P. 2006. Etude des relations entre les variables

d’environnement et les invert!ebr!es benthiques "a l’!echelle du fleuve.

1985–2004. Etude Thermique phase III – Lot 2. Report.Fruget, J.F., Tachet, H., and Pont, D. 1995. Structure of macroinvertebrate

communities in the Rhone river delta area. Abstracts of the First Inter-

national Symposium on the Ecology of Large Rivers,Krems (Autriche),04/1995.

Fruget, J.F., Dessaix, J., and Plenet, S. 1996. Macroinvertebrate communi-ties of the Doubs River prior to completion of the Rhine-Rhone con-

nection. Regulated Rivers: Research and Management 12: 617–631.Fruget, J.F., Centofanti, M., and Olivier, J.M. 1998. The fish fauna of the

Doubs River prior to the completion of the Rhine-Rhone connection.

Environmental Management 22(1): 129–144.Fruget, J.F., Centofanti, M., Dessaix, J., Olivier, J.M., Druart, J.C., and

Martinez, P.J. 2001. Temporal and spatial dynamics in large rivers:

example of a long term monitoring of the middle Rhone River. Annalesde limnologie 37(3): 237–251.

Gargani, J. 2004. Modelling of the erosion in the Rhone valley during theMessinian crisis (France).Quaternary International. Advances in Qua-

ternary studies 121: 13–22.Garot, E. 1903. La r!egularisation de la Durance. La Nature. Revue des

sciences et de leurs applications aux arts et "a l’industrie 31: 375.

Gerdeaux, D. 2001. L’omble chevalier, les Cor!egones. In: Keith, P.,Allardi, J. (coord.). Atlas des poissons d’eau douce de France,Patrimoines naturels. Publications Scientifiques du M.N.H.N., Paris,

pp. 263–264.

Gibert, J., Stanford, J.A., Dole-Olivier, M.J., and Ward, J.V. 1994. Basic

attributes of groundwater ecosystems and prospects for research. In:Gibert, J., Danielopol, D.L., Stanford, J.A. (eds).Groundwater Ecology,Academic Press, San Diego, pp. 7–40, (Chapter 1).

Gibert, J., Ginet, R., Mathieu, J., Reygrobellet, J.L., and Seyed-Reihani, A.1977. Structure et fonctionnement des !ecosyst"emes du Haut Rhonefrancais. IV. Le peuplement des eaux phr!eatiques, premiers r!esultats.Annales de limnologie 13(1): 83–97.

Girel, J., Vautier, F., and Peiry, J.L. 2003. Biodiversity and land-use historyof the Alpine riparian landscapes: the example of the Is"ere River Valley,France. In: Mander, U., Antrop, M. (eds).Multifunctional Landscapes,

Vol. 3 Continuity and Change, WIT Press, Southampton, Boston,pp. 167–200.

Giudicelli, J., Dia, A., and Legier, P. 1980. Etude hydrobiologique d’une

rivi"ere de r!egion m!editerran!eenne l’Argens (Var, France). Habitats,hydrochimie, distribution de la faune benthique. Bijdragen tot de Dier-kunde 50: 303–341.

Glennie, E.B., Littlejohn, C., Gendebien, A., Hayes, A., Palfrey, R., Sivil, D.,

and Wright, K. 2002. Phosphates and alternative detergent builders.Final report. Report No.UC 4011. EU Environment Directorate, WRcSwindon, Wiltshire.

Gobin, M. 1868. Note sur les ressources que pr!esente actuellement le Haut-Rhone au point de vue de la peche. Lu "a la Soci!et!e Imp!erialed’agriculture, histoire naturelle et arts utiles de Lyon, dans la s!eance

du 20 mars 1868.Godreau, V., Bornette, G., Frochot, B., Amoros, C., Castella, E., Oertli, B.,

Chambaud, F., Oberti, D., and Craney, E. 1999. Biodiversity in the

floodplain of Saone: a global approach. Biodiversity and Conservation

8: 839–864.Golterman, H.L., andMeyer, M. 1985a. The geochemistry of two hard water

rivers, the Rhine and the Rhone Part 1: Presentation and screening of

data. Hydrobiologia 126: 3–10.Golterman, H.L., andMeyer, M. 1985b. The geochemistry of two hard water

rivers, the Rhine and the Rhone. Part 2: The apparent solubility of

calcium carbonate. Hydrobiologia 126: 11–19.Golterman, H.L., andMeyer, M. 1985c. The geochemistry of two hard water

rivers, the Rhine and the Rhone. Part 3: The relations between calcium,

bicarbonate, sulphate and pH. Hydrobiologia 126: 21–24.Golterman, H.L., andMeyer, M. 1985d. The geochemistry of two hard water

rivers, the Rhine and the Rhone. Part 4: The determination of the

solubility product of hydroxy-apatite. Hydrobiologia 126: 25–29.Gourret P. 1897. Les !etangs saumatres du Midi de la France et leurs

pecheries Annales du Museum d’histoire naturelle de Marseille – Zool-ogie, Tome V, MN Edition Mouillot Fils Ain Marseille.

Gras, R., and Gilbert, A. 1987. R!egime thermique de la Durance entreEspinasse et Tallard. Influence des d!ebits. Direction des Etudes et

Recherches, EDF, Chatou.

Grenouillet, G., Pont, D., andOlivier, J.M. 2000. Habitat occupancy patternsof juvenile fishes in a large lowland river: interactions with macro-phytes. Archiv f#ur Hydrobiology 149(2): 307–326.

Grenouillet, G., Pont, D., and Olivier, J.M. 2001. Linking zooplanktonand juvenile fish assemblages in a large lowland river: influence ofsubmerged macrophytes. Archiv f#ur Hydrobiology 151(3):383–404.

Guilbaud, C., Hollan, E., Wahl, B., Duwe, K., Lemmin, U., Umlauf, L., andFilocha, M. 2004. EUROLAKES – Integrated Water Resource Manage-

ment for Important Deep European Lakes and their Catchment Areas.

Internal Seiche Mixing Study, European Commission. http://www.eurolakes.org/.

Holley, J.-F. and Guibert, A. 2005. DCE et plans d’eau – Compte-rendu

d’avancement 2004 – Annexes 4 et 5. Cemagref, Montpellier.


http://www.eurolakes.org/

http://www.eurolakes.org/

HYDRONAT. 2000. Troisi"eme correction du Rhone. Rapport de synth"ese.

Canton du Valais, Sion.Ilg, C., and Castella, E. 2006. Patterns of macroinvertebrates traits along

three glacial stream continuums. Freshwater Biology 51: 840–853.

Illies, J., and Botosaneanu, L. 1963. Probl"emes et m!ethodes de la classifi-cation et de la zonation !ecologique des eaux courantes, consid!er!eessurtout du point de vue faunistique. Com Ass Intern Limnol Th!eor Appl

12: 1–57.

Irz, P. 2006. Approche comparative des communaut!es piscicoles des plansd’eau. Doctoral thesis, University Montpellier II.

Jakob, A., Binderheim-Bankay, E., and Davis, J.S. 2002. National long-term

surveillance of Swiss rivers. Verhundlungen Internationale VereinigungLimnologie 28: 1101–1106.

Juberthie, C. 1995. Underground habitats and their protection. Nature and

Environment Vol. 72 Council of Europe Press, Strasbourg, .Klingeman, P.C., Bravard, J.P., Giuliani, Y., Olivier, J.M., and Pautou, G.

1998. Hydropower reach by-passing and dewatering impacts ingravel-bed rivers. In: Klingeman, P.C., Beschta, R.L., Komar, P.D.,

Bradley, J.B. (eds). Gravel-bed Rivers in the Environment, WaterResources Publications, LLC, Highlands Ranch, Colorado,pp. 313–344.

Knispel, S. 2004. Temporal and Spatial Dynamics of Benthic Invertebrate

Communities in an Alpine Glacier-fed Alluvial System. Doctoral thesis,University of Lausane.

Kreitmann, L. 1931. Etude d’hydrobiologie des eaux alpines, Carte piscicoledu d!epartement de la Haute-Savoie. Travaux du Laboratoire

d’Hydrobiologie et de Pisciculture de l’Universit!e de Grenoble 23:

145–155.Kreitmann, L. 1932. Les grandes lignes de l’!economie piscicole du bassin

francais du Rhone. Travaux du Laboratoire d’Hydrobiologie et de Pi-

sciculture de l’Universit!e de Grenoble 24: 127–130.

K#uttel, S. 2001. Bedeutung der Seitengew#asser der Rhone f#ur die nat#urlicheReproduktion der Bachforelle und Diversit#at der Fischfauna im Wallis.Doctoral thesis, Swiss Federal Institute of Technology Zurich.

Labonne, J., andGaudin, P. 2005. Exploring population dynamics patterns ina rare fish, Zingel asper, through capture-mark-recapture methods.Biological Conservation 19: 463–472.

Labonne, J., Allouche, S., and Gaudin, P. 2003. Use of generalized linearmodel to test habitat preferences: the example of Zingel asper, anendemic endangered percid of the Rhone River. Freshwater Biology

48: 687–697.Lambrechts, A., and Foulquier, L. 1987. Radioecology of the Rhone Basin:

data on the fish of the Rhone (1974–1984). Journal of EnvironmentalRadioactivity 5: 105–121.

Lamouroux, N., Olivier, J.M., Persat, H., Pouilly, M., Souchon, Y., andStatzner, B. 1999. Predicting community characteristics from habi-tat conditions: fluvial fish and hydraulics. Freshwater Biology 42:

1–25.Lamouroux, N., Olivier, J.M., Capra, H., Zylberblat,M., Chandesris, A., and

Roger, P. 2007. Fish community changes after minimum flow increase:

testing qunatitative predictions in the Rhone River at Pierre-B!enite,France. Freshwater Biology 51: 1730–1743.

Laroche, J., and Durand, J.D. 2004. Genetic structure of fragmented popula-tions of a threatened endemic percid of the Rhone river: Zingel asper.

Heredity 92(4): 329–334.Le Corre, M., Sabati!e, M.R., and Baglini"ere, J.L. 2000. Caract!erisation

d!emographique de populations d’Alosa fallax rhodanensis (Clupeidae)

de la M!editerran!ee francaise. Cybium 24(3 Suppl.): 119–128.Le Corre, M., Baglini"ere, J.L., Sabati!e, M.R., Menella, J.Y., and Pont, D.

1997. Donn!ees r!ecentes sur les caract!eristiques morphologiques et bio-

logiques de la population d’alose feinte du Rhone (Alosa fallax rhoda-

nensis Roule 1924). Bulletin Francais de Peche et de Pisciculture 346:

527–545.Le Corre, M., Alexandrino, P., Sabati!e, M.R., Aprahamian, M.W., and

Baglini"ere, J.L. 2005. Genetic characterisation of the Rhodanian twaite

shad, Alosa fallax rhodanensis. Fisheries Management and Ecology 12:275–282.

Le Corre, M., Linhares, D., Castro, F., Alexandrino, P., Sabati!e, M.R., andBaglini"ere, J.L. 1998. Premiers !el!ements de caract!erisation g!en!etique

de l’alose du Rhone (Alosa fallax rhodanensis Roule 1924). BulletinFrancais de Peche et de Pisciculture 350–351: 635–645.

Lefebvre, F., Acou, A., Poizat, G., and Crivelli, A. 2006. Epid!emiologie de

l’anguillicolose et qualit!e des individus argent!es : !etude sur 4 ans, dans4 sites de Camargue. Commission Technique du Comit!e de Gestion desPoissons Migrateurs.

L!eger, L. 1926. Carte piscicole du d!epartement de l’Ain avec une noticeexplicative. Travaux du Laboratoire d’Hydrobiologie et de Pisciculturede l’Universit!e de Grenoble 19: 149–155.

L!eger, L. 1934. Carte piscicole du d!epartement des Hautes Alpes (avec

notice explicative et consid!erations sur l’!economie piscicole r!egionale).Travaux du Laboratoire d’Hydrobiologie et de Pisciculture de

l’Universit!e de Grenoble 251–28, 28 pages + carte hors texte.

L!eger, L. 1943. Carte piscicole du d!epartement de la Savoie avec une noticesur l’Hydrographie et l’!economie piscicoles des cours d’eau et des lacs.Travaux du Laboratoire d’Hydrobiologie et de Pisciculture de

l’Universit!e de Grenoble 34–36: 51–66.L!eger, L. 1945. Etude sur l’hydrobiologie et l’!economie piscicoles du

d!epartement du Rhone avec une carte et un graphique. Travaux du

Laboratoire d’Hydrobiologie et de Pisciculture de l’Universit!e de Gre-

noble 37–40: 1–14.Loizeau, J.L., and Dominik, J. 2000. Evolution of the Upper Rhone

River discharge and suspended sediment load during the last 80

years and some implications for Lake L!eman. Aquatic Sciences

62: 54–67.Lunel, G. 1874. Histoire naturelle des poissons du basin du L!eman. Georg,

Gen"eve.Malard, F., Galassi, D., Lafont, M., Doledec, S., and Ward, J.V. 2003.

Longitudinal patterns of invertebrates in the hyporheic zone of a glacial

river. Freshwater Biology 48(10): 1709–1725.Mallet, J.P. 1999. Recherche des facteurs de controle de la dynamique des

populations d’ombre commun Thymallus thymallus (L. 1758) de la

Basse Rivi"ere d’Ain. Doctoral thesis University Lyon, p. 1.Marigo, G., Peltier, J.P., Girel, J., and Pautou, G. 2000. Success in the

demographic expansion of Fraxinus excelsior L. Trees 15: 1–13.Marmonier, P. 1988.Bioc!enoses interstitielles et circulation des eaux dans le

sous-!ecoulement d’un chenal am!enag!e du Haut Rhone francais. Doc-toral thesis, University of Lyon.

Marmonier, P., and Dole, M.J. 1986. Les Amphipodes des s!ediments d’un

bras court-circuit!e du Rhone: logique de r!epartition et r!eaction auxcrues. Sciences de l’eau 5: 461–486.

Marmonier, P., Dole-Olivier, M.J., and Creuz!e des Chatelliers, M. 1992.

Spatial distribution of interstitial assemblages in the floodplain of theRhone River. Regulated Rivers: Research & Management 7: 75–82.

Marston, R.A., Girel, J., Pautou, G., Piegay, H., Bravard, J.P., and Arneson,C. 1995. Channel metamorphosis, floodplain disturbance, and vegeta-

tion development – Ain River, France. Geomorphology 13: 121–131.Mathys, N., Brochot, S., Meunier, M., and Richard, D. 2003. Erosion quan-

tification in the small marly experimental catchments of Draix (Alpes

de Haute Provence France). Calibration of the ETC rainfall-runoff-erosion model. CATENA 50: 527–548.

Maurin, H. and Keith, P. (eds). 1994. Le livre rouge. Inventaire de la faune

menac!ee de France, Editions Nathan, Paris.


Mayaud, N. 1936. Inventaire des Oiseaux de France. Soci!et!e d’Etudes

Ornithologiques, Paris.Meier, W., Frey, M., Moosmann, L., Steinlin, S., and W#uest, A. 2004.

Schlussbericht Rhone Ist-Zustand. Rhone-Thur Projekt, Subprojekt I-

2: Wassertemperaturen und W#armehaushalt der Rhone und ihrer Sei-tenb#ache. EAWAG, Kastanienbaum.

Meile, T., Boillat, J.L., and Schleiss, A.J. 2006. Influence of dams andreservoirs on the flow regime of the Upper Rhone River. Proceedings

of the Twenty-second Congress on Large Dams, Barcelona, Spain,In-ternational Commission on Large Dams, Paris.

Meybeck, M. 1986. Composition chimique naturelle des ruisseaux non

pollu!es en France. Bulletin de la Soci!et!e G!eologique 39: 3–77.Miramont, C., Jorda, M., and Pichard, G. 1998. Fluvial morphogenesis

evolution of a mediterranean river: the example of the Durance fluvial

system (south-eastern France).Geographie physique et Quaternaire 52:381–392.

Moutin, T., Raimbault, P., Golterman, H.L., and Coste, B. 1998. Theinput of nutrients by the Rhone river into the Mediterranean Sea:

recent observations and comparison with earlier data. Hydrobiologia374: 237–246.

Ollivier, P., Radakovitch, O., and Hamelin, B. 2006. Unusual variations of

dissolved As, Sb and Ni in the Rhone River during flood events. Journalof Geochemical Exploration 88: 394–398.

Paillex, A. 2005.Etat de r!ef!erence hydrobiologique de huit annexes fluviales

avant restauration (Haut-Rhone francais). Diplome en Sciences natur-elles de l’environnement, Universit!e de Gen"eve.

Parde, M. 1925. Le R!egime du Rhone. Etude hydrologique. Doctoral thesis,

University of Grenoble.Pautou, G., and Girel, J. 1994. Interventions humaines et changements de la

v!eg!etation alluviale dans la vall!ee de l’Is"ere (de Montm!elian au Port deSt-Gervais). RGA 2: 127–146.

Persat, H. 1988. De la biologie des populations de l’ombre commun Thy-

mallus thymallus (L. 1758) "a la dynamique des communaut!es dans un

hydrosyst"eme fluvial am!enag!e, le Haut-Rhone francais. El!ements pour

un changement d’!echelles. Doctoral Thesis es Sciences, UniversityLyon, p. 1.

Persat, H., and Berrebi, P. 1990. Relative ages of present populations of

Barbus barbus and Barbus meridionalis (Cyprinidae) in southernFrance: preliminary considerations. Aquatic Living Resources 3:253–263.

Persat, H., and Eppe, R. 1997. Alevinage, pollution et cloisonnement del’espace fluvial dans les structures g!en!etiques des populations de pois-son: l’ombre commun, Thymallus thymallus, dans le Rhone au niveaude la Savoie. Bulletin Francais de la Peche et de la Pisciculture 344–

345: 287–299.Persat, H., and Keith, P. 1997. La r!epartition g!eographique des poissons

d’eau douce en France: qui est autochtone et qui ne l’est pas? Bulletin

Francais de la Peche et de la Pisciculture 344–345: 15–32.Persat H., and Fruget, J.F. (Coord). 2007. Etude et analyse du fonctionne-

ment biologique et trophique de la Saone dans le territoire du Grand

Lyon. Unpublished report to the ‘Grand Lyon’.Persat, H., Olivier, J.M., and Bravard, J.P. 1995. Stream riparian manage-

ment of large braided Mid-European rivers, and consequences for fish.In: Armantrout, N.B. (ed). Condition of the World’s Aquatic Habitats.

Proceedings of theWorld Fisheries Congress, Theme 1,Oxford and IBHPublishing Co. Pvt. Ltd., New Delhi, pp. 139–169.

Peter, A., andWeber, C. 2004. Die Rhone als Lebensraum f#ur Fische.Wasser

Energie Luft 11–12: 326–330.Pineau, O. and Sinnassamy, J.M. (eds). 2001. Plan de gestion de la Tour du

Valat 2001–2005 (Camargue, France), La Tour du Valat, Camargue,

France.

Poizat, G., and Pont, D. 1996. Multi-scale approach to species-habitat rela-

tionships: juvenile fish in a large river section. Freshwater Biology 36(3): 611–622.

Pont, D. 1997. Le Rhone: l’axe et la vall!ee. Les d!ebits solides du Rhone "a

proximit!e de son embouchure: donn!ees r!ecentes (1994–1995).Revue deG!eographie de Lyon 72: 23–33.

Pont, D., and Nicolas, Y., 2001. Habitat use by 0+ fish in an old-engineeredriver reach (lower Rhone, France): relative importance of habitat het-

erogeneity and hydrological variability. Large Rivers 12 (2–4), ArchivF#ur Hydrobiolgy Supplements 135 (2–4): 219–238.

Pont, D., Simonnet, J.P., and Walter, A.V. 2002. Medium-term changes in

suspended sediment delivery to the ocean: Consequences of catchmentheterogeneity and river management (Rhone River, France). Estuarine.Coastal and Shelf Science 54: 1–18.

Poussard, G., and Madrid, N. 1999. Qualit!e des eaux du Rhone. Evolution1969–1995. Agence de l’Eau Rhone-M!editerran!ee-Corse, Lyon.

Pr!evot, R. 1984. Hydrobiologie de la Moyenne-Durance. Etude des diff!er-

ents milieux et de leurs peuplements de macroinvert!ebr!es. Doctoral

thesis, University Aix-Marseille III.Provansal, M. 2003. Le delta du Rhone: un h!eritage menac!e. La Lettre du

Changement global 15: 60–63.

Rabotin, M. 2002. Base de donn!ees piscicoles du bassin versant de laDurance: mise en place d’un Syst"eme d’Information G!eographique.DES Gestion Int!egr!ee des Ressources Hydriques, Option Gestion des

Ressources biologiques des Eaux continentales, Universit!e de Li"ege,Fondation Universitaire Luxembourgeoise.

Reverchon, B., Bellevilee, A., Schaltenbrand, N., andMenu, S. 2006. Essais

p!eriodiques des vannes de crue des barrages et d!emarche d’am!eliorationdu transport solide en suspension engag!ee sur la Durance. Proceedingsof Twenty-second Congress on Large Dams, Barcelona, Spain,Interna-tional Commission on Large Dams, Paris, France, pp. 411–427.

Roux, A.L. 1984. The impact of emptying and cleaning reservoirs on thephysico-chemical and biological water quality of the Rhone down-stream of the dams. In: Lillehammer, A., Salveit, S.J. (eds). Regulated

Rivers, Universitetsforlaget AS, Oslo, pp. 61–70.Roux, A.L., Bravard, J.P., Amoros, C., and Pautou, G. 1989. Ecological

changes of the French Upper Rhone since 1750. In: Petts, G., Moller,

H. (eds). Historical Change of Large Alluvial Rivers, John Wiley andSons Ltd., Chichester, pp. 323–350.

Sabatier, F., and Suanez, S. 2003. Evolution of the Rhone delta coast since

the end of the 19th century. G!eomorphologie: Relief, Processus, Envir-onnement 4: 283–300.

Salen-Picard, C., Darnaude, A.M., Arlhac, D., and Harmelin-Vivien, M.L.2002. Fluctuations of macrobenthic populations: a link between cli-

mate-driven river run-off and sole fishery yields in the Gulf of Lions.Oecologia 133: 380–388.

Salvador, P.G. 1999. L’!edification holoc"ene de la plaine alluviale du Rhone

dans le bassin de Malville-St-Brenaz (Ain, Is"ere). G!eomorphologie:

Relief, Processus, Environnement 1: 3–22.Salvador, P.G., Verot, A., Bravard, J.P., Franc, O., and Mace, S. 2002. Les

crues du Rhone "a l’!epoque gallo-romaine dans la r!egion lyonnaise. In:Bravard, J.P., Magny, M. (eds). Les Fleuves ont une histoire: pal!eo-

environnement des rivi"eres et des lacs francais depuis 15000 ans, Edi-tions Errance, Collection Arch!eologie aujourd’hui, Paris, pp. 215–221.

Santiago, S., Thomas, R.L., Larbaigt, G., Corvi, C., Rossel, D., Tarradellas,J., Gregor, D.J., McCarthy, L., and Vernet, J.P. 1994. Nutrient, heavy-Metal and organic pollutant composition of suspended and bed sedi-

ments in the Rhone River. Aquatic Sciences 56: 220–242.SDAGERM&C. 1996. Sch!emaDirecteur d’Am!enagement et deGestion des

Eaux, des bassins Rhone-M!editerran!ee et Corse. Agence de l’Eau

Rhone-M!editerran!ee-Corse, Lyon.


Seyed-Reihani, A., Ginet, R., and Reygrobellet, J.L. 1982. Structure et

fonctionnement des ecosyst"emes du Haut-Rhone francais. XXX. Lepeuplement de trois stations interstitielles dans la plaine de MiribelJonage (vall!ee du Rhone en amont de Lyon), en relation avec leur

alimentation hydrog!eologique. Revue Francaise des Sciences de l’Eau1(2): 163–174.

S.F.P.N.P. Gen"eve. 2003. Inventaire piscicole des d’eau du Canton de Gen-ve. Constats et plan d’actions envisag!es pour une politique cantonale

cibl!ee. Etudes Nature, Service des Forets, de la Protection de la Natureet du Paysage, R!epublique du Canton de Gen"eve.

SGFH. 1916a. Compte-rendu et R!esultats des Etudes et Travaux – Annexe

du Tome 7 – Cartes et nivellements. Service des Grandes ForcesHydrauliques (r!egion des Alpes), Minist"ere de l’Agriculture, DirectionG!en!erale des Eaux et Forets, Eaux et am!eliorations agricoles, Paris.

SGFH. 1916b. Liste des principales usines hydrauliques en 1916. Servicedes Grandes Forces Hydrauliques (r!egion des Alpes), Minist"ere del’Agriculture, Direction G!en!erale des Eaux et Forets, Eaux et am!eliora-tions agricoles, Paris.

SOGREAH. 2000. Etude globale pour une strat!egie de r!eduction des risquesdus aux crues. Etude du transport solide. Rapport de synth"ese. Institu-tion Interd!epartementale des Bassins Rhone-Saone, Valence.

Stanford, J.A., and Ward, J.V. 1988. The hyporheic habitat of river ecosys-tem. Nature 6185: 64–66.

Stanford, J.A., and Ward, J.V. 1993. An ecosystem perspective of alluvial

rivers: connectivity and the hyporheic corridor. Journal of the North

American Benthological Society 12(1): 48–60.Surell, A. 1847. M!emoire sur l’am!elioration des embouchures du Rhone.

Imprimerie C!evenole, Nimes.Tachet, H., Gaschignard-Fossati, O., Cellot, B., and Berly, A. 1988. Le

macrobenthos de la Saone. Annales de Limnologie 24(1): 83–100.Van Der Leeuw, S.E. 2005. Climate, hydrology, land use, and environmental

degradation in the lower Rhone Valley during the Roman period.Comp-tes Rendus Geosciences 337: 9–27.

Verneaux, J., Schmitt, A., Verneaux, V., and Prouteau, C. 2003. Benthic

insects and fish of the Doubs River system: typological traits and thedevelopment of a species continuum in a theoretically extrapolatedwatercourse. Hydrobiologia 190: 63–74.

Vivian, H. 1989. Hydrological changes of the Rhone River. In: Petts, G.E., Roux, A.L. (eds). Historical Change of Large Alluvial Rivers:

Western Europe, John Wiley and Sons, Chichester, pp. 55–77.

Ward, J.V. 1989. The four dimentional nature of lotic ecosystems. Journal ofthe North American Benthological Society 8(1): 2–8.

Warner, R.F. 2000. Gross channel changes along the Durance River, South-ern France, over the last 100 years using cartographic data. Regulated

Rivers: Research & Management 16: 141–157.Weber, C., Peter, A., and Zanini, F. 2007. Spatio-temporal analysis of fish

and their habitat: a case study on a highly degraded Swiss river system

prior to extensive rehabilitation. Aquatic Sciences - Research Across

Boundaries 69: 162–172.Wilhelm, I. 1913. La Durance. Etude de l’utilisation de ses eaux et de

l’am!elioration de son r!egime par la cr!eation de barrages. Lucien

LAVEUR, Paris.

RELEVANT WEBSITES

http://www.ecologie.gouv.fr/HYDRO-banque-nationale-de-donnees.html:National Data Bank ob Hydrometry and Hydrology (France).http://rdb.eaurmc.fr/: Information about water quality, aquatic milieu andwater management on the French Rhone watershed.

http://195.167.226.100/DCE/RM/RM_etat-des-lieux.htm: data about WFDimplementation in the French Rhone waterhed.http://www.rhonealpes.ecologie.gouv.fr/orgfh/html_orgfh_fichesesp.php3:

Direction R!egionale de l’Environnement Rhone-Alpes. D!el!egation de bas-sin Rhone-M!editerran!ee. Les orientations r!egionales de la gestion de lafaune sauvage et d’am!elioration de la qualit!e de ses habitats (O.R.G.F.H.)

en Rhone-Alpes.http://www.oieau.org/5pcrd/projets/EUROLAKES.htm: EUROLAKES, In-tegrated Water Resource Management for Important Deep European Lakes

and their Catchment Areas.http://natura2000.environnement.gouv.fr/regions/idxreg.html: Natura 2000network in France.http://www.cipel.org/sp/: website of the International Commission for the

Protection of Lake Geneva (CIPEL).http://195.167.226.100/peche/carnet.html, Statistics on professional andamateur fisheries on the Rhone and Saone are available since 1988.

http://www.vs.ch/Press/DS_12/PU-2005-06-29-7882/fr/Rapport%20FR.pdf: Documentation on the third correction of the Rhone in Valais, Switzer-land.

www.irsn.org: ‘Institut de Radioprotection et de Suret!e Nucl!eaire’, theIRSN’s scientific website gives information about the Institute’s researchactivities in the field of nuclear safety and radiation protection.http://inpn.mnhn.fr/: Mus!eum national d’Histoire naturelle [Ed]. 2003–

2006. /Inventaire national du Patrimoine naturel.http://www.thonon.inra.fr/poisson/pacagelacustre/pacagesalmonides/omblechevalier/omblepacage.htm: data about fish farming in the Lake

L!eman.http://lepus.unine.ch/carto/: Swiss Center of Animal Cartography !CSCFhttp://www.karch.ch/karch/d/pro/rolia/roliafs2.html: Red-list of Amphi-

bians in Switzerland.http://www.karch.ch/karch/f/pro/rolir/rolirfs2.html: Red-list of Reptiles inSwitzerland.

http://www.cren-Rhonealpes.fr/part2/progs/life_apron.htm: Life programon the Apron.



http://www.ecologie.gouv.fr/HYDRO-banque-nationale-de-donnees.html

http://rdb.eaurmc.fr/


http://www.rhonealpes.ecologie.gouv.fr/orgfh/html_orgfh_fichesesp.php3

http://www.oieau.org/5pcrd/projets/EUROLAKES.htm


http://www.cipel.org/sp/


http://www.vs.ch/Press/DS_12/PU-2005-06-29-7882/fr/Rapport%20FR.pdf

http://www.vs.ch/Press/DS_12/PU-2005-06-29-7882/fr/Rapport%20FR.pdf

http://www.irsn.org/

http://inpn.mnhn.fr/



http://lepus.unine.ch/carto/

http://www.karch.ch/karch/d/pro/rolia/roliafs2.html

http://www.karch.ch/karch/f/pro/rolir/rolirfs2.html


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